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Plastic debris is thought to be widespread in freshwater ecosystems globally1. However, a lack of comprehensive and comparable data makes rigorous assessment of its distribution challenging2,3. Here we present a standardized cross-national survey that assesses the abundance and type of plastic debris (>250 μm) in freshwater ecosystems. We sample surface waters of 38 lakes and reservoirs, distributed across gradients of geographical position and limnological attributes, with the aim to identify factors associated with an increased observation of plastics. We find plastic debris in all studied lakes and reservoirs, suggesting that these ecosystems play a key role in the plastic-pollution cycle. Our results indicate that two types of lakes are particularly vulnerable to plastic contamination: lakes and reservoirs in densely populated and urbanized areas and large lakes and reservoirs with elevated deposition areas, long water-retention times and high levels of anthropogenic influence. Plastic concentrations vary widely among lakes; in the most polluted, concentrations reach or even exceed those reported in the subtropical oceanic gyres, marine areas collecting large amounts of debris4. Our findings highlight the importance of including lakes and reservoirs when addressing plastic pollution, in the context of pollution management and for the continued provision of lake ecosystem services.

Spatiotemporal earth system data on freshwaters are poorly organized at present and their rich potential for research and environmental management is barely exploited. Part of this deficiency relates to the spatial structure of river networks, in particular, which requires a specialized workflow for earth system data integration and use. To address this challenge, we developed the GeoFRESH online platform, available at http://geofresh.org/. GeoFRESH provides the integration, processing, management and visualization of standardized spatiotemporal earth system data related to freshwaters on the global scale. Users can (i) upload point data; (ii) automatically assign the points to a new fine-scale hydrographical network; (iii) map the points to the corresponding upstream catchment; (iv) annotate the points with site-specific and upstream environmental information based on a suite of 104 layers capturing topography, climate, landcover and soil characteristics; (v) calculate network distances among points; and (vi) download the compiled data ready for downstream analyses. GeoFRESH capitalizes on open-source PostgreSQL, PostGIS and pgRouting software, building on a new database comprising the global ‘Hydrography90m’ stream network at 90 m resolution, comprising a total of 726 million unique stream segments. The platform is readily expandable thanks to its modular structure and serves as a key element in supporting freshwater science and management relying on high-resolution geospatial analyses. GeoFRESH thus contributes significantly to FAIR data management and exchange by enabling globally standardized stream network queries and offering a powerful tool to stakeholders engaged in water research, sustainable water management and the monitoring of freshwater ecosystem services alike.

Pelagic bacteria can be classified into free‐living and particle‐attached life modes, which either dwell in the water column or attach to suspended particles. Bacteria with a generalist life style, however, can actively shift between these two habitats. Globally increasing densities of natural and artificial particles enhance habitat heterogeneity, with potential consequences for system stability and trophic transfer through aquatic food webs. To better decipher the dynamics of microbial communities, we investigated the influence of adaptive vs. fixed habitat choice on species coexistence for a simplified bacterial community by analyzing a corresponding food web model, consisting of two specialist bacterial prey species (free and attached), a generalist bacterial prey species with the ability to shift between both habitats, and two protist predators, specialized on either water or particle compartment. For simplicity we assume a shared resource pool, considering particles only for colonization but not as a source for nutrients or carbon, that is, inert particles like microplastics or inorganic sediments. The model predicts coexistence on a cyclic attractor between fixed and flexible bacteria, if the costs for adaptive habitat choice can be balanced by adaptation speed. The presence of adaptive prey dampens predator–prey cycle amplitudes, contributing to system stabilization resulting in higher mean predator biomass compared to specialist prey only. Thus, in pelagic microbial systems, flexible habitat choice at the prey level has important implications for system stability and magnitude of energy flow through the microbial loop., German Ministry of Education and Science, German Science Foundation (DFG)

Ammonia-oxidizing archaea (AOA) play a key role in the aquatic nitrogen cycle. Their genetic diversity is viewed as the outcome of evolutionary processes that shaped ancestral transition from terrestrial to marine habitats. However, current genome-wide insights into AOA evolution rarely consider brackish and freshwater representatives or provide their divergence timeline in lacustrine systems. An unbiased global assessment of lacustrine AOA diversity is critical for understanding their origins, dispersal mechanisms, and ecosystem roles. Here, we leveraged continental-scale metagenomics to document that AOA species diversity in freshwater systems is remarkably low compared to marine environments. We show that the uncultured freshwater AOA, “ Candidatus Nitrosopumilus limneticus,” is ubiquitous and genotypically static in various large European lakes where it evolved 13 million years ago. We find that extensive proteome remodeling was a key innovation for freshwater colonization of AOA. These findings reveal the genetic diversity and adaptive mechanisms of a keystone species that has survived clonally in lakes for millennia.

Scoping review protocol

If life exists on Mars, it would face several challenges including the presence of perchlorates, which destabilize biomacromolecules by inducing chaotropic stress. However, little is known about perchlorate toxicity for microorganisms on the cellular level. Here, we present the first proteomic investigation on the perchlorate‐specific stress responses of the halotolerant yeast Debaryomyces hansenii and compare these to generally known salt stress adaptations. We found that the responses to NaCl and NaClO4‐induced stresses share many common metabolic features, for example, signalling pathways, elevated energy metabolism, or osmolyte biosynthesis. Nevertheless, several new perchlorate‐specific stress responses could be identified, such as protein glycosylation and cell wall remodulations, presumably in order to stabilize protein structures and the cell envelope. These stress responses would also be relevant for putative life on Mars, which—given the environmental conditions—likely developed chaotropic defence strategies such as stabilized confirmations of biomacromolecules or the formation of cell clusters.

