A theory for context-dependent effects of mammalian trampling on ecosystem nitrogen cycling.

Large mammalian herbivores substantially impact ecosystem functioning. As their populations are dramatically altered globally, disentangling their consumptive and non-consumptive effects is critical to advance mechanistic understanding and improve prediction of effects over ecosystem and Earth-system spatial extents. Mathematical models have played an important role in clarifying potential mechanisms of herbivore zoogeochemistry, based mostly on their consumptive effects as primary consumers and recyclers of organic and inorganic matter via defecation and urination. Trampling is a ubiquitous effect among walking vertebrates, but the consequences and potential mechanisms of trampling in diverse environments remain poorly understood. We derive a novel mathematical model of large mammalian herbivore effects on ecosystem nitrogen cycling, focusing on how trampling and environmental context impact soil processes. We model herbivore trampling with a linear positive or negative additive effect on soil-mediated nitrogen cycling processes. Combining analytical and numerical analyses, we find trampling by large mammalian herbivores is likely to decrease nitrogen mineralisation rate across diverse environments, such as temperate grassland and boreal forest. These effects are mediated by multiple potential mechanisms, including trampling-induced changes to detritivore biomass and functioning (e.g. rate of organic matter consumption). We also uncover scenarios where trampling can increase nitrogen mineralisation rate, contingent on the environment-specific relative sensitivity of detritivore mineral-nitrogen release and detritivore mortality, to trampling. In contrast to some consumptive mechanisms, our results suggest the pace of soil nitrogen cycling prior to trampling has little influence over the direction of the trampling net effect on nitrogen mineralisation, but that net effects may be greater in slow-cycling systems (e.g. boreal forests) than in fast-cycling systems (e.g. grasslands). Our model clarifies the potential consequences of previously overlooked mechanisms of zoogeochemistry that are common to all terrestrial biomes. Our results provide empirically testable predictions to guide future progress in empirical and theoretical studies of herbivore effects in diverse environmental contexts. Resolving ecological contingencies around animal consumptive and non-consumptive effects will improve whole-ecosystem management efforts such as restoration and rewilding.

Towards a mechanistic understanding of animal-ecosystem interactions.

Research Highlight: Ferraro, K. M., Welker, L., Ward, E. B., Schmitz, O. J., & Bradford, M. A. (2023). Plant mycorrhizal associations mediate the zoogeochemical effects of calving subsidies by a forest ungulate. Journal of Animal Ecology, https://doi.org/10.1111/1365-2656.14002. Animals play large roles in ecosystem elemental cycling but predicting effects in diverse contexts remains a substantial challenge. Fundamental to progress is (1) identifying mechanisms by which animals impact nutrient distribution and cycling, and (2) disentangling how environmental context mediates the operation of alternative mechanisms. In an elegant field experiment, Ferraro et al. (2023) provide the first detailed exploration of the impact of nutrient inputs from mammalian parturition on soil functioning and the stoichiometry of plant tissues. The authors find that nitrogen from experimental additions of ungulate parturition material (natal fluids) is rapidly incorporated into microsite soil organic pools and plant tissues. They also find that soil processes (soil microbial biomass, rates of carbon mineralization, nitrogen mineralization and nitrification) and the nitrogen content of plant tissues above- and belowground are increased by addition of parturition material. Notably, the authors identify that increases in some soil processes and plant tissue nitrogen are weaker in microsites dominated by ericoid mycorrhizal plants than those dominated by ectomycorrhizal plants. These findings demonstrate that parturition depositions, a ubiquitous but overlooked mechanism of mammalian impacts on ecosystems, impact ecosystem processes and plant tissue stoichiometry. Furthermore, plant-fungal associations are a predictive axis of context dependency mediating zoogeochemical effects at fine scales. Ferraro et al.'s (2023) novel approach simultaneously advances mechanistic understanding of animal-ecosystem interactions at fine scales and facilitates prediction of ungulate effects on nutrient availability at landscape extents.

Strong above-ground impacts of a non-native ungulate do not cascade to impact below-ground functioning in a boreal ecosystem.

