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bioRxiv2018

Warming can destabilise predator-prey interactions by shifting the functional response from Type III to Type II

O_LIThe potential for climate change and temperature shifts to affect community stability remains relatively unknown. One mechanism by which temperature may affect stability is by altering trophic interactions. The functional response quantifies the per capita resource consumption by the consumer as a function of resource abundance and is a suitable framework for the description of nonlinear trophic interactions.\nC_LIO_LIWe studied the effect of temperature on a ciliate predator-prey pair (Spathidium sp. and Dexiostoma campylum) by estimating warming effects on the functional response and on the associated conversion efficiency of the predator.\nC_LIO_LIWe recorded prey and predator dynamics over 24 hours and at three temperature levels (15, 20 and 25{degrees}C). To these data we fitted a population dynamic model including the predator functional response, such that the functional response parameters (space clearance rate, handling time, and density dependence of space clearance rate) were estimated for each temperature separately. To evaluate the ecological significance of temperature effects on the functional response parameters we simulated predator-prey population dynamics. We considered the predator-prey system to be destabilised, if the prey was driven extinct by the predator.\nC_LIO_LIEffects of increased temperature included a transition of the functional response from a Type III to a Type II and an increase of the conversion efficiency of the predator. The simulated population dynamics showed a destabilisation of the system with warming, with greater risk of prey extinction at higher temperatures likely caused by the transition from a Type III to a Type II functional response.\nC_LIO_LIWarming-induced shifts from a Type III to II are not commonly considered in modelling studies that investigate how population dynamics respond to warming. Future studies should investigate the mechanism and generality of the effect we observed and simulate temperature effects in complex food webs including shifts in the type of the functional response as well as consider the possibility of a temperature dependent conversion efficiency.\nC_LI

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bioRxiv2018

Terminal Reproductive Investment, Physiological Trade-offs and Pleiotropic Effects: Their effects produce complex immune/reproductive interactions in the cricket Gryllus texensis

O_LIShould females increase or decrease reproduction when attacked by pathogens? Two hypotheses provide opposite predictions. Terminal reproductive investment theory predicts an increase in reproduction, but hypothesized physiological trade-offs between reproduction and immune function might be expected to produce a decrease. There is evidence for both hypotheses. What determines the choice between the two responses remains unclear. We examine the effect of age on the reproductive response to immune challenge in long-wing females of the Texas field cricket, Gryllus texensis, when fed an ecologically valid (limited) diet.\nC_LIO_LIThe limited diet reduced reproductive output. However, immune challenge had no effect on their reproductive output either in young or middle-aged crickets, which is contrary to either prediction.\nC_LIO_LIFlight muscle maintenance correlated negatively with reproductive output, suggesting a physiological trade-off between flight muscle maintenance and reproduction. Within the long-wing variant there was considerable variability in flight muscle maintenance. This variability may mask physiological trade-offs between immunity and reproduction.\nC_LIO_LIMiddle-aged crickets had higher total phenoloxidase (PO) activity in their hemolymph, compared to young females, which is contrary to the terminal investment theory. Given that PO is involved in both immunity and reproduction, the increased PO may reflect simultaneous investment in both functions.\nC_LIO_LIWe identified four proPO transcripts in a published RNA-seq dataset (transcriptome). Three of the proPO genes were expressed either in the fat body or the ovaries (supporting the hypothesis that PO is bifunctional); however, the two organs expressed different subsets. The possible bifunctionality of PO suggests that it may not be an appropriate immune measure for immune/reproductive trade-offs in some species.\nC_LIO_LIIncreasing age may not cue terminal reproductive investment prior to senescence.\nC_LI

ecology↗

bioRxiv2018

Co-occurrence of contrasting life-history strategies in a metapopulation inhabiting temporally variable and stable breeding sites

