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Trophic ecology of epibenthic communities exposed to different sea-ice concentrations across the Canadian Arctic Ocean
Abstract
Sea ice is one of the most critical environmental drivers shaping primary production and fluxes of organic inputs to benthic communities in the Arctic Ocean. Fluctuations in organic inputs influence ecological relationships, trophic cascades, and energy fluxes. However, changes in sea-ice concentration (SIC) induced by global warming could lead to significant shifts in trophic interactions, ultimately affecting the functioning of Arctic food webs. Despite the increasing concern over the need to understand benthic species and food web responses to rapid sea-ice loss, few studies have addressed this topic so far. Using multiple niche metrics based on stable isotopes, this research examined the trophic ecology of epibenthic communities in areas with different SIC across the Canadian Arctic Ocean. We found that trophic niches varied according to complex interactions between environmental conditions, resource supply, and biotic pressures such as predation and competition. Our results highlighted a lower isotopic richness (i.e., shorter food chain length and niche width) in low and high SIC areas, suggesting homogeneity of resources and a low diversity of food items ingested by individuals. In contrast, a higher isotopic richness (i.e., broad niche) was observed in the moderate SIC area, implying higher heterogeneity in basal food sources and consumers using individual trophic niches. Finally, our findings suggested a lower isotopic redundancy in areas with high SIC compared to low and moderate SIC. Overall, our results support the idea that sea ice is an important driver of benthic food web dynamics and reinforce the urgent need for further investigations of declining sea ice cover impacts on Arctic food web functioning.
... Over 1400 refereed scientific publications and 400 datasets have resulted from the CCGS Amundsen expeditions. The publications range from physical oceanography to geology, biogeochemistry, ecology and safety hazard assessments (Dmitrenko et al., 2023;Rodríguez-Cuicas et al., 2023;Stern et al., 2023;Vogt et al., 2023;Yunda-Guarin et al., 2023). Hosted at Université Laval, Amundsen Science is the organization responsible for the management of the scientific mandate of the research icebreaker CCGS Amundsen. ...
Since 2003, the state-of-the-art Canadian Coast Guard Ship (CCGS) research icebreaker Amundsen has furrowed the Canadian Arctic waters to support novel research endeavors and collect oceanographic data. This paper presents the data acquisition, the processing methods and an overview of the data collected during the 2021 expedition as the ship traveled over 30 000 km during 122 d across the Canadian Arctic Ocean, collecting sea surface, atmospheric and seabed underway measurements. A total of 266 casts of a conductivity, temperature and depth profiler mounted on a Conductivity Temperature Depth rosette (CTD Rosette) were also conducted to monitor the main physical, chemical and biological parameters of the water column. More specifically, the data presented here were collected with the CTD Rosette across historical sampling transects in Davis Strait, the North Water Polynya (NOW) and Cape Bathurst. A 182 km dedicated survey using the Moving Vessel Profiler® (MVP), equipped with CTD, transmissometer, dissolved oxygen, fluorescence and sound velocity sensors, was conducted across Hudson Strait. We also present an overview of the data collected by the underway systems (seabed, thermosalinograph and atmospheric). Such data are essential in understanding the impacts of climate warming on the unique environments of the Canadian Arctic Ocean. Amundsen Science supports and promotes easy access and sharing of such valuable data to the scientific community.
The polymer-facilitated flux of ice algae on Arctic shelves can initiate benthic activity and growth after the nutritionally constrained winter period. Lipid-rich ice algae are readily consumed by benthos and those entering the sediment can benefit deposit feeders. Ice algae assimilated by benthic organisms cascade up multiple trophic levels within the benthic sub-web, re-entering the pelagic sub web through habitat coupling species. Pelagic predators can have significant ice-algal carbon signals obtained from the benthic compartment. Sympagic-pelagic-benthic coupling on Arctic shelves is expected to weaken with ongoing sea-ice change. This review discusses the phenology, quantity, and quality of ice-algal contributions to coupling, linked to thinning snow and ice cover including multi-year ice replacement. Predicting future coupling between marine sub-webs requires focused research that considers trophic markers of multiple carbon sources.
On Arctic shelves, where primary production occurs in both the pelagic and sympagic (ice-associated) habitats, sympagic organic material (OM) can constitute a disproportionate fraction of benthic diets due to higher sinking rates and lower grazing pressure than pelagic OM. Less documented is how sympagic OM assimilation across feeding guilds varies seasonally and in relation to sea ice duation. We therefore investigated the relative abundance of sympagic vs pelagic OM in Barents Sea shelf megabenthos in the summer and winter of 2018 and 2019, from 10 stations where sea ice duration ranged from 0 to 245 days per year. We use highly branched isoprenoids, which are lipid biomarkers produced with distinct molecular structures by diatoms in sea ice and the water column, to determine the ratio of sympagic-to-pelagic OM assimilated by benthic organisms. From 114 samples of 25 taxa analysed, we found that the proportion of sympagic OM assimilated ranged from 0.4% to 95.8% and correlated strongly (r2 = 0.754) with the duration of sea ice cover. The effect of sea ice duration was more evident in fauna collected during summer than winter, indicating that sympagic signals are more evident in the summer than in the winter at higher latitudes. Our data show that sympagic production can supply a high fraction of carbon for Barents Sea benthos, although this is highly variable and likely dependent on availability and patchiness of sympagic OM deposition. These results are comparable to similar studies conducted on benthos in the Pacific Arctic and highlight the variable importance of sympagic OM in the seasonal ice zone of Arctic inflow shelves, which are the Arctic regions with highest rates of sea ice loss.
Establishing management programs to preserve the benthic communities along the NW Pacific and the Arctic Ocean (AO) requires a deep understanding of the composition of communities and their responses to environmental stressors. In this study, we thus examine patterns of benthic community composition and patterns of species richness along the NW Pacific and Arctic Seas and investigate the most important environmental drivers of those patterns. Overall we found a trend of decreasing species richness toward higher latitudes and deeper waters, peaking in coastal waters of the eastern Philippines. The most dominant taxa along the entire study area were Arthropoda, Mollusca, Cnidaria, Echinodermata, and Annelida. We found that depth, not temperature, was the main driver of community composition along the NW Pacific and neighboring Arctic Seas. Depth has been previously suggested as a factor driving species distribution in benthic fauna. Following depth, the most influential environmental drivers of community composition along the NW Pacific and the Arctic Ocean were silicate, light, and currents. For example, silicate in Hexactinellida, Holothuroidea, and Ophiuroidea; and light in Cephalopoda and Gymnolaemata had the highest correlations with community composition. In this study, based on a combination of new samples and open-access data, we show that different benthic communities might respond differently to future climatic changes based on their taxon-specific biological, physiological, and ecological characteristics. International conservation efforts and habitat preservation should take an adaptive approach and apply measures that take the differences among benthic communities in responding to future climate change into account. This facilitates implementing appropriate conservation management strategies and sustainable utilization of the NW Pacific and Arctic marine ecosystems.