Speaker presentation

Aquatic ecosystems are frequently overlooked as fungal habitats, although there is increasing evidence that their diversity and ecological importance are greater than previously considered. Aquatic fungi are critical and abundant components of nutrient cycling and food web dynamics, e.g., exerting top-down control on phytoplankton communities and forming symbioses with many marine microorganisms. However, their relevance for microphytobenthic communities is almost unexplored. In the light of global warming, polar regions face extreme changes in abiotic factors with a severe impact on biodiversity and ecosystem functioning. Therefore, this study aimed to describe, for the first time, fungal diversity in Antarctic benthic habitats along the salinity gradient and to determine the co-occurrence of fungal parasites with their algal hosts, which were dominated by benthic diatoms. Our results reveal that Ascomycota and Chytridiomycota are the most abundant fungal taxa in these habitats. We show that also in Antarctic waters, salinity has a major impact on shaping not just fungal but rather the whole eukaryotic community composition, with a diversity of aquatic fungi increasing as salinity decreases. Moreover, we determined correlations between putative fungal parasites and potential benthic diatom hosts, highlighting the need for further systematic analysis of fungal diversity along with studies on taxonomy and ecological roles of Chytridiomycota., Zweitveröffentlichungen der Universität Potsdam : Mathematisch-Naturwissenschaftliche Reihe; 1290

Global freshwater biodiversity is declining dramatically, and meeting the challenges of this crisis requires bold goals and the mobilisation of substantial resources. While the reasons are varied, investments in both research and conservation of freshwater biodiversity lag far behind those in the terrestrial and marine realms. Inspired by a global consultation, we identify 15 pressing priority needs, grouped into five research areas, in an effort to support informed stewardship of freshwater biodiversity. The proposed agenda aims to advance freshwater biodiversity research globally as a critical step in improving coordinated actions towards its sustainable management and conservation.

Plastic debris is widespread in freshwater ecosystems, but a rigorous assessment of its global distribution has been hindered by a lack of comprehensive and comparable data. We performed the first standardized global survey of lakes to assess the quantity and type of plastics (>250μm). We included 38 lakes located in 23 different countries distributed across five continents, spanning different environmental gradients and varying levels of anthropogenic stress. All samples were collected by horizontal trawling of a plankton net and subsequently treated with hydrogen peroxide. We identified 9425 plastic particles, which were classified based on shape, color, and size. Polymer identification was carried out using Raman micro-spectroscopy. Our results showed that the concentration of plastics spanned four orders of magnitude (10-3-101 particles/m3). Fibers (49%) and fragments (41%) were the most frequently detected particles, suggesting a secondary origin of plastic contamination. The most commonly identified polymers were polyester (30%), polypropylene (20%), and polyethylene (16%), which are widely used in short life-cycle products and account for the majority of global plastic production. Further, we found that urban-related attributes of lakes/watersheds influenced the occurrence and type of plastics in lentic systems and larger and deeper lakes with higher retention times are accumulating plastic debris at higher concentrations. info:eu-repo/semantics/publishedVersion

Additional file 2: Fig. S1Number (a) and relative abundance (b) of ARGs detected inriverine ecosystems of eastern China. R represents rhizosphere soil; SBrepresents surface bulk soil (0-20 cm below bulk soil surface); BB represents bottombulk soil (40-60 cm below bulk soil surface); S represents sediment (0-20 cmbelow sediment surface); SR and NR represent rhizosphere soil in the south andnorth, respectively; SSB and NSB represent surface bulk soil (0-20 cm belowbulk soil surface) in the south and north, respectively; SBB and NBB represent bottombulk soil (40-60 cm below bulk soil surface) in the south and north,respectively; SS and NS represent sediment (0-20 cm below sediment surface) inthe south and north, respectively. MLS representmacrolides-lincosamides-streptogramines. Fig.S2. Proportion of detected ARGsin terms of number (a) and relative abundance (b). (Forabbreviations, see legend of Fig. S1). Fig. S3. Relative abundance ofcore ARGs in the riverine ecosystems of eastern China. (For abbreviations, seelegend of Fig. S1). Fig. S3. Relative abundance of core ARGs in the riverineecosystems of eastern China. (For abbreviations, see legend of Fig. S1). Fig.S4. The standard deviation of ARG abundance (a) and subtype number (b)in the riverine ecosystems of eastern China. (For abbreviations, see legend ofFig. S1). Fig.S5. Proportion of detected ARGs (number (a) and abundance (b)) in different habitats in the riverine ecosystem of easternChina. (For abbreviations, see legend of Fig. S1). Fig. S6. Influence of vegetation richness, coverage and plant biomasson ARG abundance. (For abbreviations, see legend of Fig. S1). Fig. S7. SPEC-OCCU plots showing ARGs that differ between two defined groups (i.e. SR vs. SSB&SBB, NR vs. NSB&NBB); the x-axis represents occupancy, i.e. how well an ARG is distributed across all sites of each group; and the y-axis represents specificity, i.e. whether ARGs are also found in other groups. (For abbreviations, see legend of Fig. S1).Fig. S8. RDA of ARGs and physiochemical variables controlling ARGs in different habitats (rhizosphere (R), surface bulk soil (SB), bottom bulk soil (BB), and sediment (S)). Note: *0.01 < P < 0.05, **P < 0.01. (For abbreviations, see legend of Fig. S1). Fig. S9. Network diagrams showing the correlation of individual ARGs and individual parameters in the different habitats. Blue and red lines represent negative and positive relationships, respectively (ρ > 0.8 or ρ < -0.8, P > 0.01). (The complement of Fig. 7). (For abbreviations, see legend of Fig. S1).

Conventional agricultural practices negatively impact soil biodiversity, carbon stocks, and greenhouse gas emissions in ways that make them unsustainable for supporting future supply of food and fiber. Better management of agrobiodiversity will likely play a critical role in transitioning toward more sustainable practices. In particular, innovation and developments targeting the aboveground and belowground components of agroecosystems should be informed by frameworks and approaches that harness the—in particular functional—diversity of complex microbial communities. Here, we review and discuss microbial trait-based approaches that will help us understand and steer agroecosystem functioning in the face of global change. We highlight how trait-based approaches can improve agricultural practices related to soil functioning (e.g., soil fertility and aggregation); climate regulation (e.g., carbon storage and greenhouse gas emissions) and adaptation to climate change; plant health; and reduction of contaminant-related hazards for human health. We also consider how microbial trait-based approaches can be used as a tool to improve cultivated plant performance through artificial selection and microbiome engineering. Last, we discuss the inherent obstacles associated with the development and implementation of trait-based approaches owing to strong interactions within microbial communities and linkages between plants and the soil environment. Despite these obstacles, microbial trait-based approaches hold promise for the sustainable management of agricultural ecosystems needed to feed and nourish a rapidly growing human population.