Life-history theory states that, during the lifetime of an individual, resources are allocated to either somatic maintenance or reproduction. Resource allocation trade-offs determine the evolution and ecology of life-history strategies and determine an organisms position along the fast-slow continuum. Theory predicts that environmental stochasticity is an important driver of resource allocation and therefore life-history evolution. Highly stochastic environments are expected to increase uncertainty in reproductive success and select for iteroparity and a slowing down of the life history. To date, most empirical studies have used comparisons among species to examine these theoretical predictions. By contrast, few have investigated how environmental stochasticity affects life-history strategies at the intraspecific level. In this study, we examined how variation in breeding site stochasticity (among-year variability in pond volume and hydroperiod) promotes the co-occurrence of different life-history strategies in a spatially structured population, and determines life-history position along the fast-slow continuum in the yellow-bellied toad (Bombina variegata). We collected mark-recapture data from a metapopulation and used multievent capture-recapture models to estimate survival, recruitment and breeding probabilities. We found higher survival and longer lifespans in populations inhabiting variable sites compared to those breeding in stable ones. In addition, probabilities of recruitment and skipping a breeding event were higher in variable sites. The temporal variance of survival and recruitment probabilities as well as the probability to skip breeding was higher in variable sites. Taken together, these findings indicate that populations breeding in variable sites experienced a slowing down of the life-history. Our study thus revealed similarities in the macroevolutionary and microevolutionary processes shaping life-history evolution.

ecology↗

bioRxiv2018

Established rodent community delays recovery of dominant competitor following experimental disturbance

Human activities alter processes that control local biodiversity, causing changes in the abundance and identity of species in many ecosystems. However, restoring biodiversity to a previous state is rarely as simple as reintroducing lost species or restoring processes to their pre-disturbance state. Theory suggests that established species can impede shifts in species composition via a variety of mechanisms, including direct interference (e.g. territoriality), preempting resources, or habitat alteration. Here we use a long-term experimental manipulation of a desert rodent community to examine differences in the recolonization dynamics of a dominant competitor (kangaroo rats of the genus Dipodomys) when patches were already occupied by an existing rodent community relative to when patches were empty. Recovery of kangaroo rat populations was slow on plots with an established community of other rodent species, taking approximately two years. In contrast, recovery of kangaroo rat populations was rapid on empty plots with no established residents (approximately 3 months). We found little evidence that the delay in kangaroo rat colonization was due to direct interference from competitors, or could be explained by differences in habitat, implicating resource preemption by the established community as the most likely mechanism. These results demonstrate that the presence of an established alternate community inhibits recolonization by new species, even those that should be dominant in the community. This has important implications for understanding how biodiversity may change in the future, and what processes may slow or prevent this change.\n\nSignificance statementEcological communities are changing due to human activities altering the processes governing local biodiversity. However restoring these processes often fails to restore the previous biodiversity state, implying that additional mechanisms contribute to community dynamics. Here we use an experimental manipulation of a desert rodent community--in which dominant competitors (kangaroo rats) were removed and then reintroduced years later--to show that the presence of previously-established species alters the dynamics of the dominant competitors recovery. Kangaroo rat populations took two years to recover on patches where inferior competitors were already established, compared to three months on uninhabited patches. This suggests that priority effects and initial conditions are critical to consider when predicting community response to disturbance, or in ecological restoration projects.

ecology↗

bioRxiv2018

Aspects of invasiveness of Ludwigia and Nelumbo in shallow temperate fluvial lakes

The relationship between invasive plant functional traits and their invasiveness is still the subject of scientific investigation, and the backgrounds of transition from non-native to invasive species in ecosystems are therefore poorly understood. Furthermore, our current knowledge on species invasiveness is heavily biased toward terrestrial species; we know much less about the influence of allochthonous plant traits on their invasiveness in aquatic ecosystems. We studied physiological and ecological traits of two introduced and three native macrophyte species in the Mantua lakes system (northern Italy). We compared their photophysiology, pigment content, leaf reflectance, and phenology in order to assess how the invasive Nelumbo nucifera and Ludwigia hexapetala perform compared to native species, Nuphar lutea, Nymphaea alba, and Trapa natans. We found L. hexapetala to have higher photosynthetic efficiency and able to tolerate higher light intensities than N. nucifera and the native species especially at extreme weather conditions (prolonged exposure to high light and higher temperatures). Chlorophyll a and b, and carotenoid contents of both allochthonous species was substantially higher than that of the native plants suggesting adaptive response to the ecosystem of Mantua lakes system. Higher variability of recorded data in invasive species also was observed. These observations suggest advanced photosynthetic efficiency of the invasive species, especially L. hexapetala, resulting in faster growth rates and higher productivity. This was supported by the evaluation of seasonal dynamics mapped from satellite remote sensing data. This study provides empirical evidence for the relationship between specific plant physiological traits and invasiveness of aquatic plant species, highlighting the importance of trait studies in predicting ecosystem-level impacts of invasive plant species.

ecology↗

bioRxiv2018

Flowers respond to pollinator sound within minutes by increasing nectar sugar concentration.