Benthic organisms depend primarily on seasonal pulses of organic matter from primary producers. In the Arctic, declines in sea ice due to warming climate could lead to changes in this food supply with as yet unknown effects on benthic trophic dynamics. Benthic consumer diets and food web structure were studied in a seasonally ice-covered region of Baffin Bay during spring 2016 at stations ranging in depth from 199 to 2,111 m. We used a novel combination of highly branched isoprenoid (HBI) lipid biomarkers and stable isotope ratios (d 13 C, d 15 N) to better understand the relationship between the availability of carbon sources in spring on the seafloor and their assimilation and transfer within the benthic food web. Organic carbon from sea ice (sympagic carbon [SC]) was an important food source for benthic consumers. The lipid biomarker analyses revealed a high relative contribution of SC in sediments (mean SC% + standard deviation [SD] ¼ 86% + 16.0, n ¼ 17) and in benthic consumer tissues (mean SC% + SD ¼ 78% + 19.7, n ¼ 159). We also detected an effect of sea-ice concentration on the relative contribution of SC in sediment and in benthic consumers. Cluster analysis separated the study region into three different zones according to the relative proportions of SC assimilated by benthic macrofauna. We observed variation of the benthic food web between zones, with increases in the width of the ecological niche in zones with less sea-ice concentration, indicating greater diversity of carbon sources assimilated by consumers. In zones with greater sea-ice concentration, the higher availability of SC increased the ecological role that primary consumers play in driving a stronger transfer of nutrients to higher trophic levels. Based on our results, SC is an important energy source for Arctic deep-sea benthos in Baffin Bay, such that changes in spring sea-ice phenology could alter benthic food-web structure.
We studied ice algae utilization by benthic fauna from the northern Bering and Chukchi Seas using highly branched isoprenoid (HBI) biomarkers. We assessed whether various food acquisition strategies influence the observed HBI signatures. The proportion of phytoplankton to ice algae-sourced HBIs was determined through the H-Print approach that is presumed to reflect the percentage of sea ice organic carbon (iPOC) incorporated into tissues, relative to phytoplankton organic carbon. Cluster analysis separated 3 groups based on location and feeding strategy that were significantly influenced by annual sea ice persistence. Ice algae utilization was most significant in the northeast Chukchi Sea, where seasonal sea ice was present the longest. General feeding strategy was determined to be a significant factor in the degree of ice algae utilization. Predominant deposit feeders (both surface and subsurface) used more ice algae relative to suspension feeders. Organic carbon incorporated by predominant suspension feeders was primarily phytoplankton-based. The vast majority of all organisms sampled (~90%) incorporated a measurable quantity of iPOC. Sipunculids and brittle stars had the highest relative dependence on ice algae, while other taxa displayed plastic dietary responses, including the suspension/surface deposit feeder Macoma calcarea. This study indicates that ice algae are widely utilized in Pacific Arctic benthic food webs, but most benthic organisms displayed flexibility in consuming the available food sources. The elevated utilization of ice algae by deposit feeders may prove to be a disadvantage for these organisms if they cannot adapt to the ongoing decline of iPOC as seasonal sea ice declines.
The decline of sea-ice thickness, area, and volume due to the transition from multi-year to first-year sea ice has improved the under-ice light environment for pelagic Arctic ecosystems. One unexpected and direct consequence of this transition, the proliferation of under-ice phytoplankton blooms (UIBs), challenges the paradigm that waters beneath the ice pack harbor little planktonic life. Little is known about the diversity and spatial distribution of UIBs in the Arctic Ocean, or the environmental drivers behind their timing, magnitude, and taxonomic composition. Here, we compiled a unique and comprehensive dataset from seven major research projects in the Arctic Ocean (11 expeditions, covering the spring sea-ice-covered period to summer ice-free conditions) to identify the environmental drivers responsible for initiating and shaping the magnitude and assemblage structure of UIBs. The temporal dynamics behind UIB formation are related to the ways that snow and sea-ice conditions impact the under-ice light field. In particular, the onset of snowmelt significantly increased under-ice light availability (>0.1–0.2 mol photons m–2 d–1), marking the concomitant termination of the sea-ice algal bloom and initiation of UIBs. At the pan-Arctic scale, bloom magnitude (expressed as maximum chlorophyll a concentration) was predicted best by winter water Si(OH)4 and PO43– concentrations, as well as Si(OH)4:NO3– and PO43–:NO3– drawdown ratios, but not NO3– concentration. Two main phytoplankton assemblages dominated UIBs (diatoms or Phaeocystis), driven primarily by the winter nitrate:silicate (NO3–:Si(OH)4) ratio and the under-ice light climate. Phaeocystis co-dominated in low Si(OH)4 (i.e., NO3:Si(OH)4 molar ratios >1) waters, while diatoms contributed the bulk of UIB biomass when Si(OH)4 was high (i.e., NO3:Si(OH)4 molar ratios
Isotopic niches reflect the basic structure and functioning of river food webs; however, their response to riverine environments remains unclear. We used stable isotope analysis and community-wide metrics to quantify how invertebrate niches vary with environmental changes along a large subtropical river in China. Eight niche metrics, which had higher values in the wet than in the dry season, increased from headwaters to the middle river and decreased sharply near the estuarine industrial zones. The δ13C value of > − 23.8‰, which indicated consumption of epilithic diatoms, separated the invertebrates between the upper and mid-lower reaches. The δ15N values > 9.4‰ identified site-specific nitrogen sources from manure (e.g., animal effluent) and domestic sewage in agricultural area. The output of mixing models showed a downstream shift in carbon utilization by invertebrates from autochthonous periphyton and submerged hydrophytes to allochthonous C3 plants. Principle component (PC) and cluster analysis decomposed and grouped 40 environmental variables into 4 PCs that explained 84.5% of the total variance. Hierarchical partitioning revealed that the second and first PCs, which were driven mainly by biological indicators and habitat characteristics, had the highest explanatory power for niche ranges and areas (e.g., Bayesian ellipse), respectively. Our results suggest that reducing anthropogenic pressures (e.g., habitat loss and water pollution) on river ecosystems through measures, such as protecting diatom-dominated biofilms in riffles and controlling nitrogen loading in rural regions, may produce the greatest impact for river management.
Graphical abstract
The production and transfer of biomass through trophic relationships is a core ecosystem function. The movement of energy through the food web is mediated by organisms operating in their niche space. For generalists, the size of this niche space is inherently plastic and changes in response to available food sources. Therefore, this relationship between ecosystem productivity and niche size is an important determinant of ecosystem function. Competing theories about the nature of this relationship predict that as productivity increases niche size will either increase as species capitalize on a general increase in resource availability or decrease as it becomes viable to focus on preferred production channels. Here, we test these two competing theoretical frameworks using a novel approach to determine trophic niche size using stable isotope analysis and hypervolume metrics. Resource use is quantified in two generalist fish species at three productivity levels in a seagrass ecosystem. Niche size of both species was inversely related to seagrass productivity, consistent with the hypothesis that increasing productivity allows species to focus on a narrower diet. This pattern describes the relationship between ecosystem production and niche size and provides an empirical ecological explanation for the resource maximization behaviors commonly observed in nature.
The production and transfer of biomass through trophic relationships is a core ecosystem function. The movement of energy through the food web is mediated by organisms operating in their niche space. For generalists, the size of this niche space is inherently plastic and changes in response to available food sources. Therefore, this relationship between ecosystem productivity and niche size is an important determinant of ecosystem function. Competing theories about the nature of this relationship predict that as productivity increases niche size will either increase as species capitalize on a general increase in resource availability or decrease as it becomes viable to focus on preferred production channels. Here, we test these two competing theoretical frameworks using a novel approach to determine trophic niche size using stable isotope analysis and hypervolume metrics. Resource use is quantified in two generalist fish species at three productivity levels in a seagrass ecosystem. Niche size of both species was inversely related to seagrass productivity, consistent with the hypothesis that increasing productivity allows species to focus on a narrower diet. This pattern describes the relationship between ecosystem production and niche size and provides an empirical ecological explanation for the resource maximization behaviors commonly observed in nature.