Space-for-Time-Substitution surveys (SFTS) are commonly used to describe zooplankton community dynamics and to determine lake ecosystem health. SFTS surveys typically combine single point observations from many lakes to evaluate the response of zooplankton community structure and dynamics (e.g., species abundance and biomass, diversity, demographics and modeled rate processes) to spatial gradients in hypothesized environmental drivers (e.g., temperature, nutrients, predation), in lieu of tracking such responses over long time scales. However, the reliability and reproducibility of SFTS zooplankton surveys have not yet been comprehensively tested against empirically-based community dynamics from longterm monitoring efforts distributed worldwide. We use a recently compiled global data set of more than 100 lake zooplankton time series to test whether SFTS surveys can accurately capture zooplankton diversity, and the hypothesized relationship with temperature, using simulated SFTS surveys of the time series data. Specifically, we asked: (1) to what degree can SFTS surveys capture observed biodiversity dynamics; (2) how does timing and duration of sampling affect detected biodiversity patterns; (3) does biodiversity ubiquitously increase with temperature across lakes, or vary by climate zone or lake type; and (4) do results from SFTS surveys produce comparable biodiversity-temperature relationship(s) to empirical data within and among lakes? Testing biodiversity-ecosystem function (BEF) relationships, and the drivers of such relationships, requires a solid data basis. Our work provides a global perspective on the design and usefulness of (long-term) zooplankton monitoring programs and how much confidence we can place in the zooplankton biodiversity patterns observed from SFTS surveys. info:eu-repo/semantics/publishedVersion

Cellulose and chitin are the most abundant polymeric, organic carbon source globally. Thus, microbes degrading these polymers significantly influence global carbon cycling and greenhouse gas production. Fungi are recognized as important for cellulose decomposition in terrestrial environments, but are far less studied in marine environments, where bacterial organic matter degradation pathways tend to receive more attention. In this study, we investigated the potential of fungi to degrade kelp detritus, which is a major source of cellulose in marine systems. Given that kelp detritus can be transported considerable distances in the marine environment, we were specifically interested in the capability of endophytic fungi, which are transported with detritus, to ultimately contribute to kelp detritus degradation. We isolated 10 species and two strains of endophytic fungi from the kelp Ecklonia radiata. We then used a dye decolorization assay to assess their ability to degrade organic polymers (lignin, cellulose, and hemicellulose) under both oxic and anoxic conditions and compared their degradation ability with common terrestrial fungi. Under oxic conditions, there was evidence that Ascomycota isolates produced cellulose-degrading extracellular enzymes (associated with manganese peroxidase and sulfur-containing lignin peroxidase), while Mucoromycota isolates appeared to produce both lignin and cellulose-degrading extracellular enzymes, and all Basidiomycota isolates produced lignin-degrading enzymes (associated with laccase and lignin peroxidase). Under anoxic conditions, only three kelp endophytes degraded cellulose. We concluded that kelp fungal endophytes can contribute to cellulose degradation in both oxic and anoxic environments. Thus, endophytic kelp fungi may play a significant role in marine carbon cycling via polymeric organic matter degradation.

Our understanding of the small-scale processes that drive global biogeochemical cycles and the Earth’s climate is dependent on accurate estimations of interfacial diffusive fluxes to and from biologically-active substrates in aquatic environments. In this study, we present a novel model approach for accurate calculations of diffusive fluxes of dissolved gases, nutrients, and solutes from concentration profiles measured across the substrate-water interfaces using microsensors. The model offers a robust computational scheme for automatized determination of the interface position and enables precise calculations of the interfacial diffusive fluxes simultaneously. In contrast to other methods, the new approach is not restricted to any particular substrate geometry, does not require a priori determination of the interface position for the flux calculation, and, thus, reduces the uncertainties in calculated fluxes arising from partly subjective identification of the interface position. In addition, it is robust when applied to measured profiles containing scattered data points and insensitive to reasonable decreases of the spatial resolution of the data points. The latter feature allows for significantly reducing measurement time which is a crucial factor for in situ experiments.

International audience; The intensity and frequency of storms are projected to increase in many regions of the world because of climate change. Storms can alter environmental conditions in many ecosystems. In lakes and reservoirs, storms can reduce epilimnetic temperatures from wind-induced mixing with colder hypolimnetic waters, direct precipitation to the lake's surface, and watershed runoff. We analyzed 18 long-term and high-frequency lake datasets from 11 countries to assess the magnitude of wind-vs. rainstorm-induced changes in epilimnetic temperature. We found small day-today epilimnetic temperature decreases in response to strong wind and heavy rain during stratified conditions. Day-today epilimnetic temperature decreased, on average, by 0.28 C during the strongest windstorms (storm mean daily wind speed among lakes: 6.7 AE 2.7 m s −1 , 1 SD) and by 0.15 C after the heaviest rainstorms (storm mean daily rainfall: 21.3 AE 9.0 mm). The largest decreases in epilimnetic temperature were observed ≥2 d after sustained strong wind or heavy rain (top 5 th percentile of wind and rain events for each lake) in shallow and medium-depth lakes. The smallest decreases occurred in deep lakes. Epilimnetic temperature change from windstorms, but not rainstorms, was negatively correlated with maximum lake depth. However, even the largest storm-induced mean epilimnetic temperature decreases were typically

There were errors in the name of author László G. Nagy and in affiliation no. 31 in the original publication. The original article has been corrected.

Plans are currently being drafted for the next decade of action on biodiversity-both the post-2020 Global Biodiversity Framework of the Convention on Biological Diversity (CBD) and Biodiversity Strategy of the European Union (EU). Freshwater biodiversity is disproportionately threatened and underprioritized relative to the marine and terrestrial biota, despite supporting a richness of species and ecosystems with their own intrinsic value and providing multiple essential ecosystem services. Future policies and strategies must have a greater focus on the unique ecology of freshwater life and its multiple threats, and now is a critical time to reflect on how this may be achieved. We identify priority topics including environmental flows, water quality, invasive species, integrated water resources management, strategic conservation planning, and emerging technologies for freshwater ecosystem monitoring. We synthesize these topics with decades of first-hand experience and recent literature into 14 special recommendations for global freshwater biodiversity conservation based on the successes and setbacks of European policy, management, and research. Applying and following these recommendations will inform and enhance the ability of global and European post-2020 biodiversity agreements to halt and reverse the rapid global decline of freshwater biodiversity.

