Emergent Aquatic Insects Research Articles

Shift in diet composition of a riparian predator along a stream pollution gradient.

Terrestrial insectivores in riparian areas, such as spiders, can depend on emergent aquatic insects as high-quality prey. However, chemical pollution entering streams from agricultural and urban sources can alter the dynamics and composition of aquatic insect emergence, which may also affect the riparian food web. Few studies have examined the effects of stressor-induced alterations in aquatic insect emergence on spiders, especially in terms of chemical pollution and diet composition. We used DNA metabarcoding of gut content to describe the diet of Tetragnatha montana spiders collected from 10 forested streams with differing levels of pesticide and wastewater pollution. We found that spiders consumed more Chironomidae and fewer other aquatic Diptera, including Tipulidae, Ptychopteridae and Culicidae, at more polluted streams. Pollution-related effects were mainly observed in the spider diet, and were not significant for the number nor composition of flying insects trapped at each site. Our results indicate that the composition of riparian spider diets is sensitive to stream pollution, even in the absence of a change in the overall proportion of aquatic prey consumed. A high reliance on aquatic prey at polluted streams may give spiders an increased risk of dietary exposure to chemical pollutants retained by emergent insects.

Effects of Taxon and Body Size on Mercury Concentrations in Spiders from Two Rivers with Different Levels of Mercury Contamination: Implications for the Use of Riparian Spiders as Sentinels.

Due to widespread atmospheric deposition of mercury (Hg), all aquatic food webs are contaminated with toxic methyl mercury (MeHg). At high concentrations, MeHg poses a health hazard to wildlife and humans. Spiders feeding in riparian habitats (hereafter referred to as riparian spiders) have been proposed as sentinels of MeHg contamination of aquatic systems. Riparian spiders are exposed to MeHg through their diets, and the concentration of MeHg in spiders is positively related to the proportion of MeHg-contaminated emergent aquatic insects in their diets. The use of spiders as sentinels is complex because their MeHg concentrations can vary, not only among ecosystems but also between different spider taxa and as a function of spider body size. The objective of the present study was to examine how the level of ecosystem contamination, spider taxon, and spider body size interact to influence MeHg concentrations in four genera of riparian spiders from two rivers with different levels of Hg contamination. We collected four genera of riparian spiders (Tetragnatha sp., Larinioides sp., Pardosa sp., and Rabidosa sp.) from two sites along both the Clear Fork of the Trinity River and the West Fork of the Trinity River (Fort Worth, TX, USA). We analyzed concentrations of MeHg in different body sizes of spiders from each genus. We found that MeHg contamination of the river ecosystem, spider taxon, and spider body size were important determinants of MeHg concentration in riparian spiders. The results suggest that any of the four taxa of riparian spiders from the present study could be used as sentinels of aquatic MeHg contamination, but they should not be used interchangeably because of the interdependence between the effects of ecosystem contamination level, spider taxon, and body size. Future studies utilizing riparian spiders as sentinels of biomagnifying aquatic contaminants (e.g., MeHg, polychlorinated biphenyls) should consider the potentially complex interaction effects between ecosystem contamination level, spider taxon, and spider body size. Environ Toxicol Chem 2024;00:1-7. © 2024 The Author(s). Environmental Toxicology and Chemistry published by Wiley Periodicals LLC on behalf of SETAC.

Drivers of spatio-temporal variation in organic matter decomposition across a river network

Rivers are biodiverse ecosystems that play an essential role in the processing of organic matter from headwaters to the lower reaches. The biodiversity within these spatially complex ecosystems is often subject to huge seasonal variation, particularly in temperate regions, for example seasonal peaks in emergence of aquatic insects or migration of fish through the river network. However, these ecosystems are also subject to strong anthropogenic pressures that affect not only the biodiversity within this system, but the functions this biodiversity provides to key ecosystem processes. Therefore, spatial and temporally dynamics must be considered to effectively assess and understand the biodiversity and functioning within an ecosystem. Here, we assessed ecosystem function at a high spatial and temporal resolution across a large river network, specifically focussing on standardised organic–matter decomposition. Our results show—even when correcting for temperature—significant effects of both season and water chemistry parameters on microbial decomposition. However, we found no effect of space (i.e., site location within the network) on ecosystem function, contrary to expectations. We also found no imprint of the microbial community richness assessed in the water column (α–diversity and Shannon Index) on the patterns of organic–matter decomposition, however, this may be due to our focus on potential colonising microbes rather than benthic communities. Our study highlights the need to include functional indicators for the assessment of ecosystems.