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The development of the isotopic niche, an n-dimensional hypervolume (with n being the number of isotopes) occupied by a population in delta-space, has revolutionized the study of animal interactions in wild populations. While the isotopic niche offers a useful means to understand interactions at many ecological resolutions (e.g., individual, population, community, ecosystem), a variety of intrinsic and extrinsic factors drive isotopic variability and influence the ultimate geometry of observed niche dimensions. Here, we provide an updated synthesis to guide the application of bulk stable isotope ratios to study ecological niches. We summarize progress in the application of bulk stable isotope ratios for evaluating niches to synthesize a formal definition of the isotopic niche. We identify six broad categories to describe drivers of isotopic variance introduced by the animal, its environment, and the researcher, and provide recommendations to account for such variations before, during, and after sample collection and data analyses. Our synthesis illustrates the considerations that should be made before employing the isotopic niche to describe trophic ecology, and guidance for the use and interpretation of isotopic niche dynamics in future studies.
Linking mechanistic processes to the stability of ecological networks is a key frontier in ecology. In trophic networks, “modules”—groups of species that interact more with each other than with other members of the community—confer stability, mitigating effects of species loss or perturbation. Modularity, in turn, is shaped by the interplay between species’ diet breadth traits and environmental influences, which together dictate interaction structure. Despite the importance of network modularity, variation in this emergent property is poorly understood in complex natural systems. Using two years of field data, we quantified interactions between a rich community of lepidopteran herbivores and their host plants across a mosaic of low-resource serpentine and high-resource nonserpentine soils. We used literature and our own observations to categorize herbivore species as generalists (feeding on more than one plant family) or specialists (feeding on one plant family). In both years, the plant-herbivore network was more modular on serpentine than on nonserpentine soils—despite large differences in herbivore assemblage size across years. This structural outcome was primarily driven by reduction in the breadth of host plant use by generalist species, rather than by changes in the composition of species with different fundamental diet breadths. Greater modularity—and thus greater stability—reflects environmental conditions and plastic responses by generalist herbivores to low host plant quality. By considering the dual roles of species traits and ecological processes, we provide a deeper mechanistic understanding of network modularity, and suggest a role for resource availability in shaping network persistence.
The Arctic Ocean is particularly affected by climate change, with changes in sea ice cover expected to impact phytoplankton primary production. During the Green Edge expedition, the development of the late spring–early summer diatom bloom was studied in relation with the sea ice retreat by multiple transects across the marginal ice zone. Biogenic silica concentrations and uptake rates were measured. In addition, diatom assemblage structures and their associated carbon biomass were determined, along with taxon- specific contributions to total biogenic silica production using the fluorescent dye PDMPO. Results indi- cate that a diatom bloom developed in open waters close to the ice edge, following the alleviation of light limitation, and extended 20–30 km underneath the ice pack. This actively growing diatom bloom (up to 0.19 μmol Si L–1 d–1) was associated with high biogenic silica concentrations (up to 2.15 μmol L–1), and was dominated by colonial fast-growing centric (Chaetoceros spp. and Thalassiosira spp.) and ribbon-forming pennate species (Fragilariopsis spp./Fossula arctica). The bloom remained concentrated over the shallow Greenland shelf and slope, in Atlantic-influenced waters, and weakened as it moved westwards toward ice- free Pacific-influenced waters. The development resulted in a near depletion of all nutrients eastwards of the bay, which probably induced the formation of resting spores of Melosira arctica. In contrast, under the ice pack, nutrients had not yet been consumed. Biogenic silica and uptake rates were still low (respectively <0.5 μmol L–1 and <0.05 μmol L–1 d–1), although elevated specific Si uptake rates (up to 0.23 d–1) probably reflected early stages of the bloom. These diatoms were dominated by pennate species (Pseudo-nitzschia spp., Ceratoneis closterium, and Fragilariopsis spp./Fossula arctica). This study can contribute to predic- tions of the future response of Arctic diatoms in the context of climate change.
In recent years, sea ice cover along coasts of East Antarctica has tended to increase. To understand ecological implications of these environmental changes, we studied benthic food web structure on the coasts of Adélie Land during an event of unusually high sea ice cover (i.e. two successive austral summers without seasonal breakup). We used integrative trophic markers (stable isotope ratios of carbon, nitrogen and sulfur) to build ecological models and explored feeding habits of macroinvertebrates. In total, 28 taxa spanning most present animal groups and functional guilds were investigated. Our results indicate that the absence of seasonal sea ice breakup deeply influenced benthic food webs. Sympagic algae dominated the diet of many key consumers, and the trophic levels of invertebrates were low, suggesting omnivore consumers did not rely much on predation and/or scavenging. Our results provide insights about how Antarctic benthic consumers, which typically live in an extremely stable environment, might adapt their feeding habits in response to sudden changes in environmental conditions and trophic resource availability. They also show that local and/or global trends of sea ice increase in Antarctica have the potential to cause drastic changes in food web structure, and therefore to impact benthic communities.
Competition plays a central role in the maintenance of biodiversity. A backbone of classic niche theory is that local coexistence of competitors is favoured by the contraction or divergence of species' niches. However, this effect should depend on the diversity of resources available in the local environment, particularly when resources vary in multiple ecological dimensions. Here, we investigated how available resource breadth (i.e. prey diversity) and competition together shape multidimensional niche variation (between and within individuals) and interspecific niche overlap in 42 populations of congeneric tropical frog species. We modelled realized niches in two key trophic dimensions (prey size and carbon stable isotopes) and sampled available food resources to quantify two-dimensional resource breadth. We found a 14-fold variation in multidimensional population niche width across populations, most of which was accounted for by within-individual diet variation. This striking variation was predicted by an interaction whereby individual niche breadth increased with resource breadth and decreased with the number of congeneric competitors. These ecological gradients also interact to influence the degree of niche overlap between species, which surprisingly decreased with population total niche width, providing novel insights on how similar species can coexist in local communities. Together, our results emphasize that patterns of exploitation of resources in multiple dimensions are driven by both competitive interactions and extrinsic factors such as local resource breadth.
Benthic macrofauna is an important component linking pelagic and benthic ecosystems, especially in productive coastal areas. Through their metabolism and behaviour, benthic animals affect biogeochemical fluxes between the sediment and water column. Mechanistic models that quantify these benthic-pelagic links are imperative to understand the functioning of coastal ecosystems. In this study, we develop a dynamic model of benthic macrofauna to quantify the relationship between organic matter input and benthic macrofaunal biomass in the coastal zone. The model simulates the carbon dynamics of three functional groups of benthic macrofauna and their sediment food sources and is forced by a hydrodynamic-biogeochemical model simulating pelagic physical and biological dynamics. The model reproduces measured time-series of macrofaunal biomass from two coastal sites with contrasting sedimentation in the Baltic Sea in 1993-2005 with comparatively high accuracy, including a major increase at one of the sites dominated by the bivalve Limecola (Macoma) balthica. This shift in community composition suggests altered pathways of organic matter degradation: 39% of simulated sedimentation was mineralized by macrofauna in 2005 compared to 10% in 1995. From the early 2000s onward macrofaunal biomass seems to be food-limited, as ca 80% of organic carbon sedimentation was processed by the deposit-feeding macrofauna at both sites. This model is a first step to help quantify the role of macrofauna in marine coastal ecosystem functioning and biogeochemical cycles and build predictive capacity of the effects of anthropogenic stressors, such as eutrophication and climate change, on coastal ecosystems.