The intensity and frequency of storms are projected to increase in many regions of the world because of climate change. Storms can alter environmental conditions in many ecosystems. In lakes and reservoirs, storms can reduce epilimnetic temperatures from wind-induced mixing with colder hypolimnetic waters, direct precipitation to the lake's surface, and watershed runoff. We analyzed 18 long-term and high-frequency lake datasets from 11 countries to assess the magnitude of wind- vs. rainstorm-induced changes in epilimnetic temperature. We found small day-to-day epilimnetic temperature decreases in response to strong wind and heavy rain during stratified conditions. Day-to-day epilimnetic temperature decreased, on average, by 0.28°C during the strongest windstorms (storm mean daily wind speed among lakes: 6.7 ± 2.7 m s −1, 1 SD) and by 0.15°C after the heaviest rainstorms (storm mean daily rainfall: 21.3 ± 9.0 mm). The largest decreases in epilimnetic temperature were observed ≥2 d after sustainedstrong wind or heavy rain (top 5 th percentile of wind and rain events for each lake) in shallow and medium-depth lakes. The smallest decreases occurred in deep lakes. Epilimnetic temperature change from windstorms, but not rainstorms, was negatively correlated with maximum lake depth. However, even the largest storm-induced mean epilimnetic temperature decreases were typically

Insight into how environmental change determines the production and distribution of cyanobacterial toxins is necessary for risk assessment. Management guidelines currently focus on hepatotoxins (microcystins). Increasing attention is given to other classes, such as neurotoxins (e.g., anatoxin-a) and cytotoxins (e.g., cylindrospermopsin) due to their potency. Most studies examine the relationship between individual toxin variants and environmental factors, such as nutrients, temperature and light. In summer 2015, we collected samples across Europe to investigate the effect of nutrient and temperature gradients on the variability of toxin production at a continental scale. Direct and indirect effects of temperature were the main drivers of the spatial distribution in the toxins produced by the cyanobacterial community, the toxin concentrations and toxin quota. Generalized linear models showed that a Toxin Diversity Index (TDI) increased with latitude, while it decreased with water stability. Increases in TDI were explained through a significant increase in toxin variants such as MC-YR, anatoxin and cylindrospermopsin, accompanied by a decreasing presence of MC-LR. While global warming continues, the direct and indirect effects of increased lake temperatures will drive changes in the distribution of cyanobacterial toxins in Europe, potentially promoting selection of a few highly toxic species or strains., Postprints der Universität Potsdam : Mathematisch-Naturwissenschaftliche Reihe; 1105

Deep waters represent the largest biome on Earth and the largest ecosystem of Costa Rica. Fungi play a fundamental role in global biogeochemical cycling in marine sediments, yet, they remain little explored. We studied fungal diversity and community composition in several marine sediments from 16 locations sampled along a bathymetric gradient (from a depth of 380 to 3,474 m) in two transects of about 1,500 km length in the Eastern Tropical Pacific (ETP) of Costa Rica. Sequence analysis of the V7-V8 region of the 18S rRNA gene obtained from sediment cores revealed the presence of 787 fungal amplicon sequence variants (ASVs). On average, we detected a richness of 75 fungal ASVs per sample. Ascomycota represented the most abundant phylum with Saccharomycetes constituting the dominant class. Three ASVs accounted for ca. 63% of all fungal sequences: the yeast Metschnikowia (49.4%), Rhizophydium (6.9%), and Cladosporium (6.7%). We distinguished a cluster composed mainly by yeasts, and a second cluster by filamentous fungi, but we were unable to detect a strong effect of depth and the overlying water temperature, salinity, dissolved oxygen (DO), and pH on the composition of fungal communities. We highlight the need to understand further the ecological role of fungi in deep-sea ecosystems., Postprints der Universität Potsdam : Mathematisch-Naturwissenschaftliche Reihe, 1013

Plans are currently being drafted for the next decade of action on biodiversity-both the post-2020 Global Biodiversity Framework of the Convention on Biological Diversity (CBD) and Biodiversity Strategy of the European Union (EU). Freshwater biodiversity is disproportionately threatened and underprioritized relative to the marine and terrestrial biota, despite supporting a richness of species and ecosystems with their own intrinsic value and providing multiple essential ecosystem services. Future policies and strategies must have a greater focus on the unique ecology of freshwater life and its multiple threats, and now is a critical time to reflect on how this may be achieved. We identify priority topics including environmental flows, water quality, invasive species, integrated water resources management, strategic conservation planning, and emerging technologies for freshwater ecosystem monitoring. We synthesize these topics with decades of first-hand experience and recent literature into 14 special recommendations for global freshwater biodiversity conservation based on the successes and setbacks of European policy, management, and research. Applying and following these recommendations will inform and enhance the ability of global and European post-2020 biodiversity agreements to halt and reverse the rapid global decline of freshwater biodiversity.

Microbial interactions play an essential role in aquatic ecosystems and are of the great interest for both marine and freshwater ecologists. Recent development of new technologies and methods allowed to reveal many functional mechanisms and create new concepts. Yet, many fundamental aspects of microbial interactions have been almost exclusively studied for marine pelagic and benthic ecosystems. These studies resulted in a formulation of the Black Queen Hypothesis, a development of the phycosphere concept for pelagic communities, and a realization of microbial communication as a key mechanism for microbial interactions. In freshwater ecosystems, especially for periphyton communities, studies focus mainly on physiology, biodiversity, biological indication, and assessment, but the many aspects of microbial interactions are neglected to a large extent. Since periphyton plays a great role for aquatic nutrient cycling, provides the basis for water purification, and can be regarded as a hotspot of microbial biodiversity, we highlight that more in-depth studies on microbial interactions in periphyton are needed to improve our understanding on functioning of freshwater ecosystems. In this paper we first present an overview on recent concepts (e.g., the “Black Queen Hypothesis”) derived from state-of-the-art OMICS methods including metagenomics, metatranscriptomics, and metabolomics. We then point to the avenues how these methods can be applied for future studies on biodiversity and the ecological role of freshwater periphyton, a yet largely neglected component of many freshwater ecosystems.

