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1. Anthropogenic pressures, including climate change, are causing nonlinear changes in ecosystems globally. The development of reliable early warning indicators (EWIs) to predict these changes is vital for the adaptive management of ecosystems and the protection of biodiversity, natural capital and ecosystem services. Increased variance and autocorrelation are potential early warning indicators and can be readily estimated from ecological time series. Here, we undertook a comprehensive test of the consistency between early warning indicators and nonlinear abundance change across species, trophic levels and ecosystem types. 2. We tested whether long-term abundance time series of 55 taxa (126 data sets) across multiple trophic levels in marine and freshwater ecosystems showed (i) significant nonlinear change in abundance ‘turning points’ and (ii) significant increases in variance and autocorrelation (‘early warning indicators’). For each data set, we then quantified the prevalence of three cases: true positives (early warning indicators and associated turning point), false negatives (turning point but no associated early warning indicators) and false positives (early warning indicators but no turning point). 3. True positives were rare, representing only 9% (16 of 170) of cases using variance, and 13% (19 of 152) of cases using autocorrelation. False positives were more prevalent than false negatives (53% vs. 38% for variance; 47% vs. 40% for autocorrelation). False results were found in every decade and across all trophic levels and ecosystems. 4. Time series that contained true positives were uncommon (8% for variance; 6% for autocorrelation), with all but one time series also containing false classifications. Coherence between the types of early warning indicators was generally low with 43% of time series categorized differently based on variance compared to autocorrelation. 5. Synthesis and applications. Conservation management requires effective early warnings of

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Aerodynamics results in two characteristic speeds of flying birds: the minimum power speed and the maximum range speed. The minimum power speed requires the lowest rate of energy expenditure per unit time to stay airborne and the maximum range speed maximizes air distance traveled per unit of energy consumed. Therefore, if birds aim to minimize the cost of transport under a range of wind conditions, they are predicted to fly at the maximum range speed. Furthermore, take-off is predicted to be strongly affected by wind speed and direction. To investigate the effect of wind conditions on take-off and cruising flight behavior, we equipped 14 European shags Phalacrocorax aristotelis with a back-mounted GPS logger to measure position and hence ground speed, and a neck-mounted accelerometer to record wing beat frequency and strength. Local wind conditions were recorded during the deployment period. Shags always took off into the wind regardless of their intended destination and take-off duration was correlated negatively with wind speed. We combined ground speed and direction during the cruising phase with wind speed and direction to estimate air speed and direction. Whilst ground speed was highly variable, air speed was comparatively stable, although it increased significantly during strong head winds, because of stronger wing beats. The increased air speeds in head winds suggest that birds fly at the maximum range speed, not at the minimum power speed. Our study demonstrates that European shags actively adjust their flight behavior to utilize wind power to minimize the costs of take-off and cruising flight.

1. The age of the parents at the time of offspring production can influence offspring longevity, but the underlying mechanisms remain poorly understood. The effect of parental age on offspring telomere dynamics (length and loss rate) is one mechanism that could be important in this context. 2. Parental age might influence the telomere length that offspring inherit or age-related differences in the quality of parental care could influence the rate of offspring telomere loss. However, these routes have generally not been disentangled. 3. Here, we investigated whether parental age was related to offspring telomere dynamics using parents ranging in age from 2 to 22 years old in a free-living population of a long-lived seabird, the European shag (Phalacrocorax aristotelis). By measuring the telomere length of offspring at hatching and towards the end of the post-natal growth period, we could assess whether any potential parental age effect was confined to the post-natal rearing period. 4. There was no effect of maternal or paternal age on the initial telomere length of their chicks. However, chicks produced by older mothers and fathers experienced significantly greater telomere loss during the post-natal nestling growth period. We had relatively few nests in which the ages of both parents were known, and individuals in this population mate assortatively with respect to age. Thus, we could not conclusively determine whether the parental age effect was due to maternal age, paternal age, or both; however, it appears that the effect is stronger in mothers. 5. These results demonstrate that in this species, there was no evidence that parental age was related to offspring hatching telomere length. However, telomere loss during nestling growth was reduced in the offspring of older parents. This could be due to an age-related deterioration in the quality of the environment that parents provide, or because parents that invest less in offspring rearing live to an older age.