Can plants hear? That is, can they sense airborne sounds and respond to them? Here we show that Oenothera drummondii flowers, exposed to the playback sound of a flying bee or to synthetic sound-signals at similar frequencies, produced sweeter nectar within 3 minutes, potentially increasing the chances of cross pollination. We found that the flowers vibrated mechanically in response to these sounds, suggesting a plausible mechanism where the flower serves as the plants auditory sensory organ. Both the vibration and the nectar response were frequency-specific: the flowers responded to pollinator sounds, but not to higher frequency sound. Our results document for the first time that plants can rapidly respond to pollinator sounds in an ecologically relevant way. Sensitivity of plants to pollinator sound can affect plant-pollinator interactions in a wide range of ways: Plants could allocate their resources more adequately, focusing on the time of pollinator activity; pollinators would then be better rewarded per time unit; flower shape may be selected for its effect on hearing ability, and not only on signaling; and pollinators may evolve to make sounds that the flowers can hear. Finally, our results suggest that plants may be affected by other sounds as well, including antropogenic ones.

ecology↗

bioRxiv2018

Eco-evolutionary significance of ‘loners’

Loners, individuals out-of-sync with a coordinated majority, occur frequently in nature. Are loners incidental byproducts of large-scale synchronization attempts or are they part of a mosaic of life-history strategies? Here, we provide the first empirical evidence of naturally occurring heritable variation in loner behavior, using the social amoeba Dictyostelium discoideum. Moreover, we show that Dictyostelium loners--cells that do not join the multicellular life-stage-- result from a dynamic population-partitioning process. Underlying this partitioning, we find evidence that each cell makes a stochastic, signal-based decision resulting in an imperfectly synchronized multicellular development affected by both abiotic (environmental porosity) and biotic (strain-specific signaling) factors. Finally, we predict that when strains differing in their partitioning behavior co-occur, cross-signaling impacts slime-mold diversity across spatio-temporal scales. Loners are therefore critical to understanding collective and social behaviors, multicellular development, and ecological dynamics in D. discoideum. More broadly, across taxa, imperfect synchronization might be adaptive by enabling diversification of life-history strategies.

ecology↗

bioRxiv2019

A simple and effective sampler to collect undisturbed cores from tidal marshes

Core sampling is a common procedure in wetland ecology. PVC tubes are widely used to collect soil cores; however several studies fail to describe even the most typical characteristics of their sampling auger. This work aims to fill this gap and provide a simple and standardized core sampler design. A simple and inexpensive sampler is described for field use. Its main advantages are: 1) extraction of undisturbed cores; 7.6 cm diameter and up to 100 cm depth, 2) it is light-weight, sturdy and reliable, 3) it is made from widely available PVC items and 4) it requires minimal skills to assemble. The sampler is introduced in the wetland soil, by rotation, to the desired depth; the tooth-like edge can cut and penetrate through the dense root systems. The sampler is capped with an industrial-type stopper, and pulled up. The core is held in the sampler by suction - negative pressure. A plunger is then used to slowly remove the soil - core and sub - samples are collected. Alternatively, both ends of sampler could be sealed and taken to the lab. An 8 cm long sub-sample was adequate for soil physical, biogeochemical and molecular analysis. The sampler has been tested for 2 years with more than 200 cores taken from tidal wetlands in Chesapeake Bay, Virginia, USA. A negative correlation between salinity and organic matter content at 3 - 5 and 8 - 10 cm was found. For the deeper samples (48 - 50 cm), a positive correlation between salinity and organic matter was observed. The sampler worked satisfactory and it required no maintenance besides cleaning.

ecology↗

bioRxiv2019

Seasonal and directional dispersal behavior in an ongoing dove invasion

The dispersal behavior that underlies range expansions can be difficult to study. Eurasian Collared-Doves have staged independent northwestward invasions across both Europe and North America, reaching carrying capacity in Europe but continuing to increase exponentially in the Americas, where their dispersal behavior remains enigmatic. I used citizen science observations to investigate Eurasian Collared-Dove dispersal behavior along the North American Pacific coast, a natural barrier to land-bird dispersal. Using coastal and pelagic observations, I assessed the seasonality and directionality of dispersal and its prevalence across years. Dispersal flights peaked in spring, like in Europe, and were north-biased, consistent with northwestward initial colonization. A non-significant trend of increasing dispersal across years may reflect evolution of dispersal via spatial sorting and selection. These results inform management of this invasive dove, raise new questions about evolutionary mechanisms behind the invasion, and exemplify using citizen science to study dispersal behavior, a longstanding challenge in ecology.