Understanding drivers of benthic-pelagic coupling in Arctic marine ecosystems is key to identifying benthic areas that may be sensitive to climate-driven changes in hydrography and surface production. We coupled algal biomass and sedimentary characteristics with stable isotope data for 113 fishes and invertebrates in the Canadian Beaufort Sea and Amundsen Gulf to examine how trophic structure was influenced by the vertical water mass structure and by organic matter input regimes, from 20 to 1000 m depths. Indices of community-level trophic diversity (isotopic niche size, ¹³C enrichment relative to a pelagic baseline, and δ¹³C isotopic range) increased from west to east, coincident with the use of more diverse dietary carbon sources among benthic functional groups. Data suggested benthic-pelagic trophic coupling was strongest in the western study region where pelagic sinking flux is relatively high, intermediate in the central region dominated by riverine inputs of terrestrial organic matter, and weakest in the east where strong pelagic grazing is known to limit sinking flux. Differences in δ¹³C between pelagic and benthic functional groups (up to 5.7‰) increased from west to east, and from the near-shore shelf to the upper slope. On the upper slope, much of the sinking organic matter may be intercepted in the water column, and dynamic hydrography likely diversifies available food sources. In waters >750 m there were no clear trends in benthic-pelagic coupling or community-level trophic diversity. This study represents the first description of fish and invertebrate food web structure >200 m in the Canadian Beaufort Sea.
Climate change is expected to affect resource-consumer interactions underlying stability in polar food webs. Polar benthic organisms have adapted to the marked seasonality characterising their habitats by concentrating foraging and reproductive activity in summer months, when inputs from sympagic and pelagic producers increase. While this enables the persistence of biodiverse food webs, the mechanisms underlying changes in resource use and nutrient transfer are poorly understood. Thus, our understanding of how temporal and spatial variations in the supply of resources may affect food web structure and functioning is limited. By means of C and N isotopic analyses of two key Antarctic benthic consumers (Adamussium colbecki, Bivalvia, and Sterechinus neumayeri, Echinoidea) and Bayesian mixing models, we describe changes in trophic niche and nutrient transfer across trophic levels associated with the long- and short-term diet and body size of specimens sampled in midsummer in both shallow and deep waters. Samplings occurred soon after the sea-ice broke up at Tethys Bay, an area characterised by extreme seasonality in sea-ice coverage and productivity in the Ross Sea. In the long term, the trophic niche was broader and variation between specimens was greater, with intermediate-size specimens generally consuming a higher number of resources than small and large specimens. The coupling of energy channels in the food web was consequently more direct than in the short term. Sediment and benthic algae were more frequently consumed in the long term, before the sea-ice broke up, while consumers specialised on sympagic algae and plankton in the short term. Regardless of the time scale, sympagic algae were more frequently consumed in shallow waters, while plankton was more frequently consumed in deep waters. Our results suggest a strong temporal relationship between resource availability and the trophic niche of benthic consumers in Antarctica. Potential climate-driven changes in the timing and quality of nutrient inputs may have profound implications for the structure of polar food webs and the persistence of their constituent species, which have adapted their trophic niches to a highly predictable schedule of resource inputs.
Multiple hypotheses propose an ostensibly disparate array of drivers of food chain length (FCL), with contradictory support from natural settings. Here we posit that the magnitude of vertical energy flux in food webs underlies several drivers of FCL. We show that rising energy flux fuels top-heavy biomass pyramids, promoting omnivory, thereby reducing FCL. We link this theory to commonly evaluated hypotheses for environmental drivers of FCL (productivity, ecosystem size) and demonstrate that effects of these drivers should be context dependent. We evaluate support for this theory in lake and marine ecosystems and demonstrate that ecosystem size is the most important driver of FCL in low-productivity ecosystems (positive relationship) while productivity is most important in large and high productivity ecosystems (negative relationship). This work stands in contrast to classical
hypotheses, which predict a positive effect of productivity on FCL, and may help reconcile the contradictory nature of published results for drivers of FCL.
Climate change driven alterations to patterns of Arctic marine primary production, with increasing phytoplankton- and decreasing ice algal production, have the potential to change the resource utilisation and trophic structure of the benthic communities relying on the algae for food. To predict the benthic responses to dietary changes, we studied the macroinfaunal community compositions, and used the faunal δ¹³C and δ¹⁵N signatures to investigate their main food sources and trophic positions in North Water (NOW) and Lancaster Sound (LS) polynyas in the Canadian Arctic Archipelago. Macroinfaunal density (10 952 ind. m⁻²) and biomass (3190 mg C m⁻²) recorded in NOW were higher than previously found in the Arctic at depths >500m, and significantly higher than in LS (8355 ind. m⁻² and 2110 mg C m⁻²). This was attributed to higher particulate organic matter fluxes to seafloor in NOW. Polychaetes were significant taxa at both sites in terms of density and biomass, and in addition crustacean density in NOW and bivalve density in LS were high. Facultative filter and surface deposit feeders were highly prevalent at both sites, suggesting feeding plasticity is a successful strategy for accessing different food sources. The macrofaunal δ¹³C signatures reflected the signatures of pelagic particulate organic matter at the sites, and an isotope mixing model confirmed phytoplankton as the main food source for most taxa and feeding guilds. The food web length in LS was longer than in NOW (3.2 vs. 2.8 trophic levels). This was attributed to a larger reliance on reworked organic matter by the benthic community in LS, whereas the high export fluxes at the highly productive NOW resulted in higher rates of selective consumption of fresh algal matter. Despite studies suggesting that loss of ice algae from consumer diets in the Arctic might have a negative impact on the benthos, this study suggests that Arctic macrobenthic communities thrive using phytoplankton as their main food source and should thus be able to cope or even benefit from predicted changes to patterns of primary production.
In seasonal tropical regions, rainfall and/or temporary floods during the wet season generally increase the abundance and diversity of food resources to many consumers as compared to the dry season. Therefore, seasonality can affect intraspecific competition and ecological opportunity, which are two important ecological mechanisms underlying population and individual niche variations. Here, we took advantage of the strong seasonality in the Pantanal wetlands to investigate how within- and between-individual diet variations relate to seasonal population niche dynamics of the tetra fish Astyanax lacustris. We quantified dietary niche using gut contents and stable isotopes. Tetras had higher gut fullness and better body condition in the wet season, suggesting that competition is more intense in the dry season. The population niche was broader in the wet season due to an increase in diet divergence between individuals, in spite of potential stronger competition in the dry season. We posit that low ecological opportunity in the dry season limits the diversifying effect of intraspecific competition, constraining population niche expansion. Our results add new insights on how seasonality affects population and individual diets, indicating that intraspecific competition and ecological opportunity interact to determine temporal niche variations in seasonal environments.
