Climate-related range shifts in Arctic-breeding shorebirds.

Aim: To test whether the occupancy of shorebirds has changed in the eastern Canadian Arctic, and whether these changes could indicate that shorebird distributions are shifting in response to long-term climate change. Location: Foxe Basin and Rasmussen Lowlands, Nunavut, Canada. Methods: We used a unique set of observations, made 25 years apart, using general linear models to test if there was a relationship between changes in shorebird species' occupancy and their species temperature Index, a simple version of a species climate envelope. Results: Changes in occupancy and density varied widely across species, with some increasing and some decreasing. This is despite that overall population trends are known to be negative for all of these species based on surveys during migration. The changes in occupancy that we observed were positively related to the species temperature index, such that the warmer-breeding species appear to be moving into these regions, while colder-breeding species appear to be shifting out of the regions, likely northward. Main Conclusions: Our results suggest that we should be concerned about declining breeding habitat availability for bird species whose current breeding ranges are centered on higher and colder latitudes.

Impact of biological nitrogen fixation and livestock management on the manure transfer from grazing land in mixed farming systems.

Soil fertility in mixed farming systems relies on the manure produced by livestock and its recycling in the entire system. In the particular case of crop-livestock system with grazing area, the proper functioning of the system also depends on the presence of nitrogen-fixing plants in the area where livestock grazes (the grazing land). In this paper, we study the impact of biological nitrogen fixation (BNF) and livestock management on the flux of manure exported outside the grazing land. We address this issue using a modeling approach. We consider a plant-soil model composed of a set of nonlinear ordinary differential equations that represents the grazing land. We assume that the manure produced by the grazing livestock can be partially exported as a fertilizer outside of this area. Through the mathematical analysis of the model, and analytical and numerical optimization, we then determine the optimal livestock management in terms of grazing rate and manure recycling percentage that lead to the maximal flux of exported manure. We focus more precisely on the role of nitrogen-fixing plants and their impact on the optimal livestock management. When grazing rate is high and the capacity of plants to fix nitrogen is important, we showed that it is necessary to recycle some of the manure produced by the livestock in the grazing land to maximize the flux of exported manure. On the contrary, if we can optimize both the grazing rate and the manure recycling percentage, then it is better to transfer all the produced manure and to adapt the grazing rate accordingly to minimize nitrogen losses from the soil. Finally, to maximize the flux of exported manure, it is also necessary to bring the system to a state in which the plants fix nitrogen. In this way, we can benefit from the nitrogen fixation which provides an additional input of nitrogen in the system.

A consumer-driven recycling theory for the impact of large herbivores on terrestrial ecosystem stoichiometry.

Biological control of nutrient cycles is well documented in aquatic ecosystems, where consumer-driven recycling by herbivores can significantly impact ecosystem stoichiometry. In contrast, little is known in terrestrial ecosystems, where there is evidence that herbivores can also impact ecosystem stoichiometry. I studied a stoichiometric model of the soil-plant-herbivore system. The model shows that herbivores influence the ecosystem stoichiometry mainly through the direct and indirect controls of ecosystem inputs and losses, in a more complex way than predicted by the classic consumer-driven recycling theory. Overall, it shows that herbivores affect nutrient ratios in terrestrial ecosystems mostly independently of their own stoichiometric ratios, and that their impact may be different in forest versus grassland. The results highlight the sensitivity of terrestrial ecosystems to elusive actors, negligible in biomass but capable of modifying nutrient loss rates with major impacts on nutrient cycles and ecosystem stoichiometry.

Maximization of fertility transfers from rangeland to cropland: The contribution of control theory.

In traditional mixed farming systems, soil fertility in cropland relies on the transfer of fertility from rangeland through the transfer of manure produced by livestock that grazes in rangeland. In this work, we introduce a simple meta-ecosystem model in which the mixed farming system is represented by a cropland sub-system connected to a rangeland sub-system by nutrient fluxes. The livestock plays the role of nutrient-pump from the rangeland sub-system to the cropland sub-system. We use this model to study how spatial organization and practices of livestock management such as the control of grazing pressure and night corralling can help optimize both nutrient transfers and crop production. We argue that addressing the optimization of crop production requires different methods, depending on whether the agricultural practice in focus is constant or variable over time. We first used classical optimization methods at equilibrium to address optimization when the grazing pressure was assumed to be constant over time. Second, we address optimization for a more realistic configuration of our model, where grazing pressure was assumed to vary over the course of a year. In this case, we used methods developed in the field of the control theory. Classical methods showed the existence of an optimal level of constant grazing pressure that maximizes the transfers from rangeland to cropland, leading to the maximization of crop production. Control methods showed that by varying the grazing pressure adequately an additional gain of production is possible, with higher crop production and lower nutrient transfer from rangeland to cropland. This additional gain arises from the fact that the requirement of nutrient by crops is variable along the year. Consequently, a constant adjustment of the grazing pressure allows a better match between nutrient transfer and nutrient requirement over time, leading to a substantial gain of crop biomass. Our results provide new insights for a "smarter" management of fertility transfers leading to higher crop production with less rangeland surface.