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bioRxiv2019

Accounting for imperfect detection in species with sessile life cycle stages: a case study of bumble bee colonies

O_LIFor bumble bees, colonies (not individual workers) are the functional unit of the population. Estimates of colony density are thus critical for understanding population distribution and trends of this important pollinator group. Yet, surveys of bumble bee colonies and other taxa with sessile life cycle states rarely account for imperfect detection.\nC_LIO_LIHere we demonstrate the use of mark-recapture methods to estimate the density of bumble bee colonies across the landscape using standardized survey protocols.\nC_LIO_LIWe found that the probability of detecting colonies in standardized surveys varied considerably across space, through time, and among colonies.\nC_LIO_LIUsing simulations, we also show that imperfect detection can obscure true variation in density among plots, or generate spurious variation in counts even when all plots have the same density. In both cases, we show that mark-recapture can be used to generate unbiased estimates of density, with relatively low search effort compared to conventional survey methods for bumble bee colonies.\nC_LIO_LIOur study illustrates the advantages of mark-recapture for optimizing survey protocols for species with cryptic and sessile life cycle stages, which will be a valuable tool in ongoing studies of pollinator nesting ecology.\nC_LI

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bioRxiv2019

Species dynamics and interactions via metabolically informed consumer-resource models

Quantifying the strength, sign, and origin of species interactions, along with their dependence on environmental context, is at the heart of prediction and understanding in ecological communities. Pairwise interaction models like Lotka-Volterra provide an important and flexible foundation, but notably absent is an explicit mechanism mediating interactions. Consumer-resource models incorporate mechanism, but describing competitive and mutualistic interactions is more ambiguous. Here, we bridge this gap by modeling a coarse-grained version of a species true, cellular metabolism to describe resource consumption via uptake and conversion into biomass, energy, and byproducts. This approach does not require detailed chemical reaction information, but it provides a more explicit description of underlying mechanisms than pairwise interaction or consumer-resource models. Using a model system, we find that when metabolic reactions require two distinct resources we recover Liebigs Law and multiplicative co-limitation in particular limits of the intracellular reaction rates. In between these limits, we derive a more general phenomenological form for consumer growth rate, and we find corresponding rates of secondary metabolite production, allowing us to model competitive and non-competitive interactions (e.g., facilitation). Using the more general form, we show how secondary metabolite production can support coexistence even when two species compete for a shared resource, and we show how differences in metabolic rates change species abundances in equilibrium. Building on these findings, we make the case for incorporating coarse-grained metabolism to update the phenomenology we use to model species interactions.

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bioRxiv2019

The Regime Shift Detector: a model to identify changes in dynamic rules governing populations

Environmental factors interact with internal rules of population regulation, sometimes perturbing systems to alternate dynamics though changes in parameter values. Yet, pinpointing when such changes occur in naturally fluctuating populations is difficult. An algorithmic approach that can identify the timing and magnitude of parameter shifts would facilitate understanding of abrupt ecological transitions with potential to inform conservation and management of species.\n\nThe \"Dynamic Shift Detector\" is an algorithm to identify changes in parameter values governing temporal fluctuations in populations with nonlinear dynamics. The algorithm examines population time series data for the presence, location, and magnitude of parameter shifts. It uses an iterative approach to fitting subsets of time series data, then ranks the fit of break point combinations using model selection, assigning a relative weight to each break. We examined the performance of the Dynamic Shift Detector with simulations and two case studies. Under low environmental/sampling noise, the break point sets selected by the Dynamic Shift Detector contained the true simulated breaks with 70-100% accuracy. The weighting tool generally assigned breaks intentionally placed in simulated data (i.e., true breaks) with weights averaging >0.8 and those due to sampling error (i.e., erroneous breaks) with weights averaging <0.2. In our case study examining an invasion process, the algorithm identified shifts in population cycling associated with variations in resource availability. The shifts identified for the conservation case study highlight a decline process that generally coincided with changing management practices affecting the availability of hostplant resources.\n\nWhen interpreted in the context of species biology, the Dynamic Shift Detector algorithm can aid management decisions and identify critical time periods related to species dynamics. In an era of rapid global change, such tools can provide key insights into the conditions under which population parameters, and their corresponding dynamics, can shift.\n\nAuthor SummaryPopulations naturally fluctuate in abundance, and the rules governing these fluctuations are a result of both internal (density dependent) and external (environmental) processes. For these reasons, pinpointing when changes in populations occur is difficult. In this study, we develop a novel break-point analysis tool for population time series data. Using a density dependent model to describe a populations underlying dynamic process, our tool iterates through all possible break point combinations (i.e., abrupt changes in parameter values) and applies information-theoretic decision tools (i.e. Akaikes Information Criterion corrected for small sample sizes) to determine best fits. Here, we develop the approach, simulate data under a variety of conditions to demonstrate its utility, and apply the tool to two case studies: an invasion of multicolored Asian ladybeetle and declining monarch butterflies. The Dynamic Shift Detector algorithm identified parameter changes that correspond to known environmental change events in both case studies.

