Global terrestrial invasions: Where naturalised birds, mammals, and plants might spread next and what affects this process.

More species live outside their native range than at any point in human history. Yet, there is little understanding of the geographic regions that will be threatened if these species continue to spread, nor of whether they will spread. We predict the world's terrestrial regions to which 833 naturalised plants, birds, and mammals are most imminently likely to spread, and investigate what factors have hastened or slowed their spread to date. There is huge potential for further spread of naturalised birds in North America, mammals in Eastern Europe, and plants in North America, Eastern Europe, and Australia. Introduction history, dispersal, and the spatial distribution of suitable areas are more important predictors of species spread than traits corresponding to habitat usage or biotic interactions. Natural dispersal has driven spread in birds more than in plants. Whether these taxa continue to spread more widely depends partially on connectivity of suitable environments. Plants show the clearest invasion lag, and the putative importance of human transportation indicates opportunities to slow their spread. Despite strong predictive effects, questions remain, particularly why so many birds in North America do not occupy climatically suitable areas close to their existing ranges.

Cat: Empirical modelling of Felis catus population dynamics in the UK.

Domestic cats are popular companion animals, however not all live in human homes and many cats live within shelters or as free-roaming, unowned- feral or stray cats. Cats can transition between these subpopulations, but the influence of this connectivity on overall population dynamics, and the effectiveness of management interventions, remain poorly understood. We developed a UK-focused multistate Matrix Population Model (MPM), combining multiple life history parameters into an integrated model of cat demography and population dynamics. The model characterises cats according to their age, subpopulation and reproductive status, resulting in a 28-state model. We account for density-dependence, seasonality and uncertainty in our modelled projections. Through simulations, we examine the model by testing the effect of different female owned-cat neutering scenarios over a 10-year projection timespan. We also use the model to identify the vital rates to which total population growth is most sensitive. The current model framework demonstrates that increased prevalence of neutering within the owned cat subpopulation influences the population dynamics of all subpopulations. Further simulations find that neutering owned cats younger is sufficient to reduce overall population growth rate, regardless of the overall neutering prevalence. Population growth rate is most influenced by owned cat survival and fecundity. Owned cats, which made up the majority of our modelled population, have the most influence on overall population dynamics, followed by stray, feral and then shelter cats. Due to the importance of owned-cat parameters within the current model framework, we find cat population dynamics are most sensitive to shifts in owned cat husbandry. Our results provide a first evaluation of the demography of the domestic cat population in the UK and provide the first structured population model of its kind, thus contributing to a wider understanding of the importance of modelling connectivity between subpopulations. Through example scenarios we highlight the importance of studying domestic cat populations in their entirety to better understand factors influencing their dynamics and to guide management planning. The model provides a theoretical framework for further development, tailoring to specific geographies and experimental investigation of management interventions.

Multi-locus homozygosity promotes actuarial senescence in a wild mammal.

Genome-wide homozygosity, caused for example by inbreeding, is expected to have deleterious effects on survival and/or reproduction. Evolutionary theory predicts that any fitness costs are likely to be detected in late life because natural selection will filter out negative impacts on younger individuals with greater reproductive value. Here we infer associations between multi-locus homozygosity (MLH), sex, disease and age-dependent mortality risks using Bayesian analysis of the life histories of wild European badgers Meles meles in a population naturally infected with Mycobacterium bovis (the causative agent of bovine tuberculosis [bTB]). We find important effects of MLH on all parameters of the Gompertz-Makeham mortality hazard function, but particularly in later life. Our findings confirm the predicted association between genomic homozygosity and actuarial senescence. Increased homozygosity is particularly associated with an earlier onset, and greater rates of actuarial senescence, regardless of sex. The association between homozygosity and actuarial senescence is further amplified among badgers putatively infected with bTB. These results recommend further investigation into the ecological and behavioural processes that result in genome-wide homozygosity, and focused work on whether homozygosity is harmful or beneficial during early life-stages.

