Declining calcium concentration drives shifts toward smaller and less nutritious zooplankton in northern lakes.

Zooplankton community composition of northern lakes is changing due to the interactive effects of climate change and recovery from acidification, yet limited data are available to assess these changes combined. Here, we built a database using archives of temperature, water chemistry and zooplankton data from 60 Scandinavian lakes that represent broad spatial and temporal gradients in key parameters: temperature, calcium (Ca), total phosphorus (TP), total organic carbon (TOC), and pH. Using machine learning techniques, we found that Ca was the most important determinant of the relative abundance of all zooplankton groups studied, while pH was second, and TOC third in importance. Further, we found that Ca is declining in almost all lakes, and we detected a critical Ca threshold in lake water of 1.3 mg L-1 , below which the relative abundance of zooplankton shifts toward dominance of Holopedium gibberum and small cladocerans at the expense of Daphnia and copepods. Our findings suggest that low Ca concentrations may shape zooplankton communities, and that current trajectories of Ca decline could promote widespread changes in pelagic food webs as zooplankton are important trophic links from phytoplankton to fish and different zooplankton species play different roles in this context.

Fatty-acid based assessment of benthic food-web responses to multiple stressors in a large river system.

Rivers are often exposed to multiple stressors, such as nutrients and contaminants, whose impacts on the river food webs may not be distinguished by sole assessment of biological community structures. We examined the benthic algal assemblages and the fatty acids (FA) of benthic macroinvertebrates in the lower Athabasca River in Canada, aiming to assess the changes in algal support and nutritional quality of the benthic food web in response to cumulative exposure to natural bitumen, municipal sewage discharge (hereafter, "sewage"), and oil sands mining ("mining"). Data show that the decline in water quality (increases in nutrient concentrations and total suspended solids) was associated with decreases in benthic diatom abundance, and was driven mainly by sewage-induced nutrient enrichment. Responses in nutritional quality of benthic macroinvertebrates, indicated by their polyunsaturated FA (PUFA) concentrations, were taxon- and stressor-specific. Nutritional quality of the larval dragonfly predator, Ophiogomphus, decreased nonlinearly with decreasing benthic diatom abundance and was lowest at the sewage-affected sites, although exposure to natural bitumen also resulted in reduced Ophiogomphus PUFA concentrations. In contrast, the PUFA concentrations of mayfly grazers/collector-gatherers were not affected by natural bitumen exposure, and were higher at the sewage and sewage+mining sites. The PUFA concentrations of the shredder Pteronarcys larvae did not change with cumulative exposure to the stressors. Sediment metal and polycyclic aromatic compound concentrations were not associated with the macroinvertebrate FA changes. Overall, we provide evidence that sewage induced reduction in trophic support by PUFA-rich diatoms, and was the predominant driver of the observed changes in FA composition and nutritional quality of the benthic macroinvertebrates. Fatty-acid metrics are useful to untangle effects of concurrent stressors, but the assessment outcomes depend on the functional feeding guilds used. A food-web perspective using multiple trophic levels and feeding guilds supports a more holistic assessment of the stressor impacts.

Autochthony and isotopic niches of benthic fauna at shallow-water hydrothermal vents.

The food webs of shallow-water hydrothermal vents are supported by chemosynthetic and photosynthetic autotrophs. However, the relative importance of these two basal resources for benthic consumers and its changes along the physicochemical gradient caused by vent plumes are unknown. We used stable carbon and nitrogen isotopes (i.e., δ13C and δ15N) and Bayesian mixing models to quantify the dietary contribution of basal resources to the benthic fauna at the shallow-water vents around Kueishan Island, Taiwan. Our results indicated that the food chains and consumer production at the shallow-water vents were mainly driven by photoautotrophs (total algal contribution: 26-54%) and zooplankton (19-34%) rather than by chemosynthetic production (total contribution: 14-26%). Intraspecific differences in the trophic support and isotopic niche of the benthic consumers along the physicochemical gradient were also evident. For instance, sea anemone Anthopleura sp. exhibited the greatest reliance on chemosynthetic bacteria (26%) and photoautotrophs (66%) near the vent openings, but zooplankton was its main diet in regions 150-300 m (32-49%) and 300-700 m (32-78%) away from the vent mouths. The vent-induced physicochemical gradient structures not only the community but also the trophic support and isotopic niche of vent consumers.

Lowered nutritional quality of plankton caused by global environmental changes.