Biogenic greenhouse gas emissions, e.g., of methane (CH4) and carbon dioxide (CO2) from inland waters, contribute substantially to global warming. In aquatic systems, dissolved greenhouse gases are highly heterogeneous in both space and time. To better understand the biological and physical processes that affect sources and sinks of both CH4 and CO2, their dissolved concentrations need to be measured with high spatial and temporal resolution. To achieve this goal, we developed the Fast-Response Automated Gas Equilibrator (FaRAGE) for real-time in situ measurement of dissolved CH4 and CO2 concentrations at the water surface and in the water column. FaRAGE can achieve an exceptionally short response time (t95 %=12 s when including the response time of the gas analyzer) while retaining an equilibration ratio of 62.6 % and a measurement accuracy of 0.5 % for CH4. A similar performance was observed for dissolved CO2 (t95 %=10 s, equilibration ratio 67.1 %). An equilibration ratio as high as 91.8 % can be reached at the cost of a slightly increased response time (16 s). The FaRAGE is capable of continuously measuring dissolved CO2 and CH4 concentrations in the nM-to-sub mM (10−9–10−3 mol L−1) range with a detection limit of sub-nM (10−10 mol L−1), when coupling with a cavity ring-down greenhouse gas analyzer (Picarro GasScouter). FaRAGE allows for the possibility of mapping dissolved concentration in a “quasi” three-dimensional manner in lakes and provides an inexpensive alternative to other commercial gas equilibrators. It is simple to operate and suitable for continuous monitoring with a strong tolerance for suspended particles. While the FaRAGE is developed for inland waters, it can be also applied to ocean waters by tuning the gas–water mixing ratio. The FaRAGE is easily adapted to suit other gas analyzers expanding the range of potential applications, including nitrous oxide and isotopic composition of the gases.

Remote sensing and field spectroscopy of natural waters is typically performed under clear skies, low wind speeds and low solar zenith angles. Such measurements can also be made, in principle, under clouds and mixed skies using airborne or in-situ measurements; however, variable illumination conditions pose a challenge to data analysis. In the present case study, we evaluated the inversion of hyperspectral in-situ measurements for water constituent retrieval acquired under variable cloud cover. First, we studied the retrieval of Chlorophyll-a (Chl-a) concentration and colored dissolved organic matter (CDOM) absorption from in-water irradiance measurements. Then, we evaluated the errors in the retrievals of the concentration of total suspended matter (TSM), Chl-a and the absorption coefficient of CDOM from above-water reflectance measurements due to highly variable reflections at the water surface. In order to approximate cloud reflections, we extended a recent three-component surface reflectance model for cloudless atmospheres by a constant offset and compared different surface reflectance correction procedures. Our findings suggest that in-water irradiance measurements may be used for the analysis of absorbing compounds even under highly variable weather conditions. The extended surface reflectance model proved to contribute to the analysis of above-water reflectance measurements with respect to Chl-a and TSM. Results indicate the potential of this approach for all-weather monitoring.

Polynucleobacter asymbioticus strain QLW-P1DMWA-1T represents a group of highly successful heterotrophic ultramicrobacteria that is frequently very abundant (up to 70% of total bacterioplankton) in freshwater habitats across all seven continents. This strain was originally isolated from a shallow Alpine pond characterized by rapid changes in water temperature and elevated UV radiation due to its location at an altitude of 1300 m. To elucidate the strain’s adjustment to fluctuating environmental conditions, we recorded changes occurring in its transcriptomic and proteomic profiles under contrasting experimental conditions by simulating thermal conditions in winter and summer as well as high UV irradiation. To analyze the potential connection between gene expression and regulation via methyl group modification of the genome, we also analyzed its methylome. The methylation pattern differed between the three treatments, pointing to its potential role in differential gene expression. An adaptive process due to evolutionary pressure in the genus was deduced by calculating the ratios of non-synonymous to synonymous substitution rates for 20 Polynucleobacter spp. genomes obtained from geographically diverse isolates. The results indicate purifying selection., Postprints der Universität Potsdam : Mathematisch-Naturwissenschaftliche Reihe, 1011

The majority of microplastic research has focused on seawater, with fewer than 4% of microplasticsrelated studies occurring on freshwaters. The limited available information suggests that the abundance of microplastics in freshwaters is often as high or even higher than marine environments. However, comprehensive investigations on occurrence and fate of microplastics in freshwaters are scarce and highly fragmented, partly because detection and identification of microplastic particles is rather complex. In addition, up to now, harmonized and standardized protocols for the sampling and analysis of microplastics in freshwaters do not exist, and studies with different research aims and hypotheses often report unstandardized results, making comparison among studies difficult. In the present study, we performed the first global standardized sampling and analysis effort to investigate the occurrence and distribution of microplastics in surface water of lakes and reservoirs with different anthropogenic impacts. Participants aim to collect water samples of freshwater systems with different features (e.g., area, depth, thermal behavior, watershed), following a common protocol. This 18 establishes the collection of samples by horizontal trawling of a plankton net and, after treatment with hydrogen peroxide, the polymer identification through micro-Raman spectroscopy. This GLEON project will allow obtaining comparable data about microplastic contamination in different freshwater systems around the globe. With this global dataset, our goals are to determine whether a relationship exists between the abundance of microplastics and the waterbody/watershed attributes and understand which factors are likely to influence the occurrence of microplastics in surface water of lentic systems. info:eu-repo/semantics/publishedVersion

Microbial interactions play an essential role in aquatic ecosystems and are of the great interest for both marine and freshwater ecologists. Recent development of new technologies and methods allowed to reveal many functional mechanisms and create new concepts. Yet, many fundamental aspects of microbial interactions have been almost exclusively studied for marine pelagic and benthic ecosystems. These studies resulted in a formulation of the Black Queen Hypothesis, a development of the phycosphere concept for pelagic communities, and a realization of microbial communication as a key mechanism for microbial interactions. In freshwater ecosystems, especially for periphyton communities, studies focus mainly on physiology, biodiversity, biological indication, and assessment, but the many aspects of microbial interactions are neglected to a large extent. Since periphyton plays a great role for aquatic nutrient cycling, provides the basis for water purification, and can be regarded as a hotspot of microbial biodiversity, we highlight that more in-depth studies on microbial interactions in periphyton are needed to improve our understanding on functioning of freshwater ecosystems. In this paper we first present an overview on recent concepts (e.g., the “Black Queen Hypothesis”) derived from state-of-the-art OMICS methods including metagenomics, metatranscriptomics, and metabolomics. We then point to the avenues how these methods can be applied for future studies on biodiversity and the ecological role of freshwater periphyton, a yet largely neglected component of many freshwater ecosystems., Postprints der Universität Potsdam : Mathematisch-Naturwissenschaftliche Reihe, 969

International audience

The oligotrophic subtropical gyre covers a vast area of the Atlantic Ocean. Decades of time-series monitoring have generated detailed temporal information about zooplankton species and abundances at fixed locations within the gyre, but their live/dead status is often omitted, especially in the dynamic subtropical convergence zone (STCZ) where the water column stratification pattern can change considerably across the front as warm and cold water masses converge. We conducted a detailed survey in the North Atlantic STCZ and showed that over 85% of the copepods were typically concentrated in the upper 200 m. Copepod carcasses were present in all samples and their proportional numerical abundances increased with depth, reaching up to 91% at 300-400 m. Overall, 14-19% of the copepods within the upper 200 m were carcasses. Shipboard experiments showed that during carcass decomposition, microbial respiration increased, and the bacterial community associated with the carcasses diverged from that in the ambient water. Combining field and experimental data, we estimated that decomposing copepod carcasses constitute a negligible oxygen sink in the STCZ, but sinking carcasses may represent an overlooked portion of the passive carbon sinking flux and should be incorporated in future studies of carbon flux in this area.

