The evolutionary ecology of fatty-acid variation: Implications for consumer adaptation and diversification.

The nutritional diversity of resources can affect the adaptive evolution of consumer metabolism and consumer diversification. The omega-3 long-chain polyunsaturated fatty acids eicosapentaenoic acid (EPA; 20:5n-3) and docosahexaenoic acid (DHA; 22:6n-3) have a high potential to affect consumer fitness, through their widespread effects on reproduction, growth and survival. However, few studies consider the evolution of fatty acid metabolism within an ecological context. In this review, we first document the extensive diversity in both primary producer and consumer fatty acid distributions amongst major ecosystems, between habitats and amongst species within habitats. We highlight some of the key nutritional contrasts that can shape behavioural and/or metabolic adaptation in consumers, discussing how consumers can evolve in response to the spatial, seasonal and community-level variation of resource quality. We propose a hierarchical trait-based approach for studying the evolution of consumers' metabolic networks and review the evolutionary genetic mechanisms underpinning consumer adaptation to EPA and DHA distributions. In doing so, we consider how the metabolic traits of consumers are hierarchically structured, from cell membrane function to maternal investment, and have strongly environment-dependent expression. Finally, we conclude with an outlook on how studying the metabolic adaptation of consumers within the context of nutritional landscapes can open up new opportunities for understanding evolutionary diversification.

The effects of sediment traps on instream habitat and macroinvertebrates of mountain streams.

Sediment transport in mountain streams can be a major hazard to downstream infrastructure. Consequently, sediment traps are a common feature in many high gradient streams to retain large volumes of sediment and protect settlements from major sediment transport events. Despite the wide application of these instream structures, there is little knowledge regarding the environmental and ecological effects on streams. Here, we investigated the hydromorphological effects of sediment traps on instream habitats and associated macroinvertebrate communities in four impacted and three non-impacted streams in Switzerland. Sediment traps significantly disrupted the sediment regime homogenising grain size percentiles compared to the natural stepwise downstream fining in control streams. This disruption in the sediment regime resulted in finer grain size distributions upstream of the sediment trap, and reduced substrate diversity in the sediment retention basin and just downstream of the trap. The reductions in substrate diversity resulted in an altered macroinvertebrate community composition. Further, the disconnection in sediment transport led to a lack of longitudinal correlation in macroinvertebrate communities. Refugia provision downstream of the sediment trap, and resource availability within the retention basin, were diminished, potentially reducing resilience of macroinvertebrate assemblages to instream disturbances. The effects of sediment traps were most likely localised in three of the four streams with substrate diversity recovering to comparable control values within 8 wetted widths (ca. 50 m) downstream of the trap associated with natural longitudinal fining. In contrast, ecological and environmental effects propagated downstream in one impacted stream with no recovery being evident. Sediment retention basins in the impacted streams provided a local artificially unique habitat of dynamic-braided channels. Our results indicate that sediment traps can significantly disrupt the sediment regime with important consequences for instream ecology and environmental conditions, although these effects can be system specific. Further work is needed to fully understand the effects of sediment traps in mountain streams to assist resource managers in the mitigation and future construction of these structures.

Stream degradation affects aquatic resource subsidies to riparian ground-dwelling spiders.

Freshwater systems have undergone drastic alterations during the last century, potentially affecting cross-boundary resource transfers between aquatic and terrestrial ecosystems. One important connection is the export of biomass by emergent aquatic insects containing omega-3 polyunsaturated fatty acids (PUFAs), especially eicosapentaenoic acid (EPA), that is scarce in terrestrial systems. Because of taxon-specific differences in PUFA content and functional traits, the contribution of different insect groups should be considered, in addition to total biomass export. In this context, one important trait is the emergence mode. Stoneflies, in contrast to other aquatic insects, crawl to land to emerge instead of flying directly from the water surface, making them accessible to ground-dwelling predators. Because stoneflies are especially susceptible to environmental change, stream degradation might cause a mismatch of available and required nutrients, particularly for ground-dwelling predators. In this study, we estimated emergent biomass and EPA export along two streams with different levels of habitat degradation. The EPA content in aquatic insects did not differ with different degrees of habitat degradation and total biomass export in spring was with 7.9 ± 9.6 mg m-2 day-1 in the degraded and 7.3 ± 8.5 mg m-2 day-1 in the natural system, also unaffected. However, habitat degradation substantially altered the contribution of crawling emergence to the total export in spring, with no biomass export by stoneflies at the most degraded sites. The EPA content in ground-dwelling spiders was correlated with emergent stonefly biomass, making up only 16.0 ± 6.2 % of total fatty acids at sites with no stonefly emergence, but 27.3 ± 3.0 % at sites with highest stonefly emergence. Because immune function in ground-dwelling spiders has been connected to EPA levels, reduced crawling emergence might impact spider fitness. Functional traits, like emergence mode as well as nutritional quality, should be considered when assessing the effects of stream degradation on adjacent terrestrial ecosystems.