Diet Affects Egg Laying, Biomass, and Stable Isotope Values in Tetragnathid Spiders.

Riparian tetragnathid spiders are used as biosentinels of aquatic contamination because they are specialized feeders of aquatic emergent insects and are also prey items for terrestrial predators (e.g., birds). Analysis of both trophic position (e.g., stable nitrogen isotopes) and contaminant concentrations are needed to utilize tetragnathids as biosentinels, which can present challenges when collecting enough biomass to reach analytical detection limits, due to their relatively small size. The purpose of this study was to investigate the impacts of a controlled diet source on spider biomass, egg laying and stable isotope values (δ13C and δ15N). Diet significantly influenced the biomass and egg laying of tetragnathids, with some spiders losing up to 50% of their biomass in a single egg-laying event. δ13C had a faster turnover rate in the whole-body of spiders compared to legs, which is important, as spider legs are presently used as surrogates for whole-body δ13C values.

Fate of Planktothrix-derived toxins in aquatic food webs: A case study in Lake Mindelsee (Germany)

Blooms of the red, filamentous cyanobacterium Planktothrix rubescens occur frequently in pre-alpine lakes in Europe, often with concomitant toxic microcystin (MC) production. Trophic transfer of MCs has been observed in bivalves, fish, and zooplankton species, while uptake of MCs into Diptera species could facilitate distribution of MCs into terrestrial food webs and habitats. In this study, we characterized a Planktothrix bloom in summer 2019 in Lake Mindelsee and tracked possible trophic transfer and/or bioaccumulation of MCs via analysis of phytoplankton, zooplankton (Daphnia) and emergent aquatic insects (Chaoborus, Chironomidae and Trichoptera). Using 16 S rRNA gene amplicon sequencing, we found that five sequence variants of Planktothrix spp. were responsible for bloom formation in September and October of 2019, and these MC-producing variants, provisionally identified as P. isothrix and/or P. serta, occurred exclusively in Lake Mindelsee (Germany), while other variants were also detected in nearby Lake Constance. The remaining cyanobacterial community was dominated by Cyanobiaceae species with high species overlap with Lake Constance, suggesting a well-established exchange of cyanobacteria species between the adjacent lakes. With targeted LC–HRMS/MS we identified two MC-congeners, MC-LR and [Asp3]MC-RR with maximum concentrations of 45 ng [Asp3]MC-RR/L in lake water in September. Both MC congeners displayed different predominance patterns, suggesting that two different MC-producing species occurred in a time-dependent manner, whereby [Asp3]MC-RR was clearly associated with the Planktothrix spp. bloom. We demonstrate an exclusive transfer of MC-LR, but not [Asp3]MC-RR, from phytoplankton into zooplankton reaching a 10-fold bioconcentration, yet complete absence of these MC congeners or their conjugates in aquatic insects. The latter demonstrated a limited trophic transfer of MCs from zooplankton to zooplanktivorous insect larvae (e.g., Chaoborus), or direct transfer into other aquatic insects (e.g. Chironomidae and Trichoptera), whether due to avoidance or limited uptake and/or rapid excretion of MCs by higher trophic emergent aquatic insects.