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bioRxiv2019

Landscape-scale simulations as a tool in multi-criteria decision making for agri-environment schemes

Increasing concerns over the environmental impacts of agriculture in Europe has led to the introduction of agri-environment schemes (AES) to help mitigate biodiversity loss. However, effectiveness of AES has been mixed and only partially successful in achieving desired outcomes. To improve effectiveness and reduce high costs, multi-criteria decision analysis (MCDA) can help support decision-making and determine the most effective management action. Although MCDA has great potential for evaluating policy measures, it rarely considers the context-dependency of species responses to management practices across different landscapes. Landscape simulations can, therefore, be valuable for reducing the uncertainties when predicting the consequences of management actions. A potential suitable simulation system is the Animal, Landscape, and Man Simulation System (ALMaSS), a mechanistic simulation with can improve MCDA with the automatic integration of a species ecology and behaviour and landscape context. The aim of this study was to demonstrate the effectiveness of ALMaSS in evaluating AES management practices across different landscapes and estimate their ability to achieve the proposed conservation outcomes of three typical species of conservation interest. In this study, the effect of a particular management strategy on a species was dependent on the landscape context, in our case, a combination of landscape structure and the type and distribution of farms, and varied depending on the metrics being measured. Although we did not aim to make recommendations of particular management strategies, we demonstrate how simulations can be used for MCDA to select between management strategies with different costs. Despite the complexity of ALMaSS models, the simulation results provided are easy to interpret. Landscape simulations, such as ALMaSS, can be an important tool in multi-criteria decision making by simulating a wide range of managements and contexts and provide supporting information for filtering management options based on specific conservation goals.

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bioRxiv2019

Metabolic rates of prokaryotic microbes may inevitably rise with global warming

Understanding how the metabolic rates of prokaryotes respond to temperature is fun-damental to our understanding of how ecosystem functioning will be altered by climate change, as these micro-organisms are major contributors to global carbon efflux. Ecological metabolic theory suggests that species living at higher temperatures evolve higher growth rates than those in cooler niches due to thermodynamic constraints. Here, using a global prokaryotic dataset, we find that maximal growth rate at thermal optimum increases with temperature for mesophiles (temperature optima [lsim] 45{degrees}C), but not thermophiles ({gtrsim} 45{degrees}C). Furthermore, short-term (within-day) thermal responses of prokaryotic metabolic rates are typically more sensitive to warming than those of eukaryotes. Given that climatic warming will mostly impact ecosystems in the mesophilic temperature range, we conclude that as microbial communities adapt to higher temperatures, their metabolic rates and therefore, carbon efflux, will inevitably rise. Using a mathematical model, we illustrate the potential global impacts of these findings.

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bioRxiv2019

Risks to pollinators from different land-use transitions: bee species’ responses to agricultural expansion show strong phylogenetic signal

Bee species worldwide are facing a future of further land-use change and intensification. Populations of closely-related species with similar ecological characteristics are likely to respond similarly to such pressures. Such phylogenetic signal in species responses could undermine the stability of pollination services in agricultural and natural systems. We use abundance data from a global compilation of bee assemblages in different land uses to assess the sensitivity of 573 bee species to agricultural expansion, intensification and urbanization; and combine the results with the Bee Tree of Life to assess phylogenetic signal. In addition, we assess whether variation in species sensitivity to land-use change is better explained by phylogenetic or available functional trait differences. Bee species show strong phylogenetic signal in sensitivity to agricultural land expansion but only a weak signal in sensitivity to agricultural intensification and urbanisation. Sensitivities were usually best explained by a combination of functional and phylogenetic distances. This finding suggests that the commonly-recorded traits, despite being meaningful as functional response traits, do not capture all important determinants of bee species vulnerability or resistance. However, it also suggests that model-based predictions of the sensitivity of poorly known species may be sufficient to help guide conservation efforts.