1. Experimental studies across biomes demonstrate that herbivores can have significant effects on ecosystem functioning. Herbivore effects, however, can be highly variable with studies demonstrating positive, neutral or negative relationships between herbivore presence and different components of ecosystems. Mixed effects are especially likely in the soil, where herbivore effects are largely indirect mediated through effects on plants. 2. We conducted a long-term experiment to disentangle the effects of non-native moose in boreal forests on plant communities, nutrient cycling, soil composition and soil organism communities. 3. To explore the effect of moose on soils, we conduct separate analyses on the soil organic and mineral horizons. Our data come from 11 paired exclosure-control plots in eastern and central Newfoundland, Canada that provide insight into 22-25 years of moose herbivory. We fit piecewise structural equations models (SEM) to data for the organic and mineral soil horizons to test different pathways linking moose to above-ground and below-ground functioning. 4. The SEMs revealed that moose exclusion had direct positive impacts on adult tree count and an indirect negative impact on shrub percent cover mediated by adult tree count. We detected no significant impact of moose on soil microbial C:N ratio or net nitrogen mineralization in the organic or mineral soil horizon. Soil temperature and moisture, however, was more than twice as variable in the presence (i.e. control) than absence (i.e. exclosure) of moose. Overall, we observed clear impacts of moose on above-ground forest components with limited indirect effects below-ground. Even after 22-25 years of exclusion, we did not find any evidence of moose impacts on soil microbial C:N ratio and net nitrogen mineralization. 5. Our long-term study and mechanistic path analysis demonstrates that soils can be resilient to ungulate herbivore effects despite evidence of strong effects above-ground. Long-term studies and analyses such as this one are relatively rare yet critical for reconciling some of the context-dependency observed across studies of ungulates effects on ecosystem functions. Such studies may be particularly valuable in ecosystems with short growing seasons such as the boreal forest.

In defense of elemental currencies: can ecological stoichiometry stand as a framework for terrestrial herbivore nutritional ecology?

Nutritional ecologists aim to predict population or landscape-level effects of food availability, but the tools to extrapolate nutrition from small to large extents are often lacking. The appropriate nutritional ecology currencies should be able to represent consumer responses to food while simultaneously be simple enough to expand such responses to large spatial extents and link them to ecosystem functioning. Ecological stoichiometry (ES), a framework of nutritional ecology, can meet these demands, but it is typically associated with ecosystem ecology and nutrient cycling, and less often used to study wildlife nutrition. Despite the emerging zoogeochemical evidence that animals, and thus their diets, play critical roles in nutrient movement, wildlife nutritional ecology has not fully embraced ES, and ES has not incorporated nutrition in many wildlife studies. Here, we discuss how elemental currencies are "nutritionally, organismally, and ecologically explicit" in the context of terrestrial herbivore nutritional ecology. We add that ES and elemental currencies offer a means to measure resource quality across landscapes and compare nutrient availability among regions. Further, we discuss ES shortcomings and solutions, and list future directions to advance the field. As ecological studies increasingly grow in spatial extent, and attempt to link multiple levels of biological organization, integrating more simple and unifying currencies into nutritional studies, like elements, is necessary for nutritional ecology to predict herbivore occurrences and abundances across regions.

Integrating plant stoichiometry and feeding experiments: state-dependent forage choice and its implications on body mass.