Global environmental changes are causing widespread nutrient depletion, declines in the ratio of dissolved inorganic nitrogen (N) to total phosphorus (DIN:TP), and increases in both water temperature and terrestrial colored dissolved organic carbon (DOC) concentration (browning) in high-latitude northern lakes. Declining lake DIN:TP, warming, and browning alter the nutrient limitation regime and biomass of phytoplankton, but how these stressors together affect the nutritional quality in terms of polyunsaturated fatty acid (PUFA) contents of the pelagic food web components remains unknown. We assessed the fatty acid compositions of seston and zooplankton in 33 lakes across south-to-north and boreal-to-subarctic gradients in Sweden. Data showed higher lake DIN:TP in the south than in the north, and that boreal lakes were warmer and browner than subarctic lakes. Lake DIN:TP strongly affected the PUFA contents-especially eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA)-in seston, calanoids, and copepods (as a group), but not in cladocerans. The EPA+DHA contents increased by 123% in seston, 197% in calanoids, and 230% in copepods across a lake molar DIN:TP gradient from 0.17 to 14.53, indicating lower seston and copepod nutritional quality in the more N-limited lakes (those with lower DIN:TP). Water temperature affected EPA+DHA contents of zooplankton, especially cladocerans, but not seston. Cladoceran EPA+DHA contents were reduced by ca. 6% for every 1°C increase in surface water. Also, the EPA, DHA, or EPA+DHA contents of Bosmina, cyclopoids, and copepods increased in lakes with higher DOC concentrations or aromaticity. Our findings indicate that zooplankton food quality for higher consumers will decrease with warming alone (for cladocerans) or in combination with declining lake DIN:TP (for copepods), but impacts of these stressors are moderated by lake browning. Global environmental changes that drive northern lakes toward more N-limited, warmer, and browner conditions will reduce PUFA availability and nutritional quality of the pelagic food web components.

Inverting nutrient fluxes across the land-water interface - Exploring the potential of zebra mussel (Dreissena polymorpha) farming.

We studied the potential of zebra mussel farming for nutrient retention in a eutrophic lake. Duplicate experimental long-line cultivation units were deployed and mussel growth and nutrient retention were quantified after 28 months. Mussels grew well at shallow water depth (<3 m) and our 625 m2 (lake area) experimental units produced 507 and 730 kg dry biomass, respectively, of which 94% were shells. These yields corresponded to an average retention of 92.7 ± 23.1 kg C, 6.1 ± 0.68 kg N, and 0.43 ± 0.04 kg P retention, or 742 kg C, 49 kg N, and 3.5 kg P for a full-size (0.5 ha) mussel farm. We estimate that concentrating the long-lines to a depth of 2.5 m would probably have doubled these yields, based on the differences in mussel growth among depths. We further estimate that a full-size cultivation unit (0.5 ha) thus could compensate for the annual total-P run-off from 23 ha, or the biologically available P from approximately 49 ha of agricultural soils. As traditional measures have proven insufficient, decision-makers need to facilitate novel approaches to mitigate the negative effects of cultural eutrophication. We envision that zebra mussel farming, within their invaded range, provides a promising approach to invert nutrient losses in lakes and coastal lagoons.

Bromoanisoles and methoxylated bromodiphenyl ethers in macroalgae from Nordic coastal regions.

Marine macroalgae are used worldwide for human consumption, animal feed, cosmetics and agriculture. In addition to beneficial nutrients, macroalgae contain halogenated natural products (HNPs), some of which have toxic properties similar to those of well-known anthropogenic contaminants. Sixteen species of red, green and brown macroalgae were collected in 2017-2018 from coastal waters of the northern Baltic Sea, Sweden Atlantic and Norway Atlantic, and analyzed for bromoanisoles (BAs) and methoxylated bromodiphenyl ethers (MeO-BDEs). Target compounds were quantified by gas chromatography-low resolution mass spectrometry (GC-LRMS), with qualitative confirmation in selected species by GC-high resolution mass spectrometry (GC-HRMS). Quantified compounds were 2,4-diBA, 2,4,6-triBA, 2'-MeO-BDE68, 6-MeO-BDE47, and two tribromo-MeO-BDEs and one tetrabromo-MeO-BDE with unknown bromine substituent positions. Semiquantitative results for pentabromo-MeO-BDEs were also obtained for a few species by GC-HRMS. Three extraction methods were compared; soaking in methanol, soaking in methanol-dichloromethane, and blending with mixed solvents. Extraction yields of BAs did not differ significantly (p > 0.05) with the three methods and the two soaking methods gave equivalent yields of MeO-BDEs. Extraction efficiencies of MeO-BDEs were significantly lower using the blend method (p < 0.05). For reasons of simplicity and efficiency, the soaking methods are preferred. Concentrations varied by orders of magnitude among species: ∑2BAs 57 to 57 700 and ∑5MeO-BDEs < 10 to 476 pg g-1 wet weight (ww). Macroalgae standing out with ∑2BAs >1000 pg g-1 ww were Ascophyllum nodosum, Ceramium tenuicorne, Ceramium virgatum, Fucus radicans, Fucus serratus, Fucus vesiculosus, Saccharina latissima, Laminaria digitata, and Acrosiphonia/Spongomorpha sp. Species A. nodosum, C. tenuicorne, Chara virgata, F. radicans and F. vesiculosus (Sweden Atlantic only) had ∑5MeO-BDEs >100 pg g-1 ww. Profiles of individual compounds showed distinct differences among species and locations.