Our growing awareness of the microbial world's importance and diversity contrasts starkly with our limited understanding of its fundamental structure. Despite recent advances in DNA sequencing, a lack of standardized protocols and common analytical frameworks impedes comparisons among studies, hindering the development of global inferences about microbial life on Earth. Here we present a meta-analysis of microbial community samples collected by hundreds of researchers for the Earth Microbiome Project. Coordinated protocols and new analytical methods, particularly the use of exact sequences instead of clustered operational taxonomic units, enable bacterial and archaeal ribosomal RNA gene sequences to be followed across multiple studies and allow us to explore patterns of diversity at an unprecedented scale. The result is both a reference database giving global context to DNA sequence data and a framework for incorporating data from future studies, fostering increasingly complete characterization of Earth's microbial diversity. Fil: Thompson, Luke R.. University of California at San Diego; Estados Unidos. University of Southern Mississippi; Estados Unidos. National Oceanic and Atmospheric Administration; Estados Unidos Fil: Sanders, Jon G.. University of California at San Diego; Estados Unidos Fil: McDonald, Daniel. University of California at San Diego; Estados Unidos Fil: Amir, Amnon. University of California at San Diego; Estados Unidos Fil: Ladau, Joshua. University of California San Francisco; Estados Unidos Fil: Locey, Kenneth J.. Indiana University; Estados Unidos Fil: Prill, Robert J.. IBM Almaden Research Center; Estados Unidos Fil: Tripathi, Anupriya. University of California at San Diego; Estados Unidos Fil: Gibbons, Sean M.. Massachusetts Institute of Technology; Estados Unidos Fil: Ackermann, Gail. University of California San Diego; Estados Unidos Fil: Navas Molina, Jose A.. University of California at San Diego; Estados Unidos Fil: Janssen, Stefan. University of California at San Diego; Estados Unidos Fil: Kopylova, Evguenia. University of California at San Diego; Estados Unidos Fil: Vázquez Baeza, Yoshiki. University of California at San Diego; Estados Unidos Fil: González, Antonio. University of California at San Diego; Estados Unidos Fil: Morton, James T.. University of California at San Diego; Estados Unidos Fil: Mirarab, Siavash. University of California at San Diego; Estados Unidos Fil: Xu, Zhenjiang Zech. University of California at San Diego; Estados Unidos Fil: Jiang, Lingjing. University of California at San Diego; Estados Unidos Fil: Haroon, Mohamed F.. Harvard University; Estados Unidos Fil: Kanbar, Jad. University of California at San Diego; Estados Unidos Fil: Zhu, Qiyun. University of California at San Diego; Estados Unidos Fil: Song, Se Jin. University of California at San Diego; Estados Unidos Fil: Kosciolek, Tomasz. University of California at San Diego; Estados Unidos Fil: Bokulich, Nicholas A.. Northern Arizona University; Estados Unidos Fil: Lefler, Joshua. University of California at San Diego; Estados Unidos Fil: Brislawn, Colin J.. Earth and Biological Sciences Directorate; Estados Unidos Fil: Humphrey, Gregory. University of California at San Diego; Estados Unidos Fil: Owens, Sarah M.. Argonne National Laboratory; Estados Unidos Fil: Hampton Marcell, Jarrad. Argonne National Laboratory; Estados Unidos. University of Illinois at Chicago; Estados Unidos Fil: Berg Lyons, Donna. University of Colorado; Estados Unidos Fil: McKenzie, Valerie. University of Colorado; Estados Unidos Fil: Fierer, Noah. University of Colorado; Estados Unidos Fil: Fuhrman, Jed A.. University of Colorado; Estados Unidos Fil: Clauset, Aaron. University of Colorado; Estados Unidos Fil: Stevens, Rick L.. Argonne National Laboratory; Estados Unidos. University of Chicago; Estados Unidos Fil: Shade, Ashley. Michigan State University; Estados Unidos Fil: Pollard, Katherine S.. The Gladstone Institutes; Estados Unidos. University of California San Francisco; Estados Unidos Fil: Goodwin, Kelly D.. National Oceanic and Atmospheric Administration; Estados Unidos Fil: Jansson, Janet K.. Earth and Biological Sciences Directorate; Estados Unidos Fil: Gilbert, Jack A.. Argonne National Laboratory; Estados Unidos. University of Chicago; Estados Unidos Fil: Knight, Rob. University of California San Diego; Estados Unidos Fil: Dionisi, Hebe Monica. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Centro Nacional Patagónico. Centro para el Estudio de Sistemas Marinos; Argentina Fil: Lozada, Mariana. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Centro Nacional Patagónico. Centro para el Estudio de Sistemas Marinos; Argentina Fil: Mac Cormack, Walter Patricio. Universidad de Buenos Aires; Argentina. Ministerio de Relaciones Exteriores, Comercio Interno y Culto. Dirección Nacional del Antártico. Instituto Antártico Argentino; Argentina

Many microorganisms produce resting cells with low metabolic activity that allow them to survive prolonged nutrient or energy stress. We show that Prochlorococcus , the dominant phytoplankton lineage in large regions of the nutrient-poor ocean, cannot survive extended stress alone. In late stationary phase cultures, some cells retain metabolic activity (single-cell carbon and nitrogen uptake), but these axenic cultures do not re-grow when transferred into new media. However, when co-cultured with a heterotrophic bacterium, Prochlorococcus survived nutrient starvation for months. This dependence on heterotrophic bacteria is not limited to nutrient starvation - up to 10% of the Prochlorococcus cells in the oceans live under conditions of light starvation, surviving by utilizing organic compounds released by other microorganisms. We propose that reliance on co-occurring heterotrophic bacteria or on the organic compounds they produce, rather than the ability to survive extended starvation as resting cells, underlies the ecological success of Prochlorococcus .

