Networking nutrients: How nutrition determines the structure of ecological networks.

Nutrients can shape ecological interactions but remain poorly integrated into ecological networks. Concepts such as nutrient-specific foraging nevertheless have the potential to expose the mechanisms structuring complex ecological systems. Nutrients also present an opportunity to predict dynamic processes, such as interaction rewiring and extinction cascades, and increase the accuracy of network analyses. Here, we propose the concept of nutritional networks. By integrating nutritional data into ecological networks, we envisage significant advances to our understanding of ecological processes from individual to ecosystem scales. We show that networks can be constructed with nutritional data to illuminate how nutrients structure ecological interactions in natural systems through an empirical example. Throughout, we identify fundamental ecological hypotheses that can be explored in a nutritional network context, alongside methods for resolving those networks. Nutrients influence the structure and complexity of ecological networks through mechanistic processes and concepts including nutritional niche differentiation, functional responses, landscape diversity, ecological invasions and ecosystem robustness. Future research on ecological networks should consider nutrients when investigating the drivers of network structure and function.

Understanding trophic interactions in a warming world by bridging foraging ecology and biomechanics with network science.

Climate change will disrupt biological processes at every scale. Ecosystem functions and services vital to ecological resilience are set to shift, with consequences for how we manage land, natural resources, and food systems. Increasing temperatures cause morphological shifts, with concomitant implications for biomechanical performance metrics crucial to trophic interactions. Biomechanical performance, such as maximum bite force or running speed, determines the breadth of resources accessible to consumers, the outcome of interspecific interactions, and thus the structure of ecological networks. Climate change-induced impacts to ecosystem services and resilience are therefore on the horizon, mediated by disruption of biomechanical performance and, consequently, trophic interactions across whole ecosystems. Here, we argue that there is an urgent need to investigate the complex interactions between climate change, biomechanical traits and foraging ecology to help predict changes to ecological networks and ecosystem functioning. We discuss how these seemingly disparate disciplines can be connected through network science. Using an ant-plant network as an example, we illustrate how different data types could be integrated to investigate the interaction between warming, bite force and trophic interactions, and discuss what such an integration will achieve. It is our hope that this integrative framework will help to identify a viable means to elucidate previously intractable impacts of climate change, with effective predictive potential to guide management and mitigation.

Extra terrestrials: drought creates niche space for rare invertebrates in a large-scale and long-term field experiment.

Freshwater habitats are drying more frequently and for longer under the combined pressures of climate change and overabstraction. Unsurprisingly, many aquatic species decline or become locally extinct as their benthic habitat is lost during stream droughts, but less is known about the potential 'winners': those terrestrial species that may exploit emerging niches in drying riverbeds. In particular, we do not know how these transient ecotones will respond as droughts become more extreme in the future. To find out we used a large-scale, long-term mesocosm experiment spanning a wide gradient of drought intensity, from permanent flows to full streambed dewatering, and analysed terrestrial invertebrate community assembly after 1 year. Droughts that caused stream fragmentation gave rise to the most diverse terrestrial invertebrate assemblages, including 10 species with UK conservation designations, and high species turnover between experimental channels. Droughts that caused streambed dewatering produced lower terrestrial invertebrate richness, suggesting that the persistence of instream pools may benefit these taxa as well as aquatic biota. Particularly intense droughts may therefore yield relatively few 'winners' among either aquatic or terrestrial species, indicating that the threat to riverine biodiversity from future drought intensification could be more pervasive than widely acknowledged.

Long-term changes in macroinvertebrate communities across high-latitude streams.