Plant Strategies along Resource Gradients.

Plants present a variety of defensive strategies against herbivores, broadly classified into tolerance and resistance. Since resource availability can also limit plant growth, we expect plant allocation to resource acquisition and defense to vary along resource gradients. Yet, the conditions under which one defensive strategy is favored over the other are unclear. Here, we use an eco-evolutionary model to investigate plant adaptive allocation to resource acquisition, tolerance, and resistance along a resource gradient in a simple food web module inspired by plankton communities where plants compete for a single resource and are grazed on by a shared herbivore. We show that undefended, acquisition-specialist strategies dominate under low resource supplies. Conversely, high resource supplies, which lead to high herbivore abundance because of trophic transfers, result in either the dominance of very resistant strategies or coexistence between a completely resistant strategy and a fast-growing, tolerant one. We also explore the consequences of this adaptive allocation on species biomasses. Finally, we compare our predictions to a more traditional, density-independent optimization model. We show that density dependence mediated by resources and herbivores is the cause of the increase in plant resistance along the resource gradient, as the optimization model would instead have favored tolerance.

Avian Influenza Virus Surveillance in High Arctic Breeding Geese, Greenland.

The connectedness in Arctic regions between migratory waterbird populations originating from different continents and the potential for virus exchange at their shared Arctic breeding ground point to the need to explore the largely unstudied circumpolar circulation of avian influenza viruses (AIV). We here report the investigation of AIV in wild birds and lakes in a high Arctic area of Northeast Greenland. No AIV could be detected in the fecal, feather, and water samples collected from large flocks of pink-footed geese Anser brachyrhynchus and barnacle geese Branta leucopsis in and around refuge lakes, where they congregate at high density during their flightless molting period in summer.

Facilitation- vs. competition-driven succession: the key role of resource-ratio.

Symbiotic nitrogen (N)-fixing plants are abundant during primary succession, as typical bedrocks lack available N. In turn, fixed N accumulates in soils through biomass turnover and recycling, favouring more nitrophilous organisms. Yet, it is unclear how this facilitation mechanism interacts with competition for other limiting nutrients such as phosphorus (P) and how this affects succession. Here, we introduce a resource-explicit, community assembly model of N-fixing species and analyze successional trajectories along resource availability gradients using contemporary niche theory. We show that facilitation-driven succession occurs under low N and high enough P availabilities, and is characterised by autogenic ecosystem development and relatively ordered trajectories. We show that late facilitation-driven succession is sensitive to catastrophic shifts, highlighting the need to invoke other mechanisms to explain ecosystem stability near the climax. Put together with competition-driven succession, these results lead to an enriched version of Tilman's resource-ratio theory of succession.

Size evolution in microorganisms masks trade-offs predicted by the growth rate hypothesis.

Adaptation to local resource availability depends on responses in growth rate and nutrient acquisition. The growth rate hypothesis (GRH) suggests that growing fast should impair competitive abilities for phosphorus and nitrogen due to high demand for biosynthesis. However, in microorganisms, size influences both growth and uptake rates, which may mask trade-offs and instead generate a positive relationship between these traits (size hypothesis, SH). Here, we evolved a gradient of maximum growth rate (µmax) from a single bacterium ancestor to test the relationship among µmax, competitive ability for nutrients and cell size, while controlling for evolutionary history. We found a strong positive correlation between µmax and competitive ability for phosphorus, associated with a trade-off between µmax and cell size: strains selected for high µmax were smaller and better competitors for phosphorus. Our results strongly support the SH, while the trade-offs expected under GRH were not apparent. Beyond plasticity, unicellular populations can respond rapidly to selection pressure through joint evolution of their size and maximum growth rate. Our study stresses that physiological links between these traits tightly shape the evolution of competitive strategies.

Geometrical envelopes: Extending graphical contemporary niche theory to communities and eco-evolutionary dynamics.

Contemporary niche theory is a powerful structuring framework in theoretical ecology. First developed in the context of resource competition, it has been extended to encompass other types of regulating factors such as shared predators, parasites or inhibitors. A central component of contemporary niche theory is a graphical approach popularized by Tilman that illustrates the different outcomes of competition along environmental gradients, like coexistence and competitive exclusion. These food web modules have been used to address species sorting in community ecology, as well as adaptation and coexistence on eco-evolutionary time scales in adaptive dynamics. Yet, the associated graphical approach has been underused so far in the evolutionary context. In this paper, we provide a rigorous approach to extend this graphical method to a continuum of interacting strategies, using the geometrical concept of the envelope. Not only does this approach provide community and eco-evolutionary bifurcation diagrams along environmental gradients, it also sheds light on the similarities and differences between those two perspectives. Adaptive dynamics naturally merges with this ecological framework, with a close correspondence between the classification of singular strategies and the geometrical properties of the envelope. Finally, this approach provides an integrative tool to study adaptation between levels of organization, from the individual to the ecosystem.