1. The need to manage exposure to oxidative stress, which can damage macromolecules, is thought to influence the resolution of life-history trade-offs. Oxidative damage is expected to increase with age as a consequence of changes in the optimal investment in defences or repair, and/or because of senescence in antioxidant defence systems, although the pattern might differ between short and long-lived species. However, data on age-related changes in damage levels in wild populations are rare. 2. Using cross-sectional and longitudinal data collected over 3 years, we examine variation in a measure of oxidative damage exposure in known age, wild European Shags (Phalacrocorax aristotelis), a relatively long lived species. 3. The cross-sectional data showed a quadratic relationship between oxidative damage exposure and age: both relatively young and old adults had higher levels than those in middle age. In contrast, a measure of non-enzymatic antioxidant levels did not vary with age. 4. The cross-sectional increase in oxidative damage exposure in later life was consistent with longitudinal patterns observed within older birds (more than 10 years old). 5. However, the apparent decline in oxidative damage in early adulthood was not consistent with longitudinal patterns in younger birds, which showed individual variation but no consistent age-related change in the marker. This suggests that cross-sectional patterns reflect instead higher disappearance of individuals with high exposure to oxidative damage at this life stage. 6. Our data further show that oxidative damage levels are predictive of attendance at the colony in all age classes: juveniles fledging with a high damage exposure index were less likely to be resighted in the breeding colony 2 years later, and adults with high levels at the end of the breeding season had reduced return rates, irrespective of age. Since this is a species that shows high colony fidelity, this is likely to reflect mortality patterns. 7. Thes

The recent trend for journals to require open access to primary data included in publications has been embraced by many biologists, but has caused apprehension amongst researchers engaged in long-term ecological and evolutionary studies. A worldwide survey of 73 principal investigators (PIs) with long-term studies revealed positive attitudes towards sharing data with the agreement or involvement of the PI, and 93% of PIs have historically shared data. Only 8% were in favor of uncontrolled, open access to primary data while 63% expressed serious concern. We present here their viewpoint on an issue that can have non-trivial scientific consequences. We discuss potential costs of public data archiving and provide possible solutions to meet the needs of journals and researchers.

1. Ecological resilience is developing into a credible paradigm for policy development and environmental management for preserving natural capital in a rapidly changing world. However, resilience emerges from complex interactions, limiting the translation of theory into practice. 2. Main limitations include the following: (i) difficulty in quantification and detection of changes in ecological resilience, (ii) a lack of empirical evidence to support preventative or proactive management and (iii) difficulties in managing processes operating across socio-ecological systems that vary in space and time. 3. We highlight recent research with the potential to address these limitations including new and/or improved indicators of resilience and tools to assess scale as a driver of resilience. 4. Synthesis and applications. Effective resilience-based management must be adaptive in nature. To support this, we propose an operational model using resilience-based iterative management actions operating across scales.

Parasitic infection has a direct physiological cost to hosts but may also alter how hosts interact with other individuals in their environment. Such indirect effects may alter both host fitness and the fitness of other individuals in the host's social network, yet the relative impact of direct and indirect effects of infection are rarely quantified. During reproduction, a host's social environment includes family members who may be in conflict over resource allocation. In such situations, infection may alter how resources are allocated, thereby redistributing the costs of parasitism between individuals. Here, we experimentally reduce parasite burdens of parent and/or nestling European shags (Phalacrocorax aristotelis) infected with Contracaecum nematodes in a factorial design, then simultaneously measure the impact of an individual's infection on all family members. We found no direct effect of infection on parent or offspring traits but indirect effects were detected in all group members, with both immediate effects (mass change and survival) and longer-term effects (timing of parents’ subsequent breeding). Our results show that parasite infection can have a major impact on individuals other than the host, suggesting that the effect of parasites on population processes may be greater than previously thought.