In aquatic ecosystems, light availability can significantly influence microbial turnover of terrestrial organic matter through associated metabolic interactions between phototrophic and heterotrophic communities. However, particularly in streams, microbial functions vary significantly with the structure of the streambed, that is the distribution and spatial arrangement of sediment grains in the streambed. It is therefore essential to elucidate how environmental factors synergistically define the microbial turnover of terrestrial organic matter in order to better understand the ecological role of photoheterotrophic interactions in stream ecosystem processes. In outdoor experimental streams, we examined how the structure of streambeds modifies the influence of light availability on microbial turnover of leaf carbon (C). Furthermore, we investigated whether the studied relationships of microbial leaf C turnover to environmental conditions are affected by flow intermittency commonly occurring in streams. We applied leaves enriched with a C-13-stable isotope tracer and combined quantitative and isotope analyses. We thereby elucidated whether treatment induced changes in C turnover were associated with altered use of leaf C within the microbial food web. Moreover, isotope analyses were combined with measurements of microbial community composition to determine whether changes in community function were associated with a change in community composition. In this study, we present evidence, that environmental factors interactively determine how phototrophs and heterotrophs contribute to leaf C turnover. Light availability promoted the utilization of leaf C within the microbial food web, which was likely associated with a promoted availability of highly bioavailable metabolites of phototrophic origin. However, our results additionally confirm that the structure of the streambed modifies light-related changes in microbial C turnover. From our observations, we conclude that the streambed structure influences the strength of photo-heterotrophic interactions by defining the spatial availability of algal metabolites in the streambed and the composition of microbial communities. Collectively, our multifactorial approach provides valuable insights into environmental controls on the functioning of stream ecosystems.

Growing attention to phytoplankton mixotrophy as a trophic strategy has led to significant revisions of traditional pelagic food web models and ecosystem functioning. Although some empirical estimates of mixotrophy do exist, a much broader set of in situ measurements are required to (i) identify which organisms are acting as mixotrophs in real time and to (ii) assess the contribution of their heterotrophy to biogeochemical cycling. Estimates are needed through time and across space to evaluate which environmental conditions or habitats favour mixotrophy: conditions still largely unknown. We review methodologies currently available to plankton ecologists to undertake estimates of plankton mixotrophy, in particular nanophytoplankton phago-mixotrophy. Methods are based largely on fluorescent or isotopic tracers, but also take advantage of genomics to identify phylotypes and function. We also suggest novel methods on the cusp of use for phago-mixotrophy assessment, including single-cell measurements improving our capacity to estimate mixotrophic activity and rates in wild plankton communities down to the single-cell level. Future methods will benefit from advances in nanotechnology, micromanipulation and microscopy combined with stable isotope and genomic methodologies. Improved estimates of mixotrophy will enable more reliable models to predict changes in food web structure and biogeochemical flows in a rapidly changing world.

Fungi are phylogenetically and functionally diverse ubiquitous components of almost all ecosystems on Earth, including aquatic environments stretching from high montane lakes down to the deep ocean. Aquatic ecosystems, however, remain frequently overlooked as fungal habitats, although fungi potentially hold important roles for organic matter cycling and food web dynamics. Recent methodological improvements have facilitated a greater appreciation of the importance of fungi in many aquatic systems, yet a conceptual framework is still missing. In this Review, we conceptualize the spatiotemporal dimensions, diversity, functions and organismic interactions of fungi in structuring aquatic food webs. We focus on currently unexplored fungal diversity, highlighting poorly understood ecosystems, including emerging artificial aquatic habitats.

Salinity is a significant factor for structuring microbial communities, but little is known for aquatic fungi, particularly in the pelagic zone of brackish ecosystems. In this study, we explored the diversity and composition of fungal communities following a progressive salinity decline (from 34 to 3 PSU) along three transects of ca. 2000 km in the Baltic Sea, the world’s largest estuary. Based on 18S rRNA gene sequence analysis, we detected clear changes in fungal community composition along the salinity gradient and found significant differences in composition of fungal communities established above and below a critical value of 8 PSU. At salinities below this threshold, fungal communities resembled those from freshwater environments, with a greater abundance of Chytridiomycota, particularly of the orders Rhizophydiales, Lobulomycetales, and Gromochytriales. At salinities above 8 PSU, communities were more similar to those from marine environments and, depending on the season, were dominated by a strain of the LKM11 group (Cryptomycota) or by members of Ascomycota and Basidiomycota. Our results highlight salinity as an important environmental driver also for pelagic fungi, and thus should be taken into account to better understand fungal diversity and ecological function in the aquatic realm. UCR::Vicerrectoría de Docencia::Ciencias Básicas::Facultad de Ciencias::Escuela de Biología

Background Mortality is a main driver in zooplankton population biology but it is poorly constrained in models that describe zooplankton population dynamics, food web interactions and nutrient dynamics. Mortality due to non-predation factors is often ignored even though anecdotal evidence of non-predation mass mortality of zooplankton has been reported repeatedly. One way to estimate non-predation mortality rate is to measure the removal rate of carcasses, for which sinking is the primary removal mechanism especially in quiescent shallow water bodies. Objectives and Results We used sediment traps to quantify in situ carcass sinking velocity and non-predation mortality rate on eight consecutive days in 2013 for the cladoceran Bosmina longirostris in the oligo-mesotrophic Lake Stechlin; the outcomes were compared against estimates derived from in vitro carcass sinking velocity measurements and an empirical model correcting in vitro sinking velocity for turbulence resuspension and microbial decomposition of carcasses. Our results show that the latter two approaches produced unrealistically high mortality rates of 0.58-1.04 d(-1), whereas the sediment trap approach, when used properly, yielded a mortality rate estimate of 0.015 d(-1), which is more consistent with concurrent population abundance data and comparable to physiological death rate from the literature. Ecological implications Zooplankton carcasses may be exposed to water column microbes for days before entering the benthos; therefore, non-predation mortality affects not only zooplankton population dynamics but also microbial and benthic food webs. This would be particularly important for carbon and nitrogen cycles in systems where recurring mid-summer decline of zooplankton population due to non-predation mortality is observed.