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bioRxiv2019

Maximum entropy models elucidate the contribution of metabolic traits to patterns of community assembly

Aim: Maximum entropy (MaxEnt) models promise a novel approach for understanding community assembly and species abundance patterns. One of these models, the \"Maximum Entropy Theory of Ecology\" (METE) reproduces many observed species abundance patterns, but is based on an aggregated representation of community structure that does not resolve species identity or explicitly represent species-specific functional traits. In this paper, METE is compared to \"Very Entropic Growth\" (VEG), a MaxEnt model with a less aggregated representation of community structure that represents species (more correctly, functional types) in terms of their per capita metabolic rates. We examine the contribution of metabolic traits to the patterns of community assembly predicted by VEG and, through aggregation, compare the results with METE predictions in order to gain insight into the biological factors underlying observed patterns of community assembly. Innovation: We formally compare two MaxEnt-based community models, METE and VEG, that differ as to whether or not they represent species-specific functional traits. We empirically test and compare the metabolic predictions of both models, thereby elucidating the role of metabolic traits in patterns of community assembly. Main Conclusions: Our analysis reveals that a key determinant of community metabolic patterns is the \"density of species\" distribution, defined as the intrinsic number of species with metabolic rates in a given range that are available to a community prior to filtering by environmental constraints. Our analysis suggests that appropriate choice of of the density of species in VEG may lead to more realistic predictions than METE, for which this distribution is not defined, and thus opens up new ways to understanding the link between functional traits and patterns of community assembly.

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bioRxiv2019

Transmission bias's fundamental role in biodiversity change

There is a great need to understand how and why biodiversity, which we define as the variety of organisms found in a given place, changes over time. Current estimates suggest strikingly slow change in traditional measures of biodiversity. These estimates seem to contradict rapid shifts in the abundance of individual species and have led to a rethinking of the mechanisms shaping biodiversity. Conceptual models emphasize the role of competition among species or, more recently, selection on species identity (i.e. selection that favors some species at the expense of others). However, it is difficult to quantify how these mechanisms contribute to biodiversity change. To illustrate this point we present cases where strong competition or selection on species identity leads to no biodiversity change. In view of this disconnect we develop a new approach to studying biodiversity change using the Price equation. We show that biodiversity change responds to selection on species rarity, rather than to either competition or selection on species identity. We then show how this insight can be used to quantify the effects of the mechanisms previously thought to influence biodiversity: 1) selection, 2) (ecological) drift, 3) immigration and 4) speciation. Our results suggest the connection between species fates and their rarity is fundamental to understanding biodiversity change.

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bioRxiv2019

Virome heterogeneity and connectivity in waterfowl and shorebird communities

Models of host-microbe dynamics typically assume a single-host population infected by a single pathogen. In reality, many hosts form multi-species aggregations and may be infected with an assemblage of pathogens. We used a meta-transcriptomic approach to characterize the viromes of nine avian species in the Anseriformes (ducks) and Charadriiformes (shorebirds). This revealed the presence of 27 viral species, of which 24 were novel, including double-stranded RNA viruses (Picobirnaviridae and Reoviridae), single-stranded RNA viruses (Astroviridae, Caliciviridae, Picornaviridae), a retro-transcribing DNA virus (Hepadnaviridae), and a single-stranded DNA virus (Parvoviridae). These viruses comprise multi-host generalist viruses and those that are host-specific, indicative of both virome connectivity and heterogeneity. Virome connectivity was apparent in two well described multi-host virus species (avian coronavirus and influenza A virus) and a novel Rotavirus species that were shared among some Anseriform species, while heterogeneity was reflected in the absence of viruses shared between Anseriformes and Charadriiformes. Notably, within avian host families there was no significant relationship between either host taxonomy or foraging ecology and virome composition, although Anseriform species positive for influenza A virus harboured more additional viruses than those negative for influenza virus. Overall, we demonstrate complex virome structures across host species that co-exist in multi-species aggregations.

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