Long-term records of benthic macroinvertebrates in high-latitude streams are essential for understanding climatic changes, including extreme events (e.g. floods). Data extending over multiple decades are typically scarce. Here, we investigated macroinvertebrate community structural change (including alpha and beta diversity and gain and loss of species) over 22 years (1994-2016) in 10 stream systems across Denali National Park (Alaska, USA) in relation to climatological and meteorological drivers (e.g. air temperature, snowpack depth, precipitation). We hypothesised that increases in air temperature and reduced snowpack depth, due to climatic change, would reduce beta and gamma diversity but increase alpha diversity. Findings showed temporal trends in alpha diversity were variable across streams, with oscillating patterns in many snowmelt- and rainfall runoff-fed streams linked to climatic variation (temperature and precipitation), but increased over time in several streams supported by a mixture of water sources, including more stable groundwater-fed streams. Beta-diversity over the time series was highly variable, yet marked transitions were observed in response to extreme snowpack accumulation (1999-2000), where species loss drove turnover. Gamma diversity did not significantly increase or decrease over time. Investigating trends in individual taxa, several taxa were lost and gained during a relative constrained time period (2000-2006), likely in response to climatic variability and significant shifts in instream environmental conditions. Findings demonstrate the importance of long-term biological studies in stream ecosystems and highlight the vulnerability of high-latitude streams to climate change.

Mismatch between IUCN range maps and species interactions data illustrated using the Serengeti food web.

Background: Range maps are a useful tool to describe the spatial distribution of species. However, they need to be used with caution, as they essentially represent a rough approximation of a species' suitable habitats. When stacked together, the resulting communities in each grid cell may not always be realistic, especially when species interactions are taken into account. Here we show the extent of the mismatch between range maps, provided by the International Union for Conservation of Nature (IUCN), and species interactions data. More precisely, we show that local networks built from those stacked range maps often yield unrealistic communities, where species of higher trophic levels are completely disconnected from primary producers. Methodology: We used the well-described Serengeti food web of mammals and plants as our case study, and identify areas of data mismatch within predators' range maps by taking into account food web structure. We then used occurrence data from the Global Biodiversity Information Facility (GBIF) to investigate where data is most lacking. Results: We found that most predator ranges comprised large areas without any overlapping distribution of their prey. However, many of these areas contained GBIF occurrences of the predator. Conclusions: Our results suggest that the mismatch between both data sources could be due either to the lack of information about ecological interactions or the geographical occurrence of prey. We finally discuss general guidelines to help identify defective data among distributions and interactions data, and we recommend this method as a valuable way to assess whether the occurrence data that are being used, even if incomplete, are ecologically accurate.

Influence of European Beech (Fagales: Fagaceae) Rot Hole Habitat Characteristics on Invertebrate Community Structure and Diversity.

Hollows of veteran trees (i.e., rot holes) provide habitat for many rare and threatened saproxylic invertebrates. Rot holes are highly heterogeneous, particularly in terms of substrate and microclimate conditions. There is, however, a dearth of information regarding the differences in biological communities inhabiting rot holes with different environmental conditions. Invertebrates were sampled from European beech (Fagus sylvatica) rot holes in Windsor, Savernake, and Epping Forests (United Kingdom). For each rot hole, physical and environmental conditions were measured, including tree diameter, rot hole dimensions, rot hole height, substrate density, water content, and water potential. These parameters were used to assess the influence of environmental conditions and habitat characteristics on invertebrate communities. Rot hole invertebrate communities were extremely diverse, containing both woodland generalist and saproxylic specialist taxa. Large variation in community structure was observed between rot holes and across woodlands; all sites supported threatened and endangered taxa. Environmental conditions in rot holes were highly variable within and between woodland sites, and communities were predominantly structured by these environmental conditions. In particular, turnover between invertebrate communities was linked to high ß-diversity. The linked heterogeneity of environmental conditions and invertebrate communities in rot holes suggests that management of deadwood habitats in woodlands should strive to generate environmental heterogeneity to promote invertebrate diversity. Additional research is required to define how management and conservation activities can further promote enhanced biodiversity across rot holes.

Global variation in freshwater physico-chemistry and its influence on chemical toxicity in aquatic wildlife.