Intraspecific feeding choices comprise a large portion of herbivore foraging decisions. Plant resource quality is heterogeneously distributed, affected by nutrient availability and growing conditions. Herbivores navigate landscapes, foraging not only according to food qualities, but also energetic and nutritional demands. We test three non-exclusive foraging hypotheses using the snowshoe hare (Lepus americanus): (1) herbivore feeding choices and body conditions respond to intraspecific plant quality variation; (2) high energetic demands mitigate feeding responses; and (3) feeding responses are inflated when nutritional demands are high. We measured black spruce (Picea mariana) nitrogen, phosphorus and terpene compositions, as indicators of quality, within a snowshoe hare trapping grid and found plant growing conditions to explain spruce quality variation (R2 < 0.36). We then offered two qualities of spruce (H1) from the trapping grid to hares in cafeteria-style experiments and measured their feeding and body condition responses (n = 75). We proxied energetic demands (H2) with ambient temperature and coat insulation (% white coat) and nutritional demands (H3) with the spruce quality (nitrogen and phosphorus content) in home ranges. Hares with the strongest preference for high-quality spruce lost on average 2.2% less weight than hares who ate the least high-quality spruce relative to low-quality spruce. The results supported our energetic predictions as follows: hares in colder temperatures and with less-insulative coats (lower % white) consumed more spruce and were less selective towards high-quality spruce. Collectively, we found variation in plant growing conditions within herbivore home ranges substantial enough to affect herbivore body conditions, but energetic stats mediate plant-herbivore interactions.

The multiple meanings of omnivory influence empirical, modular theory and whole food web stability relationships.

The persistence of whole communities hinges on the presence of select interactions which act to stabilize communities making the identification of these keystone interactions critical. One potential candidate is omnivory, yet theoretical research on omnivory thus far has been dominated by a modular theory approach whereby an omnivore and consumer compete for a shared resource. Empirical research, however, has highlighted the presence of a broader suite of omnivory modules. Here, we integrate empirical data analysis and mathematical models to explore the influence of both omnivory module (including classic, multi-resource, higher level, mutual predation and cannibalism) and omnivore-resource interaction type on food web stability. We use six classic empirical food webs to examine the prevalence of the different types of omnivory, a multi-species consumer-resource model to determine the stability of these different kinds of omnivory within a module context, and finally extend these models to a 50 species, whole food web model to examine the influence of omnivory on whole food web persistence. Our results challenge the concept that omnivory is broadly stabilizing. In particular, we demonstrate that the impact of omnivory depends on the type of omnivory being examined with multi-resource omnivory having the largest correlation with whole food web persistence. Moreover, our results highlight that we need to exercise caution when scaling modular theory to whole food web theory. Cannibalism, for example, was the most persistent and stable omnivory module in the modular theory analysis, but only demonstrated a weak correlation with whole food web persistence. Lastly, our results demonstrate that the frequency of omnivory modules are more important for whole food web persistence than the frequency of omnivore-resource interactions. Together, these results demonstrate that the role of omnivory often depends both on the type of omnivory being examined and the food web within which it is nested. In whole food web models, omnivory acts less as a keystone interaction, rather, specific types of omnivory, particularly multi-resource omnivory, act as keystone modules. Future work integrating module and whole food web theory is critical for resolving the role of key interactions in food webs.

Forage stoichiometry predicts the home range size of a small terrestrial herbivore.

Home range size of consumers varies with food quality, but the many ways of defining food quality hamper comparisons across studies. Ecological stoichiometry studies the elemental balance of ecological processes and offers a uniquely quantitative, transferrable way to assess food quality using elemental ratios, e.g., carbon (C):nitrogen (N). Here, we test whether snowshoe hares (Lepus americanus) vary their home range size in response to spatial patterns of C:N, C:phosphorus (P), and N:P ratios of two preferred boreal forage species, lowbush blueberry (Vaccinium angustifolium) and red maple (Acer rubrum), in summer months. Boreal forests are N- and P-limited ecosystems and access to N- and P-rich forage is paramount to snowshoe hares' survival. Accordingly, we consider forage with higher C content relative to N and P to be lower quality than forage with lower relative C content. We combine elemental distribution models with summer home range size estimates to test the hypothesis that home range size will be smaller in areas with access to high, homogeneous food quality compared to areas of low, heterogeneous food quality. Our results show snowshoe hares had smaller home ranges in areas where lowbush blueberry foliage quality was higher or more spatially homogenous than in areas of lower, more heterogeneous food quality. By responding to spatial patterns of food quality, consumers may influence community and ecosystem processes by, for example, varying nutrient recycling rates. Our reductionist biogeochemical approach to viewing resources leads us to holistic insights into consumer spatial ecology.