Within the last 30 years, fisheries and conservation interests have become concerned that the foraging by mute swans Cygnus olor on water crowfoot degrades the habitat of invertebrates, fish and other animals. Mute swans are a native species and a natural part of the chalk river ecosystem, but the population in Britain has almost doubled since the 1970s. Some sport fisheries organizations have reported declining membership and income due to swan grazing. Thus, a conflict has arisen between conservationists and anglers on one side and those who seek to protect the swans on the other. Moreover, management of this grazing conflict is complicated not only because mute swans are popular with the public but also they are protected under both the EU Wild Birds Directive (79/409/EEC) and the Wildlife & Countryside Act 1981. However, there has been little research into swan grazing damage to plants or management options to alleviate the conflict.

Determining the physiological mechanisms underpinning life-history decisions is essential for understanding the constraints under which life-history strategies can evolve. In long-lived species, where the residual reproductive value of breeders is high, adult survival is a key contributor to lifetime reproductive success. We therefore expect that when adult survival is compromised during reproduction, mechanisms will evolve to redirect resources away from reproduction, with implications for reproductive hormones, adult body mass, nest attendance behaviour and breeding success. We investigated whether manipulating corticosterone, to simulate exposure to an environmental stressor, affected the secretion of prolactin and breeding success in the black-legged kittiwake Rissa tridactyla. We used implanted Alzet® osmotic pumps to administer corticosterone to incubating kittiwakes at a constant rate over a period of approximately eight days. Manipulated birds were compared with sham implanted birds and control birds, which had no implants. There was no significant difference in the body mass of captured individuals at the time of implantation and implant removal. Corticosterone-implanted males showed lower nest attendance during the chick rearing period compared to sham-implanted males; the opposite pattern was found in females. Corticosterone treated birds showed a marginally significant reduction in breeding success compared to sham-implanted individuals, with all failures occurring at least one week after implant removal. However, prolactin concentrations at implant removal were not significantly different from initial values. We were unable to measure the profile of change in corticosterone during the experiment. However, our results suggest a delayed effect of elevated corticosterone on breeding success rather than an immediate suppression of prolactin concentrations causing premature failure.

Data from geolocators deployed on adult Common Guillemots from a colony in southeast Scotland indicated that they normally winter in the North Sea up to 1000 km southeast of the colony. However, one bird unexpectedly moved 3000 km northeast to moult in the Barents Sea.

Quantifying the effects of extreme weather is a critical question in population ecology since climate models predict increased climate variability. Effects will vary among and within species due to exposure or susceptibility, yet few studies have considered these sources of variation simultaneously. We investigated the effects of a summer storm on the breeding success of 4 seabird species at a North Sea colony in relation to aspect, height above sea-level, distance to cliff edge and laying date. The storm lasted 8 h with gusts of >60 m s−1. In exposed plots, razorbills Alca torda had higher failure rates (28.5%) than European shags Phalacrocorax aristotelis (15.1%), black-legged kittiwakes Rissa tridactyla (15.6%) and common guillemots Uria aalge (10.4%). Conversely, failure rates in sheltered plots were negligible (shags 0.0%; kittiwakes 1.9%; no guillemot or razorbill plots in sheltered locations). Guillemots breeding closer to sea-level were more likely to fail, but cliff edge proximity did not affect failure rate. In razorbills, pairs that laid early were more likely to survive the storm. In all species, some failed pairs re-laid, and success of re-lays was lower than that of pairs that survived. Thus, re-laying only provided partial compensation, and, overall, the storm caused a net reduction in annual population production of 4.6, 10.7, 8.9 and 22.8% for shags, kittiwakes, guillemots and razorbills, respectively. Increased storm frequency may therefore have important consequences on seabird populations, but orientation of storms relative to colonies and timing in relation to the breeding season are likely to be critical in determining the overall effect.