Benthic-pelagic coupling is manifested as the exchange of energy, mass, or nutrients between benthic and pelagic habitats. It plays a prominent role in aquatic ecosystems and it is crucial to functions from nutrient cycling to energy transfer in food webs. Coastal and estuarine ecosystem structure and function is strongly affected by anthropogenic pressures, however there are large gaps in our understanding of the responses of inorganic nutrient and organic matter fluxes between benthic habitats and the water column. We illustrate the varied nature of physical and biological benthic-pelagic coupling processes and their potential sensitivity to three anthropogenic pressures — climate change, nutrient loading, and fishing — using the Baltic Sea as a case study, and summarize current knowledge on the exchange of inorganic nutrients and organic material between habitats. Traditionally measured benthic-pelagic coupling processes (e.g. nutrient exchange and sedimentation of organic material) are to some extent quantifiable but the magnitude and variability of biological processes are rarely assessed, preventing quantitative comparisons. Changing oxygen conditions will continue to have widespread effects on the processes that govern inorganic and organic matter exchange among habitats while climate change and nutrient load reductions may have large effects on organic matter sedimentation. Many biological processes (predation, bioturbation) are expected to be sensitive to anthropogenic drivers but the outcomes for ecosystem function are largely unknown. We emphasize how improved empirical and experimental understanding of benthic-pelagic coupling processes and their variability are necessary to inform models that can quantify the feedbacks among processes and ecosystem responses to a changing world. This article is protected by copyright. All rights reserved.
Despite significant progress in recent years, understanding the rules governing the assembly of natural communities is still challenging and knowledge of how the integration of nonnative species may disrupt community structure and function is needed. To address this challenge, we collated stable isotope data for 159 freshwater fish communities around the world with and without nonnative species and quantified spatial variation in both community isotopic functional diversity and intraspecific variation in species niches. Using a null model and partial least squares path analysis, we then evaluated how the interplay between abiotic (historical, energetic, climatic, habitat size) and biotic (niche segregation) factors shape community structure and functional diversity, and how these relationships have changed, and with what consequences, in the presence of nonnative species. We found that niche partitioning is a primary force underlying the structure and functional redundancy of native fish communities, which may be governed by a synergism between contemporary climate, productivity and habitat size. We also found evidence of a legacy of historical climate on functional diversity, independent from species richness. By contrast, path models of communities containing nonnative species demonstrated lower explanatory power and had no clear association with any of the abiotic or biotic factors. In conclusion, we demonstrated that strong spatial patterns in community structure and functional diversity of freshwater fish communities exist at the global scale, underlined by the complex interplay between external and internal filters, but that these patterns may be blurred by anthropogenic species introductions. Our results further highlighted the importance of accounting for realized species niches and species status (i.e., native and nonnative) when investigating questions related to the assembly and functional diversity of multitrophic communities.
Forecasted increases in terrestrial organic matter (OMterr) inputs to the Arctic Beaufort Sea necessitate a better understanding of the proportional contribution of this potential food source to the trophic structure of marine communities. This study investigated the relative ecological importance of OMterr across the Beaufort Sea shelf and slope by examining differences in community trophic structure concurrent with variation in terrestrial versus marine organic matter influence. Oxygen stable isotope ratios (18O) of surface water confirmed the widespread influence of Canada's Mackenzie River plume across the Beaufort Sea. Carbon stable isotope ratios (13C) of pelagic particulate organic matter (pPOM) and marine consumers indicated a significant decrease in OMterr presence and utilization by consumers with increasing distance from the Mackenzie River outflow. Food web length, based on the nitrogen stable isotope ratios (15N) of marine consumers, was greater closer to the Mackenzie River outflow both in shelf and slope locations, due to relatively higher 15N values of pelagic and benthic primary consumers. Strong microbial pro cessing of OMterr in the eastern regions of the Beaufort Sea is inferred based on a trophic gap between assumed end members and lower trophic consumers. A greater proportion of relative epifaunal biomass occupying higher trophic levels suggests that OMterr as a basal food source can provide substantial energetic support for higher marine trophic levels. These findings challenge the current conception of low terrestrial matter usage in the Arctic marine food web, and indicate the need for a more specific understanding of energy transfer through the OMterr-associated microbial loop.
Tight coupling between pelagic and benthic communities is accepted as a general principle on Arctic shelves. Whereas this paradigm has been useful for guiding ecological research, it has perhaps led to a disproportionate focus on POM and ice algae as the most likely sources of carbon for the benthic food web. Arctic shelves are complex systems, including banks, fjords, and trough systems up to 350 m or more in depth. In this stable-isotope study, 13 different potential carbon sources were analyzed for their contribution to the food-webs of Isfjorden, Svalbard. A mixing model with herbivorous copepods and grazing sea urchins as end-members was applied to determine the relative contributions of the most likely carbon sources to pelagic and benthic taxa. Most taxa from the benthos feed on a broad mixture of POM and macroalgal detritus, even at depths down to 410 m. Most suspension-feeding bivalves had isotopic signals consistent with more than a 50% contribution from kelps and rockweeds. In contrast, nearly all pelagic species had diets consistent with an overwhelming contribution of pelagic POM. These results indicate that macroalgal detritus can contribute significantly to near-shore Arctic food-webs, a trophic link that may increase if macroalgae increase in the Arctic as predicted. These weaker quantitative links between pelagic and benthic components of coastal systems highlight the need for thorough sampling of potential carbon-baselines in food-web studies. A large detrital-carbon component in diets of Arctic benthos may dampen the impacts of strong seasonality in polar primary producers, leading to higher ecosystem resilience, but may also result in lower secondary productivity.
In the preceding chapters, organic carbon sources, pathways and transformation processes at the sea floor have been presented and discussed. In the following chapter, particulate organic carbon in Arctic Ocean sediments, its composition, burial and paleoenvironmental significance will be discussed. With respect to the Arctic Ocean’s organic carbon sedimentary records, we concentrate on surface sediment data and sediment cores representing the late Quaternary time interval, because information about organic carbon accumulation during pre-Quaternary (i.e., Jurassic to Tertiary) and its paleoenvironmental implications is very sparse (e.g., Thiede et al. 1990). For completeness, however, a short overview on pre-Quaternary records will be presented here.
Trophic structure and resilience of regional benthic food webs were studied on the Alaska Beaufort Sea shelf across 3 geographic regions (east, central, and west) and 2 depth ranges (14 to 90 m [shallow] and 100 to 220 m [deep]). Carbon (δ13C) and nitrogen (δ15N) isotope ratios were measured from water column and sediment particulate organic matter (POM) as markers of primary food sources and from common benthic invertebrate taxa. Isotopic niche space (δ13C. δ15N) occupied by benthic taxa provided measures of community-wide trophic redundancy and separation. δ13C values of the water column and sediment POM were patchy but generally indicative of terrestrial input across the shelf, especially near the mouth of the Colville River (<-25.5‰). Food web structures in the shallow and deep western Beaufort regions, however, reflected marine carbon inputs, with most consumers occupying intermediate trophic levels (TLs) and food webs of intermediate trophic redundancy and separation relative to the other regions. In the central shallow and east regions, the terrestrial carbon found in sediments was mirrored in consumer tissues. The central deep region contained the highest proportions of higher TL taxa, particularly within the predator feeding guild, possibly reflecting resource partitioning of a limited prey spectrum. The comparatively large isotopic niche space in the central deep region, with high trophic niche separation but low trophic redundancy, suggests that this region may be most vulnerable to perturbations. This study provides a valuable new understanding of benthic food web dynamics in the understudied Alaska Beaufort Sea region and represents a baseline for future comparison.