Autochthonous resources are the main driver of consumer production in dystrophic boreal lakes.

Dystrophic lakes are widespread in temperate regions and intimately interact with surrounding terrestrial ecosystems in energy and nutrient dynamics, yet the relative importance of autochthonous and allochthonous resources to consumer production in dystrophic lakes remains controversial. We argue that allochthonous organic matter quantitatively dominates over photosynthetic autotrophs in dystrophic lakes, but that autotrophs are higher in diet quality and more important for consumers as they contain essential polyunsaturated fatty acids (PUFA). In a field study, we tested the hypotheses that (1) autochthonous primary production is the main driver for consumer production, despite being limited by light availability and low nutrient supplies, and greater supply of allochthonous carbon, (2) the relative contribution of autotrophs to consumers is directly related to their tissue PUFA concentrations, and (3) methane-oxidizing bacteria (MOB) provide an energy alternative for consumers. Pelagic and benthic consumer taxa representing different trophic levels were sampled from five dystrophic lakes: isopod Asellus aquaticus, megalopteran Sialis lutaria, dipteran Chaoborus flavicans, and perch Perca fluviatilis. Based on carbon and nitrogen stable isotopes, the relative contributions of autochthonous (biofilms and seston) and allochthonous (coarse particulate and dissolved organic matter) resources and MOB to these taxa were 47-79%, 9-44% and 7-12% respectively. Results from fatty acid (FA) analyses show that the relative omega3-FA and PUFA concentrations increased with trophic level (Asellus < Sialis and Chaoborus < Perca). Also, eicosapentaenoic-acid (EPA), omega3-FA and PUFA concentrations increased with the autochthonous contribution in consumers, i.e., a 47-79% biofilm and/or seston diet resulted in tissue EPA of 4.2-18.4, omega3 FAs of 11.6-37.0 and PUFA of 21.6-61.0 mg/g dry mass. The results indicate that consumers in dystrophic lakes predominantly rely on energy from autotrophs and that their PUFA concentrations are dependent on the relative contribution of these autochthonous resources. The limited energy support from MOB suggests they are not negligible and are potentially an integral part of the food webs. Our findings show that autochthonous resources are the main driver of secondary production even in dystrophic lakes and offer new insights into the functioning of these ecosystems.

Evidence of rapid shifts in the trophic base of lotic predators using experimental dietary manipulations and assimilation-based analyses.

Assimilation-based techniques such as stoichiometric analysis, bulk tissue stable isotope analysis (SIA), fatty acid (FA) profiling and compound-specific SIA of FAs can be used to resolve ambiguities in consumer-resource relationships, but comparisons of their effectiveness are lacking. Feeding trials and concurrent application of these techniques were used to investigate the trophic base of two shrimps from Hong Kong streams: the atyid Caridina cantonensis, which is a primary consumer, and predatory Macrobrachium hainanense (Palaemonidae). Leaf litter and periphyton were fed to C. cantonensis (CC-LF and CC-PF, respectively) reared in the laboratory for 2 months, when C. cantonensis, that had fed on a mixed diet (CC-WC), were collected from the field. Atyids from each group (CC-LF, CC-PF and CC-WC) were fed to M. hainanense (MH-L, MH-P and MH-W, respectively) during a further 2-month trial, at the end of which M. hainanense were also collected from the field (MH-R). FA biomarkers present in CC-WC and CC-PF indicated that C. cantonensis depended primarily on autochthonous foods, and FA profiles of CC-WC were distinct from those of CC-LF. Differences in C/N ratios and isotope signatures of leaf litter and periphyton were not reflected in tissues of atyids or palaemonid predators. FA profiles of M. hainanense groups were similar, but FA stable C isotope ratio (delta(13)C) signatures of MH-R and MH-W were distinct from those of MH-L and leaf litter. FA delta(13)C signatures of MH-R were similar to those of MH-P and MH-W, indicating that autochthonous resources constituted the trophic base of production for this predator. This is the first study using compound-specific SIA to study stream food chains. Compound-specific SIA and FA profiling allowed accurate elucidation of consumer-resource relationships that were not revealed by stoichiometry or bulk tissue SIA. Compound-specific SIA was particularly sensitive for detecting rapid shifts in the predator trophic base. This tool will have wide applicability for investigating food webs in a range of ecosystems.