Land use alters cross-ecosystem transfer of high value fatty acids by aquatic insects

BackgroundMany aquatic insects emerge as adults from water bodies to complete parts of their life cycle in terrestrial ecosystems and are potential prey for riparian predators. The benefits of riparian predators from aquatic insects include higher contents of long-chain polyunsaturated fatty acids (PUFA) compared to terrestrial insects. Aquatic insects are therefore considered a high-quality food. Food containing high levels of PUFA can enhance growth and immune response of spiders. However, agricultural stressors like nutrient increase, pesticides and habitat degradation can affect the biomass of aquatic insects and in turn the diet of spiders. Studies quantifying the influence of land use on fatty acid (FA) profiles of emergent aquatic insects and riparian predators are lacking. We quantified differences in exports of FA, saturated FA, monounsaturated FA, and PUFA, FA profiles of aquatic insects and spiders between forested and agricultural sites over the primary emergence period within one year. The FA export to the riparian food web is crucial to understand energy fluxes between ecosystems. Furthermore, we monitored environmental variables to identify associations between agricultural stressors and FA profiles of aquatic insects and spiders.ResultsWe found differences in FA export and profiles of aquatic insects between land-use types. The quantity of total FA export via aquatic insects was lower in agricultural sites (95% CI 1147–1313 µg m−2) in comparison to forested sites (95% CI 1555–1845 µg m−2), while the biomass export was higher in agricultural sites. Additionally, in spring the PUFA export was significantly lower (up to 0.06 µg d−1 m−2) in agricultural than forested sites. Agricultural stressors explained only little variation in the FA profiles of aquatic insects, e.g., 4% for caddisflies and 12% for non-biting midges. Percentage of shading and pool habitats were identified as most important variables explaining the variation in FA profiles.ConclusionThe quality of aquatic insects as food source for riparian spiders was smaller in agricultural than forested sites, which can decrease the fitness of riparian predators. To improve our capacity to predict potential adverse effects in the riparian food web, future studies should identify the mechanisms underlying a lower PUFA content.

The influence of season, hunting mode, and habitat specialization on riparian spiders as key predators in the aquatic-terrestrial linkage

Freshwater ecosystems subsidize riparian zones with high-quality nutrients via the emergence of aquatic insects. Spiders are dominant consumers of these insect subsidies. However, little is known about the variation of aquatic insect consumption across spiders of different hunting modes, habitat specializations, seasons, and systems. To explore this, we assembled a large stable isotope dataset (n > 1000) of aquatic versus terrestrial sources and six spider species over four points in time adjacent to a lotic and a lentic system. The spiders represent three hunting modes each consisting of a wetland specialist and a habitat generalist. We expected that specialists would feed more on aquatic prey than their generalist counterparts. Mixing models showed that spiders’ diet consisted of 17–99% of aquatic sources, with no clear effect of habitat specialization. Averaged over the whole study period, web builders (WB) showed the highest proportions (78%) followed by ground hunters (GH, 42%) and vegetation hunters (VH, 31%). Consumption of aquatic prey was highest in June and August, which is most pronounced in GH and WBs, with the latter feeding almost entirely on aquatic sources during this period. Additionally, the elevated importance of high-quality lipids from aquatic origin during fall is indicated by elemental analyses pointing to an accumulation of lipids in October, which represent critical energy reserves during winter. Consequently, this study underlines the importance of aquatic prey irrespective of the habitat specialization of spiders. Furthermore, it suggests that energy flows vary substantially between spider hunting modes and seasons.

Spectral composition of light-emitting diodes impacts aquatic and terrestrial invertebrate communities with potential implications for cross-ecosystem subsidies.