Stress causes interspecific facilitation within a compost community.

Ecological theory predicts interactions between species to become more positive under abiotic stress, while competition should prevail in more benign environments. However, experimental tests of this stress gradient hypothesis in natural microbial communities are lacking. We test this hypothesis by measuring interactions between 10 different members of a bacterial community inhabiting potting compost in the presence or absence of toxic copper stress. We found that copper stress caused significant net changes in species interaction signs, shifting the net balance towards more positive interactions. This pattern was at least in part driven by copper-sensitive isolates - that produced relatively small amounts of metal-detoxifying siderophores - benefitting from the presence of other species that produce extracellular detoxifying agents. As well as providing support for the stress gradient hypothesis, our results highlight the importance of community-wide public goods in shaping microbial community composition.

Counting Cats: The integration of expert and citizen science data for unbiased inference of population abundance.

Free-roaming animal populations are hard to count, and professional experts are a limited resource. There is vast untapped potential in the data collected by nonprofessional scientists who volunteer their time to population monitoring, but citizen science (CS) raises concerns around data quality and biases. A particular concern in abundance modeling is the presence of false positives that can occur due to misidentification of nontarget species. Here, we introduce Integrated Abundance Models (IAMs) that integrate citizen and expert data to allow robust inference of population abundance meanwhile accounting for biases caused by misidentification. We used simulation experiments to confirm that IAMs successfully remove the inflation of abundance estimates caused by false-positive detections and can provide accurate estimates of both bias and abundance. We illustrate the approach with a case study on unowned domestic cats, which are commonly confused with owned, and infer their abundance by analyzing a combination of CS data and expert data. Our case study finds that relying on CS data alone, either through simple summation or via traditional modeling approaches, can vastly inflate abundance estimates. IAMs provide an adaptable framework, increasing the opportunity for further development of the approach, tailoring to specific systems and robust use of CS data.

Migrant birds and mammals live faster than residents.

Billions of vertebrates migrate to and from their breeding grounds annually, exhibiting astonishing feats of endurance. Many such movements are energetically costly yet there is little consensus on whether or how such costs might influence schedules of survival and reproduction in migratory animals. Here we provide a global analysis of associations between migratory behaviour and vertebrate life histories. After controlling for latitudinal and evolutionary patterns, we find that migratory birds and mammals have faster paces of life than their non-migratory relatives. Among swimming and walking species, migrants tend to have larger body size, while among flying species, migrants are smaller. We discuss whether pace of life is a determinant, consequence, or adaptive outcome, of migration. Our findings have important implications for the understanding of the migratory phenomenon and will help predict the responses of bird and mammal species to environmental change.

Demographic amplification is a predictor of invasiveness among plants.

Invasive plant species threaten native biodiversity, ecosystems, agriculture, industry and human health worldwide, lending urgency to the search for predictors of plant invasiveness outside native ranges. There is much conflicting evidence about which plant characteristics best predict invasiveness. Here we use a global demographic survey for over 500 plant species to show that populations of invasive plants have better potential to recover from disturbance than non-invasives, even when measured in the native range. Invasives have high stable population growth rates in their invaded ranges, but this metric cannot be predicted based on measurements in the native ranges. Recovery from demographic disturbance is a measure of transient population amplification, linked to high levels of reproduction, and shows phylogenetic signal. Our results demonstrate that transient population dynamics and reproductive capacity can help to predict invasiveness across the plant kingdom, and should guide international policy on trade and movement of plants.

Prior Precision, Prior Accuracy, and the Estimation of Disease Prevalence Using Imperfect Diagnostic Tests.