Chemical pollution is one of the major threats to global freshwater biodiversity and will be exacerbated through changes in temperature and rainfall patterns, acid-base chemistry, and reduced freshwater availability due to climate change. In this review we show how physico-chemical features of natural fresh waters, including pH, temperature, oxygen, carbon dioxide, divalent cations, anions, carbonate alkalinity, salinity and dissolved organic matter, can affect the environmental risk to aquatic wildlife of pollutant chemicals. We evidence how these features of freshwater physico-chemistry directly and/or indirectly affect the solubility, speciation, bioavailability and uptake of chemicals [including via alterations in the trans-epithelial electric potential (TEP) across the gills or skin] as well as the internal physiology/biochemistry of the organisms, and hence ultimately toxicity. We also show how toxicity can vary with species and ontogeny. We use a new database of global freshwater chemistry (GLORICH) to demonstrate the huge variability (often >1000-fold) for these physico-chemical variables in natural fresh waters, and hence their importance to ecotoxicology. We emphasise that a better understanding of chemical toxicity and more accurate environmental risk assessment requires greater consideration of the natural water physico-chemistry in which the organisms we seek to protect live.

Macroinvertebrate communities in streams with contrasting water sources in the Japanese Alps.

Alpine streams are typically fed from a range of water sources including glacial meltwater, snowmelt, groundwater flow, and surface rainfall runoff. These contributions are projected to shift with climate change, particularly in the Japanese Alps where snow is expected to decrease, but rainfall events increase. The overarching aim of the study was to understand the key variables driving macroinvertebrate community composition in groundwater and snowmelt-fed streams (n = 6) in the Kamikochi region of the northern Japanese Alps (April-December 2017). Macroinvertebrate abundance, species richness, and diversity were not significantly different between the two stream types. Community structure, however, was different between groundwater and snowmelt-fed streams with macroinvertebrate taxa specialized for the environmental conditions present in each system. Temporal variation in the abundance, species richness, and diversity of macroinvertebrate communities was also significantly different between groundwater and snowmelt streams over the study period, with snowmelt streams exhibiting far higher levels of variation. Two snowmelt streams considered perennial proved to be intermittent with periodic drying of the streambed, but the macroinvertebrates in these systems rebounded rapidly after flows resumed with no reduction in taxonomic diversity. These same streams, nevertheless, showed a major reduction in diversity and abundance following periods of high flow, indicating floods rather than periodic drying was a major driver of community structure. This conclusion was also supported from functional analyses, which showed that the more variable snowmelt streams were characterized by taxa with resistant, rather than resilient, life-history traits. The findings demonstrate the potential for significant turnover in species composition with changing environmental conditions in Japanese alpine stream systems, with groundwater-fed streams potentially more resilient to future changes in comparison to snowmelt-fed streams.

Food web transfer of plastics to an apex riverine predator.

As a rapidly accelerating expression of global change, plastics now occur extensively in freshwater ecosystems, yet there is barely any evidence of their transfer through food webs. Following previous observations that plastics occur widely in their prey, we used a field study of free-living Eurasian dippers (Cinclus cinclus), to test the hypotheses that (1) plastics are transferred from prey to predators in rivers, (2) plastics contained in prey are transferred by adults to altricial offspring during provisioning and (3) plastic concentrations in faecal and regurgitated pellets from dippers increase with urbanization. Plastic occurred in 50% of regurgitates (n = 74) and 45% of faecal samples (n = 92) collected non-invasively from adult and nestling dippers at 15 sites across South Wales (UK). Over 95% of particles were fibres, and concentrations in samples increased with urban land cover. Fourier transform infrared spectroscopy identified multiple polymers, including polyester, polypropylene, polyvinyl chloride and vinyl chloride copolymers. Although characterized by uncertainty, steady-state models using energetic data along with plastic concentration in prey and excreta suggest that around 200 plastic particles are ingested daily by dippers, but also excreted at rates that suggest transitory throughput. As some of the first evidence revealing that plastic is now being transferred through freshwater food webs, and between adult passerines and their offspring, these data emphasize the need to appraise the potential ecotoxicological consequences of increasing plastic pollution.

Estimating the size distribution of plastics ingested by animals.