Resource exchanges in the form of invertebrate fluxes are a key component of aquatic-terrestrial habitat coupling, but this interface is susceptible to human activities, including the imposition of artificial light at night. To better understand the effects of spectral composition of light-emitting diodes (LEDs)-a technology that is rapidly supplanting other lighting types-on emergent aquatic insects and terrestrial insects, we experimentally added LED fixtures that emit different light spectra to the littoral zone and adjacent riparian habitat of a pond. We installed four replicate LED treatments of different wavelengths (410, 530 and 630 nm), neutral white (4000 k) and a dark control, and sampled invertebrates in both terrestrial and over-water littoral traps. Invertebrate communities differed among light treatments and between habitats, as did total insect biomass and mean individual insect size. Proportional allochthonous biomass was greater in the riparian habitat and among some light treatments, demonstrating an asymmetrical effect of differently coloured LEDs on aquatic-terrestrial resource exchanges. Overall, our findings demonstrate that variation in wavelength from LEDs may impact the flux of resources between systems, as well as the communities of insects that are attracted to particular spectra of LED lighting, with probable implications for consumers. This article is part of the theme issue 'Light pollution in complex ecological systems'.

Increased bat hunting at polluted streams suggests chemical exposure rather than prey shortage

Streams and their riparian areas are important habitats and foraging sites for bats feeding on emergent aquatic insects. Chemical pollutants entering freshwater streams from agricultural and wastewater sources have been shown to alter aquatic insect emergence, yet little is known about how this impacts insectivorous bats in riparian areas. In this study, we investigate the relationships between the presence of wastewater effluent, in-stream pesticide toxicity, the number of emergent and flying aquatic insects, and the activity and hunting behaviour of bats at 14 streams in southwestern Germany. Stream sites were located in riparian forests, sheltered from direct exposure to pollutants from agricultural and urban areas. We focused on three bat species associated with riparian areas: Myotis daubentonii, M. cf. brandtii, and Pipistrellus pipistrellus. We found that streams with higher pesticide toxicity and more frequent detection of wastewater also tended to be warmer and have higher nutrient and lower oxygen concentrations. We did not observe a reduction of insect emergence, bat activity or hunting rates in association with pesticide toxicity and wastewater detections. Instead, the activity and hunting rates of Myotis spp. were higher at more polluted sites. The observed increase in bat hunting at more polluted streams suggests that instead of reduced prey availability, chemical pollution at the levels measured in the present study could expose bats to pollutants transported from the stream by emergent aquatic insects.

DNA metabarcoding reveals rangewide variation in aquatic diet of a riparian avian insectivore, the Prothonotary Warbler

Abstract Riparian avian insectivores not only depend on terrestrial insect prey but also benefit from the inclusion of aquatic prey during critical life-history periods. Diets identified herein show that Prothonotary Warbler (Protonotaria citrea) nestlings were provisioned with aquatic prey throughout the breeding season across their range, but with variation in prey frequency of occurrence and taxonomy. Anthropogenic activity and climate change may impact the trophic link especially between aquatic and riparian habitats by altering the presence, abundance, and timing of prey availability. Thus, we used DNA metabarcoding of fecal samples to quantify the frequency of occurrence of nestling diet items at 9 sites across their breeding range that differed in expected aquatic prey consumption. We analyzed spatial and temporal differences in the occurrence and multivariate diet assemblages of each prey source. Lepidoptera was the predominant terrestrial prey occurring in diets across space and time, whereas emergent aquatic insects and freshwater mollusks in aquatic diet exhibited greater variation. The frequency of emergent aquatic prey occurrence in nestling diets ranged from 61% to 100% across sites and was greater for early-season nestlings. The seasonal decrease in aquatic prey consumption indicates a potential temporal shift in the nutritional landscape from aquatic to terrestrial prey sources and a possible nutritional phenological mismatch for early nestlings as climate change advances the timing of insect emergence. Our findings also suggest that Prothonotary Warblers respond to environmental variability by consuming alternative prey and argue for future research investigating the extent to which shifting diets have nutritional consequences for riparian nestlings.

Enhanced Mercury Accumulation in Riparian Spiders: An Evidence of Insects' Emergence Effect in Aquatic and Upland Terrestrial Crossed Habitat.