The strength of ecological subsidies across ecosystems: a latitudinal gradient of direct and indirect impacts on food webs.

Material and energy flows among ecosystems can directly and indirectly drive ecosystem functions. Yet, how populations of consumers respond to allochthonous inputs at a macroecological scale is still unclear. Using a meta-analysis spanning several biomes, we show that the abundance of recipient populations is 36-57% larger with increased allochthonous inputs. The strength of direct effects on the recipients of these inputs as well as the indirect effects on the consumers of these recipients (i.e. ascending indirect effects) are constant across a latitudinal gradient spanning subtropical, arid, temperate, boreal and arctic ecosystems. However, indirect effect on the in situ resources of the input recipient (i.e. descending indirect effects) decreases with latitude. Our results suggest that the influence of allochthonous inputs can vary across large-scale gradients of ecosystem productivity and may be driven by the types of trophic interactions within recipient food webs.

Stoichiometric distribution models: ecological stoichiometry at the landscape extent.

Human activities are altering the fundamental geography of biogeochemicals. Yet we lack an understanding of how the spatial patterns in organismal stoichiometry affect biogeochemical processes and the tools to predict the impacts of global changes on biogeochemical processes. In this contribution we develop stoichiometric distribution models (StDMs), which allow us to map spatial structure in resource elemental composition across a landscape and evaluate spatial responses of consumers. We parameterise StDMs for a consumer-resource (moose-white birch) system and demonstrate that we can develop predictive models of resource stoichiometry across a landscape and that such models could improve our predictions of consumer space use. With results from our study system application, we argue that explicit consideration of the spatial patterns in organismal elemental composition may uncover emergent individual, population, community and ecosystem properties that are not revealed at the local extents routinely used in ecological stoichiometry. We discuss perspectives for further developments and application of StDMs to advance three emerging frameworks for spatial ecosystem ecology in an era of global change; meta-ecosystem theory, macroecological stoichiometry and remotely sensed biogeochemistry. Progress on these emerging frameworks will allow for the integration of ecological stoichiometry and individual space use and fitness.

Mechanistic models for the spatial spread of species under climate change.

Global climate change is a major threat to biodiversity. The most common methods for predicting the response of biodiversity to changing climate do not explicitly incorporate fundamental evolutionary and ecological processes that determine species responses to changing climate, such as reproduction, dispersal, and adaptation. We provide an overview of an emerging mechanistic spatial theory of species range shifts under climate change. This theoretical framework explicitly defines the ecological processes that contribute to species range shifts via biologically meaningful dispersal, reproductive, and climate envelope parameters. We present methods for estimating the parameters of the model with widely available species occurrence and abundance data and then apply these methods to empirical data for 12 North American butterfly species to illustrate the potential use of the theory for global change biology. The model predicts species persistence in light of current climate change and habitat loss. On average, we estimate that the climate envelopes of our study species are shifting north at a rate of 3.25 +/- 1.36 km/yr (mean +/- SD) and that our study species produce 3.46 +/- 1.39 (mean +/- SD) viable offspring per individual per year. Based on our parameter estimates, we are able to predict the relative risk of our 12 study species for lagging behind changing climate. This theoretical framework improves predictions of global change outcomes by facilitating the development and testing of hypotheses, providing mechanistic predictions of current and future range dynamics, and encouraging the adaptive integration of theory and data. The theory is ripe for future developments such as the incorporation of biotic interactions and evolution of adaptations to novel climatic conditions, and it has the potential to be a catalyst for the development of more effective conservation strategies to mitigate losses of biodiversity from global climate change.

Land development in and around protected areas at the wilderness frontier.