The ingestion of plastics appears to be widespread throughout the animal kingdom with risks to individuals, ecosystems and human health. Despite growing information on the location, abundance and size distribution of plastics in the environment, it cannot be assumed that any given animal will ingest all sizes of plastic encountered. Here, we use published data to develop an allometric relationship between plastic consumption and animal size to estimate the size distribution of plastics feasibly ingested by animals. Based on more than 2000 gut content analyses from animals ranging over three orders of magnitude in size (lengths 9 mm to 10 m), body length alone accounts for 42% of the variance in the length of plastic an animal may ingest and indicates a size ratio of roughly 20:1 between animal body length and the largest plastic the animal may ingest. We expect this work to improve global assessments of plastic pollution risk by introducing a quantifiable link between animals and the plastics they can ingest.

Environment and food web structure interact to alter the trophic magnification of persistent chemicals across river ecosystems.

Legacy organic pollutants persist in freshwater environments, but there is limited understanding of how their trophic transfer and effects vary across riverine ecosystems with different land use, biological communities and food webs. Here, we investigated the trophic magnification of polybrominated diphenyl ethers (PBDEs), polychlorinated biphenyls (PCBs) and a suite of organochlorines (OCs) across nine riverine food webs in contrasting hydrological catchments across South Wales (United Kingdom). Pollutants biomagnified through the food webs in all catchments studied, in some cases reaching levels sufficient for biological effects on invertebrates, fish and river birds such as the Dipper (Cinclus cinclus). Trophic magnification differed across food webs depending on pollutant characteristics (e.g. octanol-water partitioning coefficient) and site-specific environmental conditions (e.g. land use, water chemistry and basal resource composition). The trophic magnification of PBDEs, PCBs and OCs also reflected food-web structure, with greater accumulation in more connected food webs with more generalist taxa. These data highlight interactions between pollutant properties, environmental conditions and biological network structure in the transfer and biomagnification of POPs in river ecosystems. We advocate the need for further investigations of system-specific transfers of contaminants through aquatic food webs as these factors appear to have important implications for risk assessment.

Natural or synthetic - how global trends in textile usage threaten freshwater environments.

As the global demand for textiles increases, so to do the potential environmental impacts that stem from their production, use and disposal. Freshwater ecosystems are particularly at risk: rivers often act as the primary recipients of waste generated during the production of textiles and are subject to pollutants released during the broader lifecycle of a textile product. Here, we investigate how global technological and societal processes shape the way we produce, use and dispose of textiles, and what this means for the environmental quality and ecological health of freshwaters. We examine two predominant 'natural' and synthetic textiles (wool and Polyethylene terephthalate (PET), respectively), and find that risks to freshwater ecosystems vary throughout the lifecycle of these textiles; and across geographies, in-line with regulatory and economic landscapes. Woollen textiles pose most risk during the Production Phase, while PET textiles pose most risk during the Use and Disposal Phases. Our findings show that: (i) both 'natural' and synthetic textiles present substantial challenges for freshwater environments; and (ii) bespoke solutions are needed in areas of the world where the global division of labour and less stringent environmental regulations have concentrated textile production; but also in regions where high textile consumption combines with unsustainable disposal behaviours. Effective mitigation may combine technological advances with societal changes in market mechanisms, regulations, textile use and disposal.

Biological Traits and the Transfer of Persistent Organic Pollutants through River Food Webs.

Freshwater organisms remain at risk from bioaccumulation and biomagnification of persistent organic pollutants (POPs), but factors affecting their transfer through food webs are poorly understood. Here, we investigate transfer pathways of polychlorinated biphenyls, polybrominated diphenyl ethers, and organochlorine through a river food web, assessing the distribution and flux between basal resources (n = 3), macroinvertebrates (n = 22), and fish (n = 1). We investigate the effects of biological traits on the observed patterns and use trait-based models to predict POP bioaccumulation. Transfer pathways differed among POPs and traits such as habitat affinity, feeding behavior, and body size explained some variation in POP burdens between organisms. Trait-based models indicated that relationships between POPs, trophic transfers, and traits were relatively well conserved across a wider array of river food webs. Although providing more consistent predictions of POP bioaccumulation than steady-state models, variability in bioaccumulation across food webs limited the accuracy of trait-model predictions. As some of the first data to illustrate how ecological processes alter the flux of pollutants through river food webs, these results reveal important links between POPs and contrasting energetic pathways. These data also show the utility of trait-based methods in the assessment of persistent contaminants, but further field validations are required.