Unifying sources and sinks in ecology and Earth sciences.

The paired source and sink concepts are used increasingly in ecology and Earth sciences, but they have evolved in divergent directions, hampering communication across disciplines. We propose a conceptual framework that unifies existing definitions, and review their most significant consequences for the various disciplines. A general definition of the source and sink concepts that transcends disciplines is based on net flows between the components of a system: a source is a subsystem that is a net exporter of some living or non-living entities of interest, and a sink is a net importer of these entities. Sources and sinks can further be classified as conditional and unconditional, depending on the intrinsic propensity of subsystems to either produce (source) or absorb (sink) a surplus of these entities under some (conditional) or all (unconditional) conditions. The distinction between conditional and unconditional sources and sinks, however, is strongly context dependent. Sources can turn into sinks, and vice versa, when the context is changed, when systems are subject to temporal fluctuations or evolution, or when they are considered at different spatial and temporal scales. The conservation of ecosystem services requires careful consideration of the source-sink dynamics of multiple ecosystem components. Our synthesis shows that source-sink dynamics has profound consequences for our ability to understand, predict, and manage species and ecosystems in heterogeneous landscapes.

Landscape fragmentation generates spatial variation of diet composition and quality in a generalist herbivore.

Forest fragmentation may benefit generalist herbivores by increasing access to various substitutable food resources, with potential consequences for their population dynamics. We studied a European roe deer (Capreolus capreolus) population living in an agricultural mosaic of forest, woodlots, meadows and cultivated crops. We tested whether diet composition and quality varied spatially across the landscape using botanical analyses of rumen contents and chemical analyses of the plants consumed in relation to landscape metrics. In summer and non-mast winters, roe deer ate more cultivated seeds and less native forest browse with increasing availability of crops in the local landscape. This spatial variation resulted in contrasting diet quality, with more cell content and lower lignin and hemicellulose content (high quality) for individuals living in more open habitats. The pattern was less marked in the other seasons when diet composition, but not diet quality, was only weakly related to landscape structure. In mast autumns and winters, the consumption of acorns across the entire landscape resulted in a low level of differentiation in diet composition and quality. Our results reflect the ability of generalist species, such as roe deer, to adapt to the fragmentation of their forest habitat by exhibiting a plastic feeding behavior, enabling them to use supplementary resources available in the agricultural matrix. This flexibility confers nutritional advantages to individuals with access to cultivated fields when their native food resources are depleted or decline in quality (e.g. during non-mast years) and may explain local heterogeneities in individual phenotypic quality.

Plant coexistence depends on ecosystem nutrient cycles: extension of the resource-ratio theory.

We present a model of plant-nutrient interactions that extends classical resource competition theory to environments in which essential nutrients (resources) are recycled between plant and soil pools and dissolved nutrients are lost through plant-available (i.e., inorganic forms) or plant-unavailable (i.e., complex organic forms) pathways. Losses by dissolved organic pathways can alter ratios of nutrients that are recycled and supplied within the plant-soil system, thereby influencing competition and coexistence among plant species. In special cases, our extended model does not differ from classical models, but in more realistic cases our model introduces new dynamical behavior that influences competitive outcomes. At equilibrium, coexistence still depends on nutrient supply and consumption, but nutrient supply includes recycling and is highly sensitive to whether a species promotes more organic losses of the nutrient that limits its own growth than of nutrients that limit its competitors. Because recycling operates with a time delay compared with consumption, recycling-mediated effects on competition can, under certain conditions, lead to sustained population oscillations. Our findings have implications for how we understand nutrient competition, nutrient cycles, and plant evolutionary strategies.

Optimal nitrogen-to-phosphorus stoichiometry of phytoplankton.

Redfield noted the similarity between the average nitrogen-to-phosphorus ratio in plankton (N:P = 16 by atoms) and in deep oceanic waters (N:P = 15; refs 1, 2). He argued that this was neither a coincidence, nor the result of the plankton adapting to the oceanic stoichiometry, but rather that phytoplankton adjust the N:P stoichiometry of the ocean to meet their requirements through nitrogen fixation, an idea supported by recent modelling studies. But what determines the N:P requirements of phytoplankton? Here we use a stoichiometrically explicit model of phytoplankton physiology and resource competition to derive from first principles the optimal phytoplankton stoichiometry under diverse ecological scenarios. Competitive equilibrium favours greater allocation to P-poor resource-acquisition machinery and therefore a higher N:P ratio; exponential growth favours greater allocation to P-rich assembly machinery and therefore a lower N:P ratio. P-limited environments favour slightly less allocation to assembly than N-limited or light-limited environments. The model predicts that optimal N:P ratios will vary from 8.2 to 45.0, depending on the ecological conditions. Our results show that the canonical Redfield N:P ratio of 16 is not a universal biochemical optimum, but instead represents an average of species-specific N:P ratios.