Across a landscape, aquatic-terrestrial interfaces within and between ecosystems are hotspots of organic matter (OM) mineralization. These interfaces are characterized by sharp spatio-temporal changes in environmental conditions, which affect OM properties and thus control OM mineralization and other transformation processes. Consequently, the extent of OM movement at and across aquatic-terrestrial interfaces is crucial in determining OM turnover and carbon (C) cycling at the landscape scale. Here, we propose expanding current concepts in aquatic and terrestrial ecosystem sciences to comprehensively evaluate OM turnover at the landscape scale. We focus on three main concepts toward explaining OM turnover at the landscape scale: the landscape spatio-temporal context, OM turnover described by priming and ecological stoichiometry, and anthropogenic effects as a disruptor of natural OM transfer magnitudes and pathways. A conceptual framework is introduced that allows for discussing the disparities in spatial and temporal scales of OM transfer, changes in environmental conditions, ecosystem connectivity, and microbial–substrate interactions. The potential relevance of priming effects in both terrestrial and aquatic systems is addressed. For terrestrial systems, we hypothesize that the interplay between the influx of OM and its corresponding elemental composition and the elemental demand of the microbial communities – stoichiometric question – may alleviate spatial and metabolic thresholds. In comparison, substrate level OM dynamics may be substantially different in aquatic systems due to matrix effects that accentuate the role of abiotic conditions, substrate quality, and microbial community dynamics. We highlight the disproportionate impact anthropogenic activities can have on OM cycling across the landscape including reversing natural OM flows through the landscape, disrupting ecosystem connectivity, and nutrient additions that cascade across the landscape. This knowledge is crucial for a better understanding of OM cycling in a landscape context, in particular since terrestrial and aquatic compartments may respond differently to the ongoing changes in climate, land use, and other anthropogenic interferences.

We describe a low-cost three-dimensional underwater particle tracking velocimetry system to directly measure particle settling rate and flux in low-turbulence aquatic environments. The system consists of two waterproof cameras that acquire stereoscopic videos of sinking particles at 48 frames s���1 over a tunable sampling volume of about 45 �� 25 �� 24 cm. A dedicated software package has been developed to allow evaluation of particle velocities, concentration and flux, but also of morphometric parameters such as particle area, sinking angle, shape irregularity, and density. Our method offers several advantages over traditional approaches, like sediment trap or expensive in situ camera systems: (1) it does not require beforehand particle collection and handling; (2) it is not subjected to sediment trap biases from turbulence, horizontal advection, or presence of swimmers, that may alter particulate load and flux; (3) the camera system enables faster data processing and flux computation at higher spatial resolution; (4) apart from the particle settling rates, the particle size distribution, and morphology is determined. We tested the camera system in Lake Stechlin (Germany) in low turbulence and mean flow, and analyzed the morphological properties and settling rates of particles to determine their sinking behavior. The particle flux assessed from conventional sediment trap measurements agreed well with that determined by our system. By this, the low-cost approach demonstrated its reliability in low turbulence environments and a strong potential to provide new insights into particulate carbon transport in aquatic systems. Extension of the method to more turbulent and advective conditions is also discussed.

Aquatic hyphomycetes equipped with a rich array of extracellular enzymes are involved in bioremediation of anthropogenic pollutants, plastics, aromatic contaminants and petroleum hydrocarbons. The current study contributes to the knowledge of Iranian aquatic hyphomycetes. Sampling was conducted in Anzali lagoon, Gilan province, Iran during 2017. Among 55 fungal isolates obtained during this study, eight species including Arthrobotrys oligosporus, Ceratorhiza hydrophila, Fusarium cf. ensiforme, F. incarnatum species complex, Myrmecridium schulzeri, Paecilomyces variotii, Sarocladium subulatum, and Volutella citrinella were identified based on their morphological characteristics. Furthermore, three species assigned to the genera Arthrobotrys, Fusarium and Sarocladium remained unidentified. Molecular studies using mainly ITS, LSU and SSU rDNA and in some cases β-tubulin (tub2) and translation elongation factor 1-a (tef1) determined the phylogenetic position of the isolates among closely related species. The occurrence of members of Fusarium, Sarocladium, and Ceratorhiza, known as rice disease agents, in freshwater ecosystems is interesting. This is the first report of aquatic hyphomycetes communities in Anzali lagoon, Iran. In this study, Sarocladium subulatum, Myrmecridium schulzeri, and Volutella citrinella are reported for the first time for Iran. UCR::Vicerrectoría de Docencia::Ciencias Básicas::Facultad de Ciencias::Escuela de Biología

River ecosystems receive and process vast quantities of terrestrial organic carbon, the fate of which depends strongly on microbial activity. Variation in and controls of processing rates, however, are poorly characterized at the global scale. In response, we used a peer-sourced research network and a highly standardized carbon processing assay to conduct a global-scale field experiment in greater than 1000 river and riparian sites. We found that Earth’s biomes have distinct carbon processing signatures. Slow processing is evident across latitudes, whereas rapid rates are restricted to lower latitudes. Both the mean rate and variability decline with latitude, suggesting temperature constraints toward the poles and greater roles for other environmental drivers (e.g., nutrient loading) toward the equator. These results and data set the stage for unprecedented “next-generation biomonitoring” by establishing baselines to help quantify environmental impacts to the functioning of ecosystems at a global scale. This research was supported by awards to S.D.T. from the Ecuadorian Ministry of Science [Secretaría de Educación Superior Ciencia, Tecnología e Innovación (SENESCYT)] through the PROMETEO scholar exchange program, the Oakland University Research Development Grant program, and a Huron Mountain Wildlife Foundation research grant. N.A.G. was supported by the U.S. Department of Energy’s Office of Science, Biological and Environmental Research. Oak Ridge National Laboratory is managed by UT-Battelle, LLC, for the U.S. Department of Energy under contract DE-AC05-00OR22725. We are grateful for open-access-publishing funds from Kresge Library at Oakland University and Queen’s University Belfast. This research was supported by awards to S.D.T. from the Ecuadorian Ministry of Science [Secretaría de Educación Superior Ciencia, Tecnología e Innovación (SENESCYT)] through the PROMETEO scholar exchange program, the Oakland University Research Development Grant program, and a Huron Mountain Wildlife Foundation research grant. N.A.G. was supported by the U.S. Department of Energy’s Office of Science, Biological and Environmental Research. Oak Ridge National Laboratory is managed by UT-Battelle, LLC, for the U.S. Department of Energy under contract DE-AC05-00OR22725. We are grateful for open-access-publishing funds from Kresge Library at Oakland University and Queen’s University Belfast.