Stable isotope analysis has become a common tool for mapping trophic relationships, describing foodweb changes, and assessing ecosystem health. Clear interpretation of stable isotopes is facilitated by understanding how environmental factors can affect isotopic values; in estuarine systems, these factors may include salinity, land use, and shoreline habitat. To evaluate these factors, fish were collected from shallow-water habitats next to hardened (bulkhead and riprap) and unhardened (beach and marsh) shorelines within five subestuaries of the Chesapeake Bay that differed in predominant land use and salinity. This study focused on three common mid-Atlantic fish species: mummichog, Fundulus heteroclitus, Atlantic silverside, Menidia menidia, and white perch, Morone americana. Multiple regression analyses pointed to standard length, salinity, of watershed as developed or crop land, and shoreline habitat type as important predictors for N-15 in all three species and for C-13 in mummichog and white perch. Further analysis controlling for the effects of salinity, land use, and fish size demonstrated that C-13 and N-15 were lower in tissues of fish collected next to marsh compared with hardened or beach habitat. Habitat effects were strongest for mummichog. This study focused on overarching patterns driving stable isotope signatures in fish; however, it also indicated potentially important interactions between nearshore habitat type and land use or salinity that deserve further analysis. Results have implications for the scale of isotope inquiry and give justification for more detailed follow-up studies of foodweb structure along modified and natural shorelines.
We analyzed bulk carbon and nitrogen stable isotope values (δ13C and δ15N) of the benthic clams Nuculana radiata and Macoma moesta from the Bering Sea during controlled feeding experiments (spring of 2009 and 2010) using isotopically labeled sea ice algae. The aim was to determine the ability of these clam species to assimilate carbon and nitrogen from sea ice algae. Specimens were collected in the Bering Sea and placed into jars without sediment (2009, N. radiata only) or into natural sediment cores (2010, both species). The clams were offered isotopically enriched (both C and N) or non-enriched algal feeds for time periods of 42 (2009) and 18 d (2010). Isotopic assimilation rates for carbon and nitrogen were calculated using the change in the isotope ratios of the clams over the experimental time. N. radiata in the jar experiments had slow isotopic assimilation rates (0.01 to 0.23‰ d− 1), with solvent-extractable organic matter/lipids taking up both of the isotope markers fastest and muscle tissue the slowest. Lipids may thus be particularly suitable to track the immediate ingestion of sea ice algal production in benthic consumers. M. moesta showed 30% higher isotopic assimilation compared to N. radiata in sediment cores, likely reflecting the different feeding behaviors of these two species. Based on our results, N. radiata is likely better able to utilize food sources buried in the sediment and may be more competitive over the sediment surface feeding M. moesta under conditions of reduced ice algal production in the northern Bering Sea.
Arctic sea ice decline has led to an amplification of surface warming and is projected to continue to decline from anthropogenic forcing, although the exact timing of ice-free summers is uncertain owing to large natural variability. Sea ice reductions affect surface heating patterns and the atmo-spheric pressure distribution, which may alter midlatitude extreme weather patterns. Increased light penetration and nutrient availability during spring from earlier ice breakup enhances primary production in the Arctic Ocean and its adjacent shelf seas. Ice-obligate marine mammals may be losers, Further ANNUAL REVIEWS EG39CH03-Bhatt ARI 7 October 2014 12:33 whereas seasonally migrant species may be winners from rapid sea ice decline. Tundra greening is occurring across most of the Arctic, driven primarily by warming temperatures, and is display-ing complex spatial patterns that are likely tied to other factors. Sea ice changes are affecting greenhouse gas exchanges as well as halogen chemistry in the Arctic. This review highlights the heterogeneous nature of Arctic change, which is vital for researchers to better understand.
Environmental gradients and their influence on benthic community structure vary over different spatial scales; yet, few studies in the Arctic have attempted to study the influence of environmental gradients of differing spatial scales on megabenthic communities across continental-scales. The current project studied for the first time how megabenthic community structure is related to several environmental factors over 2000 km of the Canadian Arctic, from the Beaufort Sea to northern Baffin Bay. Faunal trawl samples were collected between 2007 and 2011 at 78 stations from 30 to 1000 m depth and patterns in biomass, density, richness, diversity, and taxonomic composition were examined in relation to indirect/spatial gradients (e.g., depth), direct gradients (e.g., bottom oceanographic variables), and resource gradients (e.g., food supply proxies). Six benthic community types were defined based on their biomass-based taxonomic composition. Their distribution was significantly, but moderately, associated with large-scale (100-1000 km) environmental gradients defined by depth, physical water properties (e.g., bottom salinity), and meso-scale (10-100 km) environmental gradients defined by substrate type (hard vs. soft) and sediment organic carbon content. We did not observe a strong decline of bulk biomass, density and richness with depth or a strong increase of those community characteristics with food supply proxies, contrary to our hypothesis. We discuss how local- to meso-scale environmental conditions, such as bottom current regimes and polynyas, sustain biomass-rich communities at specific locations in oligotrophic and in deep regions of the Canadian Arctic. This study demonstrates the value of considering the scales of variability of environmental gradients when interpreting their relevance in structuring of communities.
Stable isotopes have been used extensively to
study food-web functioning, that is, the flow of energy and
matter among organisms. Traditional food-web studies are
based on the natural variability of isotopes and are limited to
larger organisms that can be physically separated from their
environment. Recent developments allow isotope ratio measurements
of microbes and this in turn allows the measurement
of entire food webs, in other words, from small producers
at the bottom to large consumers at the top. Here, I
provide a concise review on the use and potential of stable
isotopes to reconstruct end-to-end food webs. I will first discuss
food web reconstruction based on natural abundances
isotope data and will then show that the use of stable isotopes
as deliberately added tracers provides complementary information.
Finally, challenges and opportunities for end-to-end
food web reconstructions in a changing world are discussed.
Recently revisited, the concept of niche ecology has lead to the formalisation of functional and trophic niches using stable isotope ratios. Isotopic diversity indices (IDI) derived from a set of measures assessing the dispersion/distribution of points in the δ-space were recently suggested and increasingly used in the literature. However, three main critics emerge from the use of these IDI: 1) they fail to account for the isotopic sources overlap, 2) some indices are highly sensitive to the number of species and/or the presence of rare species, and 3) the lack of standardization prevents any spatial and temporal comparisons. Using simulations we investigated the ability of six commonly used IDI to discriminate among different trophic food web structures, with a focus on the first two critics. We tested the sensitivity of the IDI to five food web structures along a gradient of sources overlap, varying from two distinct food chains with differentiated sources to two superimposed food chains sharing two sources. For each of the food web structure we varied the number of species (from 10 to 100 species) and the type of species feeding behaviour (i.e. random or selective feeding). Values of IDI were generally larger in food webs with distinct basal sources and tended to decrease as the superimposition of the food chains increased. This was more pronounced when species displayed food preferences in comparison to food webs where species fed randomly on any prey. The number of species composing the food web also had strong effects on the metrics, including those that were supposedly less sensitive to small sample size. In all cases, computing IDI on food webs with low numbers of species always increases the uncertainty of the metrics. A threshold of ∼20 species was detected above which several metrics can be safely used.