Resilience to changes in lake trophic state: Nutrient allocation into Daphnia resting eggs.

During past decades, many lakes underwent drastic human-caused changes in trophic state with strong implications for population dynamics and food web processes. We investigated the influence of trophic state on nutrient allocation into Daphnia resting eggs. The production of resting eggs is an important survival strategy, allowing Daphnia to cope with unfavorable environmental conditions. Allocation of essential nutrients into resting eggs may crucially influence embryonic development and offspring survival and thus is of great ecological and evolutionary interest. The capacity of Daphnia to adjust the allocation of nutrients into resting eggs may depend on the dietary nutrient supply, which may vary with trophic state-related changes in the phytoplankton community composition. Resting eggs were isolated from sediment cores taken from Lake Constance, a large prealpine lake with a distinct eutrophication and reoligotrophication history, and analyzed for elemental (carbon, nitrogen, and phosphorus) and biochemical (sterols and fatty acids) nutrients. Carbon allocation into Daphnia resting eggs continuously decreased over time, irrespective of changes in trophic state. The allocation of nitrogen into Daphnia resting eggs followed the changes in trophic state, that is, nitrogen concentrations in resting eggs increased with eutrophication and decreased again with reoligotrophication. The allocation of phosphorus, sterols and long-chain polyunsaturated fatty acids, such as eicosapentaenoic acid, into Daphnia resting eggs did not change significantly over time. Changes in trophic state strikingly influenced all trophic levels in Lake Constance. However, nutrient allocation into Daphnia resting eggs was mostly resilient to changes in lake trophic state.

Cross-ecosystem fluxes: Export of polyunsaturated fatty acids from aquatic to terrestrial ecosystems via emerging insects.

Cross-ecosystem fluxes can crucially influence the productivity of adjacent habitats. Emerging aquatic insects represent one important pathway through which freshwater-derived organic matter can enter terrestrial food webs. Aquatic insects may be of superior food quality for terrestrial consumers because they contain high concentrations of essential polyunsaturated fatty acids (PUFA). We quantified the export of PUFA via emerging insects from a midsize, mesotrophic lake. Insects were collected using emergence traps installed above different water depths and subjected to fatty acid analyses. Insect emergence from different depth zones and seasonal mean fatty acid concentrations in different insect groups were used to estimate PUFA fluxes. In total, 80.5mg PUFA m-2yr-1 were exported, of which 32.8mgm-2yr-1 were eicosapentaenoic acid (EPA), 7.8mgm-2yr-1 were arachidonic acid (ARA), and 2.6mgm-2yr-1 were docosahexaenoic acid (DHA). While Chironomidae contributed most to insect biomass and total PUFA export, Chaoborus flavicans contributed most to the export of EPA, ARA, and especially DHA. The export of total insect biomass from one square meter declined with depth and the timing at which 50% of total insect biomass emerged was correlated with the water depths over which the traps were installed, suggesting that insect-mediated PUFA fluxes are strongly affected by lake morphometry. Applying a conceptual model developed to assess insect deposition rates on land to our insect-mediated PUFA export data revealed an average total PUFA deposition rate of 150mgm-2yr-1 within 100m inland from the shore. We propose that PUFA export can be reliably estimated using taxon-specific information on emergent insect biomass and seasonal mean body PUFA concentrations of adult insects provided here. Our data indicate that insect-mediated PUFA fluxes from lakes are substantial, implying that freshwater-derived PUFA can crucially influence food web processes in adjacent terrestrial habitats.