Many studies have focused on mercury (Hg) accumulation in both aquatic and terrestrial organisms, but the effects of aquatic Hg on terrestrial organisms have rarely been documented. Here we report the accumulation of Hg in two species of spiders, Argiope bruennichi, inhabiting paddy fields, and Nephila clavata, inhabiting small forests in the riparian zones of two hydroelectric reservoirs in Guiyang, southwest China. The mean concentration of total mercury (THg) was higher in N. clavata (0.38mgkg-1) than in A. bruennichi (0.20mgkg-1). The monthly average THg in N. clavata, collected consecutively from May to October, and the highest values for THg in June (1.2mgkg-1) could be related to the emergence of aquatic insects during early summer, suggesting that emerging insects play a crucial role in the accumulation of Hg in riparian spiders. The high values could also be attributable to the different times of spider sampling or individual differences.

Chronic Engineered Nanoparticle Additions Alter Insect Emergence and Result in Metal Flux from Aquatic Ecosystems into Riparian Food Webs.

Freshwater ecosystems are exposed to engineered nanoparticles (NPs) through discharge from wastewater and agricultural runoff. We conducted a 9-month mesocosm experiment to examine the combined effects of chronic NP additions on insect emergence and insect-mediated contaminant flux to riparian spiders. Two NPs (copper, gold, plus controls) were crossed by two levels of nutrients in 18 outdoor mesocosms open to natural insect and spider colonization. We collected adult insects and two riparian spider genera, Tetragnatha and Dolomedes, for 1 week on a monthly basis. We estimated a significant decrease in cumulative insect emergence of 19% and 24% after exposure to copper and gold NPs, irrespective of nutrient level. NP treatments led to elevated copper and gold tissue concentrations in adult insects, which resulted in terrestrial fluxes of metals. These metal fluxes were associated with increased gold and copper tissue concentrations for both spider genera. We also observed about 25% fewer spiders in the NP mesocosms, likely due to reduced insect emergence and/or NP toxicity. These results demonstrate the transfer of NPs from aquatic to terrestrial ecosystems via emergence of aquatic insects and predation by riparian spiders, as well as significant reductions in insect and spider abundance in response to NP additions.

Extreme climatic events alter the aquatic insect community in a pristine German stream

As a result of ongoing climate change, extreme climatic events (ECEs) are expected to become more frequent and severe. The high biodiversity of riverine ecosystems is susceptible to ECEs, especially to water temperature (extreme heat and extreme cold) and discharge-related (flood and drought) events. Long time series are needed to unravel the effects of ECEs on ecological communities. Here, we used 20 years (1986–2005) of unusually high-resolution data from a pristine first-order stream in Germany. Daily recordings of species-level identified aquatic insect (Ephemeroptera, Plecoptera, Trichoptera: EPT) emergence, water temperature and discharge data were used to examine the effects of four types of ECEs (extreme heat, extreme cold, flood, and drought events) on insect abundance, common taxonomic diversity metrics, and selected traits after five different time lags (2 weeks, 1, 3, 6, and 12 months). Extreme heat events increased from 1.8 ± 1.9 SE events per year before 2000 to 5.3 ± 1.9 SE events per year after 2000. Water temperature-related ECEs restructured the EPT community in abundance, species richness, and traits (community temperature index: CTI, and dispersal capacity metric: DCM). The strongest effects on the EPT community were found when it was exposed to multiple ECEs and 1 and 3 months after an ECE. The changing frequencies and durations of ECEs, especially the increasing frequency of extreme heat events and the negative cumulative effects of ECEs, paint a worrisome picture for the future of EPT communities in headwater streams. High-resolution, long-term data across sites is needed to further disentangle the effects of different ECE stressors.