ABSTRACT: Understanding the mechanisms that support feeding interactions and species cooccurrence in regions subject to rapid environmental changes is becoming increasingly important to predict future trends in population dynamics. However, there is still little information available on the trophic ecology for many benthic species to help us better understand trophic interactions and individual trophic roles. Here, we used stable isotopes (δ13C, δ15N) in conjunction with the
Bayesian ellipses approach to explore spatial trends in isotopic niche width and overlap of 3 syntopic arctic brittle stars (Echinodermata: Ophiuroidea; Ophiacantha bidentata, Ophiocten sericeum, and Ophiopleura borealis) in Baffin Bay (BB), the Canadian Arctic Archipelago (CAA), and the North Water Polynya (NOW). These 3 coexisting ophiuroids displayed great interspecific plasticity in foraging behaviors and showed a high degree of inter-individual dietary flexibility. However,
differences in surface carbon composition drove the variability of resource utilization at the individual level across stations, which in turn affected trophic interactions, niche overlaps, and isotopic niche breadth of ophiuroids. Greater niche overlap was found in the highly productive
region of the NOW, where consumers exhibited similar food selectivity, whereas an increase in niche segregation occurred in regions with greater sea-ice concentration. These results suggest that isotopic niche size reflects individual responses to fluctuations in food availability and possibly past competition, both induced by local oceanographic features. Our study indicates that niche parameters of ophiuroids can respond quickly to ecological and environmental gradients, which suggests important adaptability of these species facing multiple stressors.
Antarctic sea stars can occupy different trophic niches and display different trophic levels, but, while the impacts of their body size and environmental features on their trophic niches are potentially important, they are presently understudied. Here we assessed the trophic ecology in relation to the size and habitat of sea stars in a fjord on King George Island (South Shetland Islands) using stable isotope values of carbon (δ13C), nitrogen (δ15N), and sulphur (δ34S). The disc radius influenced δ13C and δ15N values, whereas more limited changes in δ13C or δ34S values were related to arm length. Specifically, δ13C and δ15N values were linked to disc radius in generalist species (Diplasterias brandti and Odontaster validus), which could indicate ontogenetic diet shifts, while this relationship occurred less frequently in more specialised species (Bathybiaster loripes, Notasterias bongraini, and Perknaster sladeni). O. validus had a smaller isotopic niche size in the inner than the outer fjord. The niche overlap between D. brandti and O. validus was low in the inner fjord. Low resource availability within the fjord, linked to higher turbidity, could induce trophic niche constriction and interspecific resource segregation. This could represent a mechanism for competition avoidance in a resource-limited system. Conversely, higher resource availability could allow O. validus to expand and share its isotopic niche with D. brandti in the outer fjord with a limited risk of competition. This trophic plasticity will likely influence how O. validus copes with the present and future modification of environmental conditions induced by climate change.
Stable isotope ratios of nitrogen (δ¹⁵N) of benthic primary consumers are often significantly related to water depth. This relationship is commonly attributed to preferential uptake of ¹⁴N from sinking particulate organic matter (POM) by microbes, and suggests that relationships between δ¹⁵N and water depth may be affected by local POM sources and flux dynamics. We examined the relationships between δ¹⁵N and water depth (20–500 m) for six trophic functional groups using a mixed effects modelling approach, and compared relationships between two contiguous Arctic marine ecosystems with different POM sources and sinking export dynamics: the Canadian Beaufort Sea and Amundsen Gulf. We demonstrate for the first time in the Arctic that δ¹⁵N values of mobile epifaunal carnivores increased as a function of depth when considered separately from benthopelagic and infaunal carnivores, which contrarily did not exhibit increasing δ¹⁵N with depth. The δ¹⁵N of suspension/filter feeders, infaunal deposit feeders and bulk sediment also increased with water depth, and the slopes of the relationships were steeper in the Amundsen Gulf than in the Beaufort Sea. We propose that regional differences in slopes reflect differences in POM sources exported to the benthos. In the Beaufort Sea, terrestrial POM discharged from the Mackenzie River quantitatively dominates the sedimentary organic matter across the continental shelf and slope, dampening change in δ¹⁵N of benthic POM with depth. In the Amundsen Gulf, we attribute a faster rate of change in δ¹⁵N of POM with increasing depth to larger contributions of marine-derived POM to the benthic sedimentary pool, which had likely undergone extensive biological transformation in the productive offshore pelagic zone. Differences in POM input regimes among regions should be considered when comparing food webs using stable isotopes, as such differences may impact the rate at which consumer δ¹⁵N changes with depth.
Phenological cascades can occur in food webs when the timing of biological activity at one trophic level responds to timing at an adjacent trophic level. Such cascades may therefore be precipitated by changes in abiotic factors that constrain phenology. Arctic marine food webs comprise a series of coupled trophic interactions from primary producers to tertiary consumers under strong temporal control by the annual timing of sea ice melt onset, thinning, and retreat. Under recent climatic warming, the extent, thickness, and age of arctic sea ice have all declined markedly. Simultaneously, the annual timing of sea ice melt has advanced across the Arctic at a rate of − 2 days per decade, with individual regions experiencing advances in melt onset of nearly 25 to 30 days since 1979. This review highlights phenological sensitivity across trophic levels to earlier onset of the annual productive season in arctic marine food webs related to advancing timing of sea ice melt. Phenological dynamics are evident from organisms with the simplest life cycles in these food webs, algae and phytoplankton, to those with the most complex life cycles, bowhead and beluga whales. Phenological responses of arctic marine organisms to the timing of sea ice melt may, however, be stronger at lower trophic levels than at higher trophic levels. Under continued warming, maintenance of the integrity of arctic marine food webs will be sensitive not only to the increasing loss of sea ice cover during summer, but also to increasingly earlier timing of annual sea ice melt onset.
The abiotic degradation state of sea ice algae released during a late spring ice melt process was determined by sampling the underlying waters and measuring certain well-known algal lipids and their oxidation products, including those derived from epi-brassicasterol, 24-methylenecholesterol, palmitoleic acid and the phytyl side-chain of chlorophyll. More specifically, parent lipids and some of their oxidation products were quantified in suspended (collected by filtration) and sinking (collected with sediment traps at 5 and 30 m) particles from Resolute Passage (Canada) during a period of spring ice melt in 2012 and the outcomes compared with those obtained from related sea ice samples analyzed previously. Our data show that suspended cells in the near surface waters appeared to be only very weakly affected by photooxidative processes, likely indicative of a community of unaggregated living cells with high seeding potential for further growth. In contrast, we attribute the strong photooxidation state of the organic matter in the sediment traps deployed at 5 m to the presence of senescent and somewhat aggregated sea ice algae that descended only relatively slowly within the euphotic zone, and was thus susceptible to photochemical degradation. On the other hand, the increased abiotic preservation of the sinking material collected in the sediment traps deployed at 30 m, likely reflected more highly aggregated senescent sea ice algae that settled sufficiently rapidly out of the euphotic zone to avoid significant photooxidation. This better-preserved sinking material in the deeper sediment traps may therefore contribute more strongly to the underlying sediments. A three-component conceptual scheme summarizing the abiotic behavior of Arctic sea ice algae in underlying waters is proposed.