Parasites play key ecological and evolutionary roles through the costs they impose on their host. In wild populations, the effect of parasitism is likely to vary considerably with environmental conditions, which may affect the availability of resources to hosts for defense. However, the interaction between parasitism and prevailing conditions is rarely quantified. In addition to environmental variation acting on hosts, individuals are likely to vary in their response to parasitism, and the combined effect of both may increase heterogeneity in host responses. Offspring hierarchies, established by parents in response to uncertain rearing conditions, may be an important source of variation between individuals. Here, we use experimental antiparasite treatment across 5 years of variable conditions to test how annual population productivity (a proxy for environmental conditions) and parasitism interact to affect growth and survival of different brood members in juvenile European shags (Phalacrocorax aristotelis). In control broods, last-hatched chicks had more plastic growth rates, growing faster in more productive years. Older siblings grew at a similar rate in all years. Treatment removed the effect of environment on last-hatched chicks, such that all siblings in treated broods grew at a similar rate across environmental conditions. There were no differences in nematode burden between years or siblings, suggesting that variation in responses arose from intrinsic differences between chicks. Whole-brood growth rate was not affected by treatment, indicating that within-brood differences were driven by a change in resource allocation between siblings rather than a change in overall parental provisioning. We show that gastrointestinal parasites can be a key component of offspring's developmental environment. Our results also demonstrate the value of considering prevailing conditions for our understanding of parasite effects on host life-history traits. Establishing how envir

Effective wildlife management is needed for conservation, economic and human well-being objectives. However, traditional population control methods are frequently ineffective, unpopular with stakeholders, may affect non-target species, and can be both expensive and impractical to implement. New methods which address these issues and offer effective wildlife management are required. We used an individual-based model to predict the efficacy of a sacrificial feeding area in preventing grazing damage by mute swans (Cygnus olor) to adjacent river vegetation of high conservation and economic value. The accuracy of model predictions was assessed by a comparison with observed field data, whilst prediction robustness was evaluated using a sensitivity analysis. We used repeated simulations to evaluate how the efficacy of the sacrificial feeding area was regulated by (i) food quantity, (ii) food quality, and (iii) the functional response of the forager. Our model gave accurate predictions of aquatic plant biomass, carrying capacity, swan mortality, swan foraging effort, and river use. Our model predicted that increased sacrificial feeding area food quantity and quality would prevent the depletion of aquatic plant biomass by swans. When the functional response for vegetation in the sacrificial feeding area was increased, the food quantity and quality in the sacrificial feeding area required to protect adjacent aquatic plants were reduced. Our study demonstrates how the insights of behavioural ecology can be used to inform wildlife management. The principles that underpin our model predictions are likely to be valid across a range of different resource-consumer interactions, emphasising the generality of our approach to the evaluation of strategies for resolving wildlife management problems.

In partially migratory populations, individuals from a single breeding area experience a range of environments during the non-breeding season. If individuals show high within- and among- year fidelity to specific locations, any annual environmental effect on individual life histories could be reinforced, causing substantial demographic heterogeneity. Quantifying within- and among- individual variation and repeatability in non-breeding season location is therefore key to predicting broad-scale environmental impacts on the dynamics of partially migratory populations. We used field resightings of colour-ringed adult European shags known to have bred on the Isle of May, Scotland, to quantify individual variation and repeatability in winter location within and among three consecutive winters. In total, 3797 resightings of 882 individuals were recorded over 622km of coastline, including the Isle of May. These individuals comprised over 50% of the known breeding population, and encompassed representative distributions of ages and sexes. The distances from the Isle of May at which individuals were resighted during winter varied substantially, up to 486km and 136km north and south respectively and including the breeding colony on the Isle of May. However, resighting distances were highly repeatable within individuals; within- and among-winter repeatabilities were >0.72 and >0.59 respectively across the full September-March observation period, and >0.95 and >0.79 respectively across more restricted mid-winter periods. Repeatability did not differ significantly between males and females or among different age classes, either within or among winters. These data demonstrate that the focal shag population is partially migratory, and moreover that individuals show highly repeatable variation in winter location and hence migration strategy across consecutive winters. Such high among-individual variation and within-individual repeatability, both within and among winters, could lead