River organisms as indicators of the distribution and sources of persistent organic pollutants in contrasting catchments.

Persistent organic pollutants (POPs) continue to threaten aquatic organisms, but risk assessments are restricted by poor knowledge of the distribution and quantity of these substances in different biota. Assessments on aquatic invertebrates are particularly scarce. Here, we investigate variation in polybrominated diphenyl ethers (PBDEs), polychlorinated biphenyls (PCBs) and organochlorines (OCs) in sediments, biofilms, macroinvertebrates and fish across rivers in South Wales (UK). Persistent PCB (-118, -153, -180) and PBDE congeners (BDE-47, -99, -100), and OCs (p,p'-dichlorodiphenyldichloroethylene [p,p'-DDE] and dieldrin [HEOD]) dominated the POPs detected, indicating links to historical emissions. Low concentrations of less persistent PBDEs, PCBs and OCs, however, suggest more contemporary sources. Concentrations of POPs were 2-22 times greater in fish than invertebrates, but their detection frequency (>90%) and concentrations (0-304 ng g-1 wet weight) were higher in these organisms than in sediments or biofilms (<10%; 0-12 ng g-1 wet weight). Invertebrates and fish also contained several PCB congeners (28, 52, 77 and 105) and p,p'-dichlorodiphenyltrichloroethane (p,p'-DDT) that were not detected in the environmental samples. Concentrations of PBDEs, PCBs and OCs differed among invertebrate taxa and feeding guilds. After controlling for significant variation among sample types and taxa, PBDEs were found to increase with urban land cover, while increased PCBs were associated with urban land cover and wastewater discharge. These data illustrate how body burdens of POPs across invertebrate and fish taxa provide valuable information on the spatial variation and likely sources of persistent pollutants in freshwater ecosystems. More work is required to resolve differences in POP contamination between taxonomic groups.

Persistent contaminants as potential constraints on the recovery of urban river food webs from gross pollution.

Urban areas contribute substantially to xenobiotic contaminant loads in rivers, but their effects have been investigated more for individual organisms and sensitive taxa, rather than through the emergent properties of communities. Here, we use replicated, catchment-scale sampling of benthic invertebrates and novel multivariate techniques to assess whether urban wastewater contaminants affected the structure and function of river food webs. We postulated that the continued occurrence of selected contaminants in river systems might explain the incomplete recovery of urban rivers from legacy gross pollution. Benthic invertebrate communities were sampled monthly over a year (2016-2017) at 18 sites across 3 river systems in South Wales (United Kingdom). Contaminant sources were characterised using remote sensing, water quality data from routine monitoring and measured concentrations of selected persistent xenobiotic pollutants (polychlorinated biphenyls and polybrominated diphenyl ethers). Urban wastewater discharges had relatively limited effects on river water quality, with small increases in nitrate, phosphate, temperature, conductivity and total dissolved solids in urban systems. Concentrations of polychlorinated biphenyls and polybrominated diphenyl ethers in invertebrates, however, were significantly higher under greater urban land cover and wastewater discharge. Food webs at the most highly contaminated urban sites were characterised by: (i) reduced taxonomic and functional diversity; (ii) simplified food web structure with reduced network connectance; and (iii) reductions in the abundance of prey important for apex predators such as the Eurasian dipper (Cinclus cinclus). Although correlative and partially confounded by other effects, these data provide support for the hypothesis that impairment to food webs resulting from urban pollutants might explain population, community and ecosystem-level effects in urban river systems, and hence incomplete recovery from past pollution.

A catchment-scale perspective of plastic pollution.