Latitudinal patterns of aquatic insect emergence driven by climate

AbstractAimEmerging aquatic insects link aquatic and terrestrial ecosystems across the Earth. Their diversity, abundance and functional importance means their emergence is an important phenological event. Nevertheless, aquatic insect emergence is understudied at a global scale compared to other phenological events, despite changing phenology being one of the most significant ecological responses to climate change. Here, we quantitatively describe the global patterns, and key proposed drivers, of seasonal aquatic insect emergence, to further understand how these patterns might change in the future.LocationGlobal.Time Period1950–2018.Major Taxa StudiedEmerging aquatic insects.MethodsWe extracted monthly emergence data from 86 studies across 163 sites to construct 1053 annual emergence curves. We parameterized the curves using two complementary metrics of seasonality, which were modelled against geographical and climatic variables to determine the direct and indirect relationships between them.ResultsWe found clear global trends in aquatic insect emergence patterns across latitude and underlying climates. Between‐month variation and temporal restriction of emergence increased from the equator to the poles, going from small, aseasonal fluctuations in the warm, thermally stable tropics to large, seasonal peaks at cooler, thermally unstable higher latitudes. While emergence trends were associated with gradients of precipitation, temperature was the dominant climatic driver of the latitudinal trend.Main ConclusionsThese findings suggest that with climate warming, aquatic insects will emerge over longer periods, diluted in abundances and displaying less seasonal emergence patterns with smaller between‐month fluctuations. This may result in disruption of ecosystem functions seasonally dependent on aquatic insects, such as riparian predation, pollination and disease transmission. The cross‐ecosystem life cycle of aquatic insects means changes to their seasonal patterns of emergence will have impacts in both aquatic and terrestrial ecosystems.

Subsidy Quality Affects Common Riparian Web-Building Spiders: Consequences of Aquatic Contamination and Food Resource.

Anthropogenic stressors can affect the emergence of aquatic insects. These insects link aquatic and adjacent terrestrial food webs, serving as high-quality subsidy to terrestrial consumers, such as spiders. While previous studies have demonstrated that changes in the emergence biomass and timing may propagate across ecosystem boundaries, the physiological consequences of altered subsidy quality for spiders are largely unknown. We used a model food chain to study the potential effects of subsidy quality: Tetragnatha spp. were exclusively fed with emergent Chironomus riparius cultured in the absence or presence of either copper (Cu), Bacillus thuringiensis var. israelensis (Bti), or a mixture of synthetic pesticides paired with two basal resources (Spirulina vs. TetraMin®) of differing quality in terms of fatty acid (FA) composition. Basal resources shaped the FA profile of chironomids, whereas their effect on the FA profile of spiders decreased, presumably due to the capacity of both chironomids and spiders to modify (dietary) FA. In contrast, aquatic contaminants had negligible effects on prey FA profiles but reduced the content of physiologically important polyunsaturated FAs, such as 20:4n-6 (arachidonic acid) and 20:5n-3 (eicosapentaenoic acid), in spiders by approximately 30% in Cu and Bti treatments. This may have contributed to the statistically significant decline (40%-50%) in spider growth. The observed effects in spiders are likely related to prey nutritional quality because biomass consumption by spiders was, because of our experimental design, constant. Analyses of additional parameters that describe the nutritional quality for consumers such as proteins, carbohydrates, and the retention of contaminants may shed further light on the underlying mechanisms. Our results highlight that aquatic contaminants can affect the physiology of riparian spiders, likely by altering subsidy quality, with potential implications for terrestrial food webs. Environ Toxicol Chem 2023;42:1346-1358. © 2023 The Authors. Environmental Toxicology and Chemistry published by Wiley Periodicals LLC on behalf of SETAC.

Emergence phenology of the giant salmonfly and responses by birds in Idaho river networks