Estimates of disease prevalence in any host population are complicated by uncertainty in the outcome of diagnostic tests on individuals. In the absence of gold standard diagnostics (tests that give neither false positives nor false negatives), Bayesian latent class inference can be applied to batteries of diagnostic tests, providing posterior estimates of the sensitivity and specificity of each test, alongside posterior estimates of disease prevalence. Here we explore the influence of precision and accuracy of prior information on the precision and accuracy of posterior estimates of these key parameters. Our simulations use three diagnostic tests, yielding eight possible diagnostic outcomes per individual. Seven degrees of freedom allow the estimation of seven parameters: sensitivity and specificity of each test, and disease prevalence. We show that prior precision begets posterior precision but only when priors are accurate. We also show that analyses without gold standard can use imprecise priors as long as they are initialised with accuracy. Imprecise priors risk the divergence of MCMC chains towards inaccurate posterior estimates, if inaccurate initial values are used. We note that inaccurate priors can yield inaccurate and imprecise inference. Bounded priors should certainly not be used unless their accuracy is well established. Inaccurate estimates of sensitivity or specificity can yield wildly inaccurate estimates of disease prevalence. Our analyses are motivated by studies of bovine tuberculosis in a wild badger population.

Inbreeding intensifies sex- and age-dependent disease in a wild mammal.

The mutation accumulation theory of senescence predicts that age-related deterioration of fitness can be exaggerated when inbreeding causes homozygosity for deleterious alleles. A vital component of fitness, in natural populations, is the incidence and progression of disease. Evidence is growing for natural links between inbreeding and ageing; between inbreeding and disease; between sex and ageing; and between sex and disease. However, there is scant evidence, to date, for links among age, disease, inbreeding and sex in a single natural population. Using ecological and epidemiological data from a long-term longitudinal field study, we show that in wild European badgers (Meles meles) exposed naturally to bovine tuberculosis (bTB), inbreeding (measured as multilocus homozygosity) intensifies a positive correlation between age and evidence of progressed infection (measured as an antibody response to bTB), but only among females. Male badgers suffer a steeper relationship between age and progressed infection than females, with no influence of inbred status. We found no link between inbreeding and the incidence of progressed infection during early life in either sex. Our findings highlight an age-related increase in the impact of inbreeding on a fitness-relevant trait (disease state) among females. This relationship is consistent with the predictions of the mutation accumulation theory of senescence, but other mechanisms could also play a role. For example, late-life declines in condition, arising through mechanisms other than mutation accumulation might have increased the magnitude of inbreeding depression in late life. Whichever mechanism causes the observed patterns, we have shown that inbreeding can influence age-dependent patterns of disease and, by extension, is likely to affect the magnitude and timing of the late-life declines in components of fitness that characterise senescence. Better understanding of sex-specific links between inbreeding, disease and ageing provides insights into population-level pathogen dynamics and could influence management strategies for wildlife reservoirs of zoonotic disease.

Social structure contains epidemics and regulates individual roles in disease transmission in a group-living mammal.

Population structure is critical to infectious disease transmission. As a result, theoretical and empirical contact network models of infectious disease spread are increasingly providing valuable insights into wildlife epidemiology. Analyzing an exceptionally detailed dataset on contact structure within a high-density population of European badgers Meles meles, we show that a modular contact network produced by spatially structured stable social groups, lead to smaller epidemics, particularly for infections with intermediate transmissibility. The key advance is that we identify considerable variation among individuals in their role in disease spread, with these new insights made possible by the detail in the badger dataset. Furthermore, the important impacts on epidemiology are found even though the modularity of the Badger network is much lower than the threshold that previous work suggested was necessary. These findings reveal the importance of stable social group structure for disease dynamics with important management implications for socially structured populations.

Ecological selection of siderophore-producing microbial taxa in response to heavy metal contamination.

Some microbial public goods can provide both individual and community-wide benefits, and are open to exploitation by non-producing species. One such example is the production of metal-detoxifying siderophores. Here, we investigate whether conflicting selection pressures on siderophore production by heavy metals - a detoxifying effect of siderophores, and exploitation of this detoxifying effect - result in a net increase or decrease. We show that the proportion of siderophore-producing taxa increases along a natural heavy metal gradient. A causal link between metal contamination and siderophore production was subsequently demonstrated in a microcosm experiment in compost, in which we observed changes in community composition towards taxa that produce relatively more siderophores following copper contamination. We confirmed the selective benefit of siderophores by showing that taxa producing large amounts of siderophore suffered less growth inhibition in toxic copper. Our results suggest that ecological selection will favour siderophore-mediated decontamination, with important consequences for potential remediation strategies.