Plastic pollution is distributed across the globe, but compared with marine environments, there is only rudimentary understanding of the distribution and effects of plastics in other ecosystems. Here, we review the transport and effects of plastics across terrestrial, freshwater and marine environments. We focus on hydrological catchments as well-defined landscape units that provide an integrating scale at which plastic pollution can be investigated and managed. Diverse processes are responsible for the observed ubiquity of plastic pollution, but sources, fluxes and sinks in river catchments are poorly quantified. Early indications are that rivers are hotspots of plastic pollution, supporting some of the highest recorded concentrations. River systems are also likely pivotal conduits for plastic transport among the terrestrial, floodplain, riparian, benthic and transitional ecosystems with which they connect. Although ecological effects of micro- and nanoplastics might arise through a variety of physical and chemical mechanisms, consensus and understanding of their nature, severity and scale are restricted. Furthermore, while individual-level effects are often graphically represented in public media, knowledge of the extent and severity of the impacts of plastic at population, community and ecosystem levels is limited. Given the potential social, ecological and economic consequences, we call for more comprehensive investigations of plastic pollution in ecosystems to guide effective management action and risk assessment. This is reliant on (a) expanding research to quantify sources, sinks, fluxes and fates of plastics in catchments and transitional waters both independently as a major transport routes to marine ecosystems, (b) improving environmentally relevant dose-response relationships for different organisms and effect pathways, (c) scaling up from studies on individual organisms to populations and ecosystems, where individual effects are shown to cause harm and; (d) improving biomonitoring through developing ecologically relevant metrics based on contemporary plastic research.

Microplastic ingestion by riverine macroinvertebrates.

Although microplastics are a recognised pollutant in marine environments, less attention has been directed towards freshwater ecosystems despite their greater proximity to possible plastic sources. Here, we quantify the presence of microplastic particles (MPs) in river organisms upstream and downstream of five UK Wastewater Treatment Works (WwTWs). MPs were identified in approximately 50% of macroinvertebrate samples collected (Baetidae, Heptageniidae and Hydropsychidae) at concentrations up to 0.14 MP mg tissue-1 and they occurred at all sites. MP abundance was associated with macroinvertebrate biomass and taxonomic family, but MPs occurred independently of feeding guild and biological traits such as habitat affinity and ecological niche. There was no increase in plastic ingestion downstream of WwTW discharges averaged across sites, but MP abundance in macroinvertebrates marginally increased where effluent discharges contributed more to total runoff and declined with increasing river discharge. The ubiquity of microplastics within macroinvertebrates in this case study reveals a potential risk from MPs entering riverine food webs through at least two pathways, involving detritivory and filter-feeding, and we recommend closer attention to freshwater ecosystems in future research.

Endocrine disruption in aquatic systems: up-scaling research to address ecological consequences.

Endocrine-disrupting chemicals (EDCs) can alter biological function in organisms at environmentally relevant concentrations and are a significant threat to aquatic biodiversity, but there is little understanding of exposure consequences for populations, communities and ecosystems. The pervasive nature of EDCs within aquatic environments and their multiple sub-lethal effects make assessments of their impact especially important but also highly challenging. Herein, we review the data on EDC effects in aquatic systems focusing on studies assessing populations and ecosystems, and including how biotic and abiotic processes may affect, and be affected by, responses to EDCs. Recent research indicates a significant influence of behavioural responses (e.g. enhancing feeding rates), transgenerational effects and trophic cascades in the ecological consequences of EDC exposure. In addition, interactions between EDCs and other chemical, physical and biological factors generate uncertainty in our understanding of the ecological effects of EDCs within aquatic ecosystems. We illustrate how effect thresholds for EDCs generated from individual-based experimental bioassays of the types commonly applied using chemical test guidelines [e.g. Organisation for Economic Co-operation and Development (OECD)] may not necessarily reflect the hazards associated with endocrine disruption. We argue that improved risk assessment for EDCs in aquatic ecosystems urgently requires more ecologically oriented research as well as field-based assessments at population-, community- and food-web levels.