Carry-over effects have major implications for individual fitness and population and evolutionary dynamics. The strength of these effects is dependent on an individual's intrinsic performance and the environmental conditions it experiences. However, understanding the relative importance of environmental and intrinsic effects underpinning seasonal interactions has proved extremely challenging, since they co-vary. A powerful approach is longitudinal measurement of individuals across a range of conditions whereby each animal is effectively acting as its own control. We related time spent foraging during the nonbreeding period to subsequent breeding performance in European shags Phalacrocorax aristotelis. By following individuals for up to six years, we could test simultaneously for extrinsic and intrinsic effects using random regression modelling. We detected significant annual and among-individual variation in daily foraging time during the late winter, and clear variation among individuals in the quadratic relationship between foraging time and date. Shorter foraging times were associated with earlier and more successful breeding, driven by differences among years and individuals, with no evidence of individual variation in the slope of these relationships. That both environmental and intrinsic variation shape carry-over effects has important implications for population responses to environmental change.

Exposure to stressors early in life is associated with faster ageing and reduced longevity. One important mechanism that could underlie these late life effects is increased telomere loss. Telomere length in early post-natal life is an important predictor of subsequent lifespan, but the factors underpinning its variability are poorly understood. Recent human studies have linked stress exposure to increased telomere loss. These studies have of necessity been non-experimental and are consequently subjected to several confounding factors; also, being based on leucocyte populations, where cell composition is variable and some telomere restoration can occur, the extent to which these effects extend beyond the immune system has been questioned. In this study, we experimentally manipulated stress exposure early in post-natal life in nestling European shags (Phalacrocorax aristotelis) in the wild and examined the effect on telomere length in erythrocytes. Our results show that greater stress exposure during early post-natal life increases telomere loss at this life-history stage, and that such an effect is not confined to immune cells. The delayed effects of increased telomere attrition in early life could therefore give rise to a ‘time bomb’ that reduces longevity in the absence of any obvious phenotypic consequences early in life.

Marine spatial planning aims to deliver sustainable use of marine resources by minimizing environmental impacts of human activities and designating Marine Protected Areas. This poses a challenge where species’ distributions show spatio-temporal heterogeneity. However, due to logistic constraints and challenging timescales many studies of distribution are undertaken over few years or on a restricted subset of the population. Long-term studies can help identify the degree of uncertainty in those less comprehensive in space and time. We quantify inter-annual and sub-colony variation in the summer foraging distribution of a population of European shags Phalacrocorax aristotelis, using a tracking data set comprising 320 individuals and 1106 foraging trips in 15 years from 1987 to 2010. Foraging distribution over the study period was concentrated in three areas. Data from one and two years captured an average of 54% and 64% of this distribution, respectively, but it required 8 years’ data to capture over 90% of the distribution. Foraging range increased with population size when breeding success was low, suggesting interplay between extrinsic and intrinsic effects. Furthermore, females had foraging ranges on average 36% greater than males. Finally, sub-colony segregation occurred in foraging areas up to 4 km from the colony and in the most distant locations (>10 km), whilst there was considerable overlap at intermediate distances (6–10 km). Our study highlights important considerations for marine spatial planning in particular, and species conservation in general, notably the proportion of the population distribution identified, the prevailing conditions experienced and the need for balanced sampling across the population.