Emergence of adult aquatic insects from rivers is strongly influenced by water temperature, and emergence timing helps to determine the availability of this ephemeral food resource for birds and other terrestrial insectivores. It is poorly understood how spatial heterogeneity in riverine habitat mediates the timing of emergence. Such spatiotemporal variation may have consequences for terrestrial insectivores that rely on aquatic-derived prey resources. We investigated emergence phenology of the giant salmonfly, Pteronarcys californica, at three spatial scales in two Idaho river networks. We examined the influence of tributary confluences on salmonfly emergence timing and associated insectivorous bird responses. Salmonfly emergence timing was highly variable at the basin-scale during the period we sampled (May–June). Within sub-drainage pathways not punctuated by major tributaries, emergence followed a downstream-to-upstream pattern. At the scale of reaches, abrupt changes in thermal regimes created by 10 major tributary confluences created asynchrony in emergence of 1–6 days among the 20 reaches bracketing the confluences. We observed 10 bird species capturing emerged salmonflies, including 5 species typically associated with upland habitats (e.g., American robin, red-tailed hawk, American kestrel) but that likely aggregated along rivers to take advantage of emerging salmonflies. Some birds (e.g., Lewis’s woodpecker, western tanager, American dipper) captured large numbers of salmonflies, and some of these fed salmonflies to nestlings. Emergence asynchrony created by tributaries was associated with shifts in bird abundance and richness which both nearly doubled, on average, during salmonfly emergence. Thermal heterogeneity in river networks created asynchrony in aquatic insect phenology which prolonged the availability of this pulsed prey resource for insectivorous birds during key breeding times. Such interactions between spatial and temporal heterogeneity and organism phenology may be critical to understanding the consequences of fluxes of resources that link water and land. Shifts in phenology or curtailment of life history diversity in organisms like salmonflies may have implications for these organisms, but could also contribute to mismatches or constrain availability of pulsed resources to dependent consumers. These could be unforeseen consequences, for both aquatic and terrestrial organisms, of human-driven alteration and homogenization of riverscapes.

Cascading impacts of changes in subsidy quality on recipient ecosystem functioning.

Resource quantity and quality can differ between adjacent ecosystems, and these differences can impact subsidies exchanged between ecosystems. The quantity and quality of subsidies are rapidly changing in response to stressors associated with global environmental change, but while we have models to predict the effects of changes in subsidy quantity, we currently lack models to predict the effects of changes in subsidy quality on recipient ecosystem functioning. We developed a novel model to predict the effects of subsidy quality on recipient ecosystem biomass distribution, recycling, production, and efficiency. We parameterized the model for a case study of a riparian ecosystem subsidized by pulsed emergent aquatic insects. In this case study we focused on a common measure of subsidy quality that differs between riparian and aquatic ecosystems: the higher content of long-chain polyunsaturated fatty acids (PUFAs) in aquatic ecosystems. We analyzed how changes in the PUFA concentration of aquatic subsidies affect the dynamics in biomass stocks and functions of the riparian ecosystem. We also conducted a global sensitivity analysis to identify key drivers of subsidy impacts. Our analysis showed that subsidy quality increased the functioning of the recipient ecosystem. Recycling increased more strongly than production per unit subsidy quality increase, meaning there was a threshold where an increase in subsidy quality led to stronger effects of subsidies on recycling relative to the production of the recipient ecosystem. Our predictions were most sensitive to basal nutrient input, highlighting the relevance of recipient ecosystem nutrient levels to understanding the effects of ecosystem connections. We argue that recipient ecosystems that rely on high-quality subsidies, such as aquatic-terrestrial ecotones, are highly sensitive to changes in subsidy-recipient ecosystem connections. Our novel model unifies the subsidy hypothesis and food quality hypothesis and provides testable predictions to understand the effects of ecosystem connections on ecosystem functioning under global changes.

Trophic transfer of polyunsaturated fatty acids across the aquatic-terrestrial interface: An experimental tritrophic food chain approach.