Divergent demographic strategies of plants in variable environments.

One of the best-supported patterns in life history evolution is that organisms cope with environmental fluctuations by buffering their most important vital rates against them. This demographic buffering hypothesis is evidenced by a tendency for temporal variation in rates of survival and reproduction to correlate negatively with their contribution to fitness. Here, we show that widespread evidence for demographic buffering can be artefactual, resulting from natural relationships between the mean and variance of vital rates. Following statistical scaling, we find no significant tendency for plant life histories to be buffered demographically. Instead, some species are buffered, whereas others have labile life histories with higher temporal variation in their more important vital rates. We find phylogenetic signal in the strength and direction of variance-importance correlations, suggesting that clades of plants are prone to being either buffered or labile. Species with simple life histories are more likely to be demographically labile. Our results suggest important evolutionary nuances in how species deal with environmental fluctuations.

Blood thicker than water: kinship, disease prevalence and group size drive divergent patterns of infection risk in a social mammal.

The importance of social- and kin-structuring of populations for the transmission of wildlife disease is widely assumed but poorly described. Social structure can help dilute risks of transmission for group members, and is relatively easy to measure, but kin-association represents a further level of population sub-structure that is harder to measure, particularly when association behaviours happen underground. Here, using epidemiological and molecular genetic data from a wild, high-density population of the European badger (Meles meles), we quantify the risks of infection with Mycobacterium bovis (the causative agent of tuberculosis) in cubs. The risk declines with increasing size of its social group, but this net dilution effect conceals divergent patterns of infection risk. Cubs only enjoy reduced risk when social groups have a higher proportion of test-negative individuals. Cubs suffer higher infection risk in social groups containing resident infectious adults, and these risks are exaggerated when cubs and infectious adults are closely related. We further identify key differences in infection risk associated with resident infectious males and females. We link our results to parent-offspring interactions and other kin-biased association, but also consider the possibility that susceptibility to infection is heritable. These patterns of infection risk help to explain the observation of a herd immunity effect in badgers following low-intensity vaccination campaigns. They also reveal kinship and kin-association to be important, and often hidden, drivers of disease transmission in social mammals.

Transients drive the demographic dynamics of plant populations in variable environments.

The dynamics of structured plant populations in variable environments can be decomposed into the 'asymptotic' growth contributed by vital rates, and 'transient' growth caused by deviation from stable stage structure.We apply this framework to a large, global data base of longitudinal studies of projection matrix models for plant populations. We ask, what is the relative contribution of transient boom and bust to the dynamic trajectories of plant populations in stochastic environments? Is this contribution patterned by phylogeny, growth form or the number of life stages per population and per species?We show that transients contribute nearly 50% or more to the resulting trajectories, depending on whether transient and stable contributions are partitioned according to their absolute or net contribution to population dynamics.Both transient contributions and asymptotic contributions are influenced heavily by the number of life stages modelled. We discuss whether the drivers of transients should be considered real ecological phenomena, or artefacts of study design and modelling strategy. We find no evidence for phylogenetic signal in the contribution of transients to stochastic growth, nor clear patterns related to growth form. We find a surprising tendency for plant populations to boom rather than bust in response to temporal changes in vital rates and that stochastic growth rates increase with increasing tendency to boom. Synthesis. Transient dynamics contribute significantly to stochastic population dynamics but are often overlooked in ecological and evolutionary studies that employ stochastic analyses. Better understanding of transient responses to fluctuating population structure will yield better management strategies for plant populations, and better grasp of evolutionary dynamics in the real world.

Demographic buffering and compensatory recruitment promotes the persistence of disease in a wildlife population.