The evidence shows that swan grazing can reduce plant abundance, prevent flowering, reduce water depth and reduce fishery value. However, these effects seem to be limited to a small number of sites on larger chalk streams. The results of attempted management have been disappointing, and we currently have no simple effective means of preventing grazing damage. However, our understanding of the effects of swans on the chalk stream ecosystem has been growing rapidly, which gives us hope for future solutions. In particular, combining strategies which improve river condition and move swans away from sensitive areas could offer a way of managing grazing effects.

Understanding how prey capture rates are influenced by feeding ecology and environmental conditions is fundamental to assessing anthropogenic impacts on marine higher predators. We compared how prey capture rates varied in relation to prey size, prey patch distribution and prey density for two species of alcid, common guillemot (Uria aalge) and razorbill (Alca torda) during the chick-rearing period. We developed a Monte Carlo approach parameterised with foraging behaviour from birdborne data loggers, observations of prey fed to chicks, and adult diet from wateroffloading, to construct a bio-energetics model. Our primary goal was to estimate prey capture rates, and a secondary aim was to test responses to a set of biologically plausible environmental scenarios. Estimated prey capture rates were 1.5±0.8 items per dive (0.8±0.4 and 1.1±0.6 items per minute foraging and underwater, respectively) for guillemots and 3.7±2.4 items per dive (4.9±3.1 and 7.3±4.0 items per minute foraging and underwater, respectively) for razorbills. Based on species' ecology, diet and flight costs, we predicted that razorbills would be more sensitive to decreases in 0- group sandeel (Ammodytes marinus) length (prediction 1), but guillemots would be more sensitive to prey patches that were more widely spaced (prediction 2), and lower in prey density (prediction 3). Estimated prey capture rates increased non-linearly as 0- group sandeel length declined, with the slope being steeper in razorbills, supporting prediction 1. When prey patches were more dispersed, estimated daily energy expenditure increased by a factor of 3.0 for guillemots and 2.3 for razorbills, suggesting guillemots were more sensitive to patchier prey, supporting prediction 2. However, both species responded similarly to reduced prey density (guillemot expenditure increased by 1.7; razorbill by 1.6), thus not supporting prediction 3. This bio-energetics approach complements other foraging models in predicting likely impacts of

Capsule: The Mute Swan, a large generalist herbivore, showed patterns of habitat use influenced by social grouping and season. Territorial swans showed strong preferences for river and lake habitat in all seasons, while the non-territorial birds known to cause grazing conflicts preferred river in summer–autumn and pasture in winter–spring. Aims: To quantify the habitat preferences across different seasons of two types of Mute Swan social group, territorial and non-territorial, and assess the implications for the grazing conflict between swans and valuable plant communities. Methods: Repeated surveys of the River Frome catchment, Dorset, UK, over a two-year period allowed us to record the numbers of swans in each habitat type. An electivity index was used to calculate habitat preference scores for territorial and non-territorial swans across different seasons. Results: We found strong seasonal switches in habitat use for both territorial and non-territorial swans, but preferences for some habitats differed between these social groupings. In particular, non-territorial swans preferred pasture in winter and spring, and river in summer and autumn, while territorial swans preferred river and lake habitats throughout the year. Conclusion: The Mute Swan, a large generalist herbivore that can cause grazing damage to river plant communities, showed patterns of habitat use influenced by social grouping and season. These seasonal patterns of habitat use suggest that the grazing conflict with the river plant community caused by flocks of non-territorial swans is currently limited to summer and autumn.