Predicting the ability of the biosphere to continue to deliver ecosystem services in the face of biodiversity loss and environmental change is a major challenge. The results of short-term and small-scale experimental studies are both equivocal and difficult to extrapolate from. In this study we use data on benthic palaeocommunities covering 4,000,000 years (in the Late Jurassic when temperate coastal seas in NW Europe experienced fluctuations in oxygenation). The biological traits associated with each species in the palaeocommunities were combined to index the delivery of ecological functions. Five ecosystem functions were examined: food for large mobile predators, biogenic habitat provision, nutrient recycling/regeneration, inorganic carbon sequestration and food-web dynamics. In modern systems these ecological functions underpin ecosystem services that are important for human well-being. Our results show that the supply of food for higher predators was remarkably constant during the 4,000,000 years, suggesting that redundancy amongst species in the assemblage drives the biodiversity–ecosystem function (BEF) relationship. By contrast, the provision of biogenic habitat varied with the occurrence of a relatively few taxa, a pattern consistent with a rivet type model of BEF. For nutrient regeneration, carbon sequestration and food-web dynamics the patterns were complex and suggestive of an idiosyncratic model of BEF. To our knowledge this is the first study to quantify ecological functioning through deep time and demonstrates the utility of this approach to understanding long-term patterns of BEF in both ancient and contemporary marine ecosystems. The delivery of all five ecological functions studied became increasingly variable as the regional climate became drier, thus modifying the supply of terrigenous nutrient inputs.
Marine bio-production in the Arctic and Subarctic is based primarily on microscopic unicellular algae (phytoplankton) in the water column and micro-algae associated with ice. Algae utilise light to produce organic matter by photosynthesis, thereby reducing CO2 while releasing O2 and producing carbohydrates which, according to the needs of the algae, can be converted into essential compounds such as proteins and nucleic acids by incorporating nitrogen, phosphorus, sulphur, and other elements.
A key to many scientific studies of ocean and sea floor processes is an accurate description of the sea floor morphology. For Arctic scientists this has posed a particular problem because less is known about the Arctic Ocean bathymetry and physiography than for the other oceans. For nearly 60 years after Nansen’s collection of deep bathymetric soundings during his epic expedition with the vessel Fram drifting in the Arctic pack ice from the New Siberian Islands to the Fram Strait (Nansen 1902), it was a common belief that the entire central Arctic Ocean consisted of one deep basin. The perennial sea ice cover, within which Fram drifted, has severely hampered the systematic collection of bathymetric information from surface vessels. Following World War II the former Soviet Union started a sparse, but systematic, collection of bathymetric soundings in the central Arctic from ice stations established at great risk using airplanes to land scientists on the pack ice in order to carry out soundings through the ice.
Introduction.- Data management and software.- Advice for teachers.- Exploration.- Linear regression.- Generalised linear modelling.- Additive and generalised additive modelling.- Introduction to mixed modelling.- Univariate tree models.- Measures of association.- Ordination--first encounter.- Principal component analysis and redundancy analysis.- Correspondence analysis and canonical correspondence analysis.- Introduction to discriminant analysis.- Principal coordinate analysis and non-metric multidimensional scaling.- Time series analysis--Introduction.- Common trends and sudden changes.- Analysis and modelling lattice data.- Spatially continuous data analysis and modelling.- Univariate methods to analyse abundance of decapod larvae.- Analysing presence and absence data for flatfish distribution in the Tagus estuary, Portugual.- Crop pollination by honeybees in an Argentinean pampas system using additive mixed modelling.- Investigating the effects of rice farming on aquatic birds with mixed modelling.- Classification trees and radar detection of birds for North Sea wind farms.- Fish stock identification through neural network analysis of parasite fauna.- Monitoring for change: using generalised least squares, nonmetric multidimensional scaling, and the Mantel test on western Montana grasslands.- Univariate and multivariate analysis applied on a Dutch sandy beach community.- Multivariate analyses of South-American zoobenthic species--spoilt for choice.- Principal component analysis applied to harbour porpoise fatty acid data.- Multivariate analysis of morphometric turtle data--size and shape.- Redundancy analysis and additive modelling applied on savanna tree data.- Canonical correspondence analysis of lowland pasture vegetation in the humid tropics of Mexico.- Estimating common trends in Portuguese fisheries landings.- Common trends in demersal communities on the Newfoundland-Labrador Shelf.- Sea level change and salt marshes in the Wadden Sea: a time series analysis.- Time series analysis of Hawaiian waterbirds.- Spatial modelling of forest community features in the Volzhsko-Kamsky reserve.
[1] The Arctic-wide melt season has lengthened at a rate of 5 days dec-1 from 1979 to 2013, dominated by later autumn freeze-up within the Kara, Laptev, East Siberian, Chukchi and Beaufort seas between 6 and 11 days dec-1. While melt onset trends are generally smaller, the timing of melt onset has a large influence on the total amount of solar energy absorbed during summer. The additional heat stored in the upper ocean of approximately 752 MJ m-2 during the last decade, increases sea surface temperatures by 0.5 to 1.5 °C and largely explains the observed delays in autumn freeze-up within the Arctic Ocean's adjacent seas. Cumulative anomalies in total absorbed solar radiation from May through September for the most recent pentad locally exceed 300-400 MJ m-2 in the Beaufort, Chukchi and East Siberian seas. This extra solar energy is equivalent to melting 0.97 to 1.3 m of ice during the summer.
Sea ice is exchanged between the Arctic Ocean and Canadian Arctic Archipelago (CAA) but has not been quantified over long time periods. The corresponding mechanisms responsible for recent variability and change also remain unidentified. To address this, we estimated the sea ice area flux between the Arctic Ocean and the M’Clure Strait and Queen Elizabeth Islands (QEI) from 1997 to 2012 for the months of May to November. Over the period, there was a mean flux of -1x10^3 km^2 (±21x10^3 km^2) at the M’Clure Strait and mean flux of +8x10^3 km^2 (±8x10^3 km^2) at the QEI (positive and negative flux signs correspond to Arctic Ocean ice inflow and outflow, respectively). The M’Clure Strait had a mean flux of +5x10^3 km^2 from May to September and a mean flux of -7x10^3 km^2 from October to November. The QEI gates had a mean flux of +4 x10^3 km^2 from August to September with negligible ice exchange from May to July and October to November. More frequent high sea level pressure anomalies over the Beaufort Sea and Canadian Basin since 2007 have reduced Arctic Ocean multiyear ice (MYI) inflow into the M’Clure Strait. The presence of MYI in the CAA originating from the Arctic Ocean has been maintained by inflow at the QEI, which has increased since 2005. These recent increases in Arctic Ocean MYI inflow into the QEI can be attributed to increased open water area within the CAA that have provided more leeway for inflow to occur.
1] Previous comprehensive investigations of the Canadian Arctic revealed that subsurface chlorophyll maxima (SCM) are widespread and long-lived structures that can contribute significantly to daily primary production in the water column. However, estimating the annual contribution of SCM to production with in situ or remote-sensing approaches is challenging in the high Arctic. For this reason and to estimate the impacts of fluctuating or changing environmental conditions on SCM, a numerical approach combining a turbulence model and an ecosystem model was implemented for the coastal Beaufort Sea. An ensemble analysis of simulations suggested that SCM contribute 65–90% of total annual primary production and that this proportion is weakly affected by ice regime, winter nitrogen (N) concentration, parameter values determining phytoplankton growth and decay or the physical forcing imposed, all varying within realistic values. Due to the persistent association between the SCM and the shallow nitracline, the pelagic ecosystem of the coastal Beaufort Sea is apparently characterized by a high ratio of new to total production, contrasting with the common assumption that oligotrophic systems are predominantly supported by recycled N and regenerated production. This study demonstrated that the use of a simple model in combination with in situ data leads to novel insights into biogeochemical processes that are otherwise very difficult to measure and track.