Demographic buffering allows populations to persist by compensating for fluctuations in vital rates, including disease-induced mortality. Using long-term data on a badger (Meles meles Linnaeus, 1758) population naturally infected with Mycobacterium bovis, we built an integrated population model to quantify impacts of disease, density and environmental drivers on survival and recruitment. Badgers exhibit a slow life-history strategy, having high rates of adult survival with low variance, and low but variable rates of recruitment. Recruitment exhibited strong negative density-dependence, but was not influenced by disease, while adult survival was density independent but declined with increasing prevalence of diseased individuals. Given that reproductive success is not depressed by disease prevalence, density-dependent recruitment of cubs is likely to compensate for disease-induced mortality. This combination of slow life history and compensatory recruitment promotes the persistence of a naturally infected badger population and helps to explain the badger's role as a persistent reservoir of M. bovis.

Robust set-point regulation for ecological models with multiple management goals.

Population managers will often have to deal with problems of meeting multiple goals, for example, keeping at specific levels both the total population and population abundances in given stage-classes of a stratified population. In control engineering, such set-point regulation problems are commonly tackled using multi-input, multi-output proportional and integral (PI) feedback controllers. Building on our recent results for population management with single goals, we develop a PI control approach in a context of multi-objective population management. We show that robust set-point regulation is achieved by using a modified PI controller with saturation and anti-windup elements, both described in the paper, and illustrate the theory with examples. Our results apply more generally to linear control systems with positive state variables, including a class of infinite-dimensional systems, and thus have broader appeal.

Aphids Pick Their Poison: Selective Sequestration of Plant Chemicals Affects Host Plant Use in a Specialist Herbivore.

In some plant-insect interactions, specialist herbivores exploit the chemical defenses of their food plant to their own advantage. Brassica plants produce glucosinolates that are broken down into defensive toxins when tissue is damaged, but the specialist aphid, Brevicoryne brassicae, uses these chemicals against its own natural enemies by becoming a "walking mustard-oil bomb". Analysis of glucosinolate concentrations in plant tissue and associated aphid colonies reveals that not only do aphids sequester glucosinolates, but they do so selectively. Aphids specifically accumulate sinigrin to high concentrations while preferentially excreting a structurally similar glucosinolate, progoitrin. Surveys of aphid infestation in wild populations of Brassica oleracea show that this pattern of sequestration and excretion maps onto host plant use. The probability of aphid infestation decreases with increasing concentrations of progoitrin in plants. Brassica brassicae, therefore, appear to select among food plants according to plant secondary metabolite profiles, and selectively store only some compounds that are used against their own enemies. The results demonstrate chemical and behavioral mechanisms that help to explain evidence of geographic patterns and evolutionary dynamics in Brassica-aphid interactions.

Reconciling actual and perceived rates of predation by domestic cats.

The predation of wildlife by domestic cats (Felis catus) is a complex problem: Cats are popular companion animals in modern society but are also acknowledged predators of birds, herpetofauna, invertebrates, and small mammals. A comprehensive understanding of this conservation issue demands an understanding of both the ecological consequence of owning a domestic cat and the attitudes of cat owners. Here, we determine whether cat owners are aware of the predatory behavior of their cats, using data collected from 86 cats in two UK villages. We examine whether the amount of prey their cat returns influences the attitudes of 45 cat owners toward the broader issue of domestic cat predation. We also contribute to the wider understanding of physiological, spatial, and behavioral drivers of prey returns among cats. We find an association between actual prey returns and owner predictions at the coarse scale of predatory/nonpredatory behavior, but no correlation between the observed and predicted prey-return rates among predatory cats. Cat owners generally disagreed with the statement that cats are harmful to wildlife, and disfavored all mitigation options apart from neutering. These attitudes were uncorrelated with the predatory behavior of their cats. Cat owners failed to perceive the magnitude of their cats' impacts on wildlife and were not influenced by ecological information. Management options for the mitigation of cat predation appear unlikely to work if they focus on "predation awareness" campaigns or restrictions of cat freedom.