Aquatic and their adjacent terrestrial ecosystems are linked via the flux of organic and inorganic matter. Emergent aquatic insects are recognized as high-quality food for terrestrial predators, because they provide more physiologically relevant long-chain polyunsaturated fatty acids (PUFA) than terrestrial insects. The effects of dietary PUFA on terrestrial predators have been explored mainly in feeding trials conducted under controlled laboratory conditions, hampering the assessment of the ecological relevance of dietary PUFA deficiencies under field conditions. We assessed the PUFA transfer across the aquatic-terrestrial interface and the consequences for terrestrial riparian predators in two outdoor microcosm experiments. We established simplified tritrophic food chains, consisting of one of four basic food sources, an intermediary collector gatherer (Chironomus riparius, Chironomidae), and a riparian web-building spider (Tetragnatha sp.). The four basic food sources (algae, conditioned leaves, oatmeal, and fish food) differed in PUFA profiles and were used to track the trophic transfer of single PUFA along the food chain and to assess their potential effects on spiders, that is, on fresh weight, body condition (size-controlled measurement of nutritional status), and immune response. The PUFA profiles of the basic food sources, C.riparius and spiders differed between treatments, except for spiders in the second experiment. The PUFA α-linolenic acid (ALA, 18:3n-3) and ɣ-linolenic acid (GLA, 18:3n-6) were major contributors to the differences between treatments. PUFA profiles of the basic food sources influenced the fresh weight and body condition of spiders in the first experiment, but not in the second experiment, and did not affect the immune response, growth rate, and dry weight in both experiments. Furthermore, our results indicate that the examined responses are dependent on temperature. Future studies including anthropogenic stressors would deepen our understanding of the transfer and role of PUFA in ecosystems.

Metal Exposure and Sex Shape the Fatty Acid Profile of Midges and Reduce the Aquatic Subsidy to Terrestrial Food Webs.

Aquatic micropollutants can be transported to terrestrial systems and their consumers by emergent aquatic insects. However, micropollutants, such as metals, may also affect the flux of physiologically important polyunsaturated fatty acids (PUFAs). As certain PUFAs have been linked to physiological fitness and breeding success of terrestrial consumers, reduced fluxes from aquatic systems could affect terrestrial populations and food webs. We chronically exposed larvae of the aquatic insect Chironomus riparius to a range of environmentally relevant sediment contents of cadmium (Cd) or copper (Cu) in a 28-day microcosm study. Since elevated water temperatures can enhance metals' toxic effects, we used two temperature regimes, control and periodically elevated temperatures (heat waves) reflecting an aspect of climate change. Cd and Cu significantly reduced adult emergence by up to 95% and 45%, respectively, while elevated temperatures had negligible effects. Both metal contents were strongly reduced (∼90%) during metamorphosis. Furthermore, the chironomid FA profile was significantly altered during metamorphosis with the factors sex and metal exposure being relevant predictors. Consequently, fluxes of physiologically important PUFAs by emergent adults were reduced by up to ∼80%. Our results suggest that considering fluxes of physiologically important compounds, such as PUFAs, by emergent aquatic insects is important to understand the implications of aquatic micropollutants on aquatic-terrestrial meta-ecosystems.

Body stores of emergent aquatic insects are associated with body size, sex, swarming behaviour and dispersal strategies

Abstract Dispersal is a determining step in the life cycle of insects and a key factor of their energy budget. If the body stores of terrestrial insects are relatively well‐documented, those of merolimnic insects (i.e., with aquatic larvae and terrestrial winged adults) remain poorly investigated. We assayed the body stores (protein, carbohydrate and lipid contents) of 18 families of merolimnic insects, encompassing a wide diversity of biological and ecological traits. We highlighted allometry between body stores and mass. The proportion of triglyceride and free carbohydrate relatively decreases with an increase in absolute insect mass, whereas protein and glycogen contents are proportional. We found a significant effect of insect order, Diptera having relatively more proteins than Trichoptera. For swarming taxa, males have relatively more triglycerides than their conspecific female. Passive dispersers are characterised by higher protein and glycogen contents Each body store was associated with specific life‐history traits, emphasising the importance of considering each body store independently to better understand the associated functions. Dispersal strategies seem to be the main structuring factor underlying body store patterns of merolimnic insects, being significantly associated with the variation in adjusted triglycerides (30%), proteins (21%), glycogen (29%) and free carbohydrates (17%). However, our study estimated the role of flight only indirectly through the biological traits of insects. Future works should focus on identifying which substrates are used for flight, by comparing the proportion of body stores before and after flight, as well as protein identification to distinguish proteins associated with flight or reproduction. These further studies will help researchers in understanding the link between body stores of emerging merolimnic insects and the various facets of their ecology.