1. Parasites are considered to be a key driver of evolutionary processes in wild animal populations. However, assessing host-endoparasite burdens non-destructively is problematic. Collection of faecal samples can be difficult, and faecal egg counts may not always be a reliable indicator of infection intensity. 2. Here we report on endoscopy as a method for assessing natural burdens of nematode parasites Contracaecum rudolphii Hartwich, 1964, in a wild seabird, the European shag (Phalacrocorax aristotelis, L.). We aimed to measure natural individual parasite burdens and repeatability of burdens over time, and to verify that treatment with ivermectin removed parasites. 3. Endoscopy was rapid, averaging 6 min (n = 157), with no obvious adverse effects on behaviour or breeding success compared to non-endoscoped birds. 4. Nematode burdens in the oesophagus and proventriculus of conscious shags were counted and classified as absent, low, medium or high using a flexible gastroscope with a camera attachment that recorded video footage. 5. Re-assessment of worm burdens was highly accurate, with 94% of randomly selected videos (n = 50) giving identical categorical scores, and 70% of worm counts (n = 40) giving the same total or differing by only one worm. 6. All birds were parasitised by C. rudolphii. Natural burdens were significantly higher in males and in late breeders. 7. Individuals had highly repeatable categorical parasite scores over time with 65% of control birds sampled more than once (n = 17; mean interval between assessments= 10·8 days) showing no change in scores. However, although the rank ordering of birds based on categorical scores remained constant, more finely resolved quantification indicated a slight seasonal decline in worm counts within individuals. 8. Treatment with ivermectin (4 mg kg−1 of bird weight) resulted in complete removal of parasites. There was some evidence of temporal declines in worm counts with lower doses of ivermectin, including a dose

Abundant herbivores can damage plants and so cause conflict with conservation, agricultural, and fisheries interests. Management of herbivore populations is a potential tool to alleviate such conflicts but may raise concerns about the economic and ethical costs of implementation, especially if the herbivores are ‘charismatic’ and popular with the public. Thus it is critical to evaluate the probability of achieving the desired ecological outcomes before proceeding to a field trial. Here we assessed the potential for population control to resolve a conflict of non-breeding swans grazing in river catchments. We used a mathematical model to evaluate the consequences of three population management strategies; (a) reductions in reproductive success, (b) removal of individuals, and (c) reduced reproductive success and removal of individuals combined. This model gave accurate projections of historical changes in population size for the two rivers for which data were available. Our model projected that the River Frome swan population would increase by 54 %, from 257 to 397 individuals, over 17 years in the absence of population control. Removal of ≥ 60 % of non-breeding individuals each year was projected to reduce the catchment population below the level for which grazing conflicts have been previously reported. Reducing reproductive success, even to 0 eggs per nest, failed to achieve the population reduction required. High adult and juvenile survival probabilities (> 0.7) and immigration from outside of the catchment limited the effects of management on population size. Given the high, sustained effort required, population control does not represent an effective management option for preventing the grazing conflicts in river catchments. Our study highlights the need to evaluate the effects of different management techniques, both alone and in combination, prior to field trials. Population models, such as the one presented here, can provide a cost-effective and ethical mean

Existing descriptions of bi-directional ammonia (NH3) land–atmosphere exchange incorporate temperature and moisture controls, and are beginning to be used in regional chemical transport models. However, such models have typically applied simpler emission factors to upscale the main NH3 emission terms. While this approach has successfully simulated the main spatial patterns on local to global scales, it fails to address the environment- and climate-dependence of emissions. To handle these issues, we outline the basis for a new modelling paradigm where both NH3 emissions and deposition are calculated online according to diurnal, seasonal and spatial differences in meteorology. We show how measurements reveal a strong, but complex pattern of climatic dependence, which is increasingly being characterized using ground-based NH3 monitoring and satellite observations, while advances in process-based modelling are illustrated for agricultural and natural sources, including a global application for seabird colonies. A future architecture for NH3 emission–deposition modelling is proposed that integrates the spatio-temporal interactions, and provides the necessary foundation to assess the consequences of climate change. Based on available measurements, a first empirical estimate suggests that 5°C warming would increase emissions by 42 per cent (28–67%). Together with increased anthropogenic activity, global NH3 emissions may increase from 65 (45–85) Tg N in 2008 to reach 132 (89–179) Tg by 2100.