Protected areas' chief conservation objectives are to include species within their boundaries and protect them from negative external pressures. Many protected areas are not achieving these goals, perhaps in part due to land development inside and outside protected areas. We conducted spatial analyses to evaluate the ability of Canadian protected areas to mitigate the effects of nearby land development. We investigated correlations of national patterns of land development in and around protected areas and then examined national patterns of roads, urban area, and croplands in protected areas. We calculated the amount of developed land in protected areas and within 25-100 km of protected-area borders, the density of roads, and extent of urban and cropland area in protected areas. We constructed logistic-regression models to test whether development in a protected area was associated with landscape and protected-area characteristics. Land development was far less extensive inside than outside protected areas. However, several protected areas, particularly small southern areas near small urban centers had substantial development inside their boundaries, and nearly half of protected areas had roads. The cumulative extent of development within 50 km of protected areas was the best predictor of the probability of land development in protected areas. Canadian First Nations, industries, government, and nongovernmental organizations are currently planning an unprecedented number of new protected areas. Careful management of areas beyond protected-area boundaries may prove critical to meeting their long-term conservation objectives.

Cascading impacts of changes in subsidy quality on recipient ecosystem functioning.

Resource quantity and quality can differ between adjacent ecosystems, and these differences can impact subsidies exchanged between ecosystems. The quantity and quality of subsidies are rapidly changing in response to stressors associated with global environmental change, but while we have models to predict the effects of changes in subsidy quantity, we currently lack models to predict the effects of changes in subsidy quality on recipient ecosystem functioning. We developed a novel model to predict the effects of subsidy quality on recipient ecosystem biomass distribution, recycling, production, and efficiency. We parameterized the model for a case study of a riparian ecosystem subsidized by pulsed emergent aquatic insects. In this case study we focused on a common measure of subsidy quality that differs between riparian and aquatic ecosystems: the higher content of long-chain polyunsaturated fatty acids (PUFAs) in aquatic ecosystems. We analyzed how changes in the PUFA concentration of aquatic subsidies affect the dynamics in biomass stocks and functions of the riparian ecosystem. We also conducted a global sensitivity analysis to identify key drivers of subsidy impacts. Our analysis showed that subsidy quality increased the functioning of the recipient ecosystem. Recycling increased more strongly than production per unit subsidy quality increase, meaning there was a threshold where an increase in subsidy quality led to stronger effects of subsidies on recycling relative to the production of the recipient ecosystem. Our predictions were most sensitive to basal nutrient input, highlighting the relevance of recipient ecosystem nutrient levels to understanding the effects of ecosystem connections. We argue that recipient ecosystems that rely on high-quality subsidies, such as aquatic-terrestrial ecotones, are highly sensitive to changes in subsidy-recipient ecosystem connections. Our novel model unifies the subsidy hypothesis and food quality hypothesis and provides testable predictions to understand the effects of ecosystem connections on ecosystem functioning under global changes.

Predation risk, stoichiometric plasticity and ecosystem elemental cycling.

It is widely held that herbivore growth and production is limited by dietary nitrogen (N) that in turn constrains ecosystem elemental cycling. Yet, emerging evidence suggests that this conception of limitation may be incomplete, because chronic predation risk heightens herbivore metabolic rate and shifts demand from N-rich proteins to soluble carbohydrate-carbon (C). Because soluble C can be limiting, predation risk may cause ecosystem elemental cycling rates and stoichiometric balance to depend on herbivore physiological plasticity. We report on a stoichiometrically explicit ecosystem model that investigates this problem. The model tracks N, and soluble and recalcitrant C through ecosystem compartments. We evaluate how soluble plant C influences C and N stocks and flows in the presence and absence of predation risk. Without risk, herbivores are limited by N and respire excess C so that plant-soluble C has small effects only on elemental stocks and flows. With predation risk, herbivores are limited by soluble C and release excess N, so plant-soluble C critically influences ecosystem elemental stocks flows. Our results emphasize that expressing ecosystem stoichiometric balance using customary C:N ratios that do not distinguish between soluble and recalcitrant C may not adequately describe limitations on elemental cycling.

Ecoregion and community structure influences on the foliar elemental niche of balsam fir (Abies balsamea (L.) Mill.) and white birch (Betula papyrifera Marshall).

Changes in foliar elemental niche properties, defined by axes of carbon (C), nitrogen (N), and phosphorus (P) concentrations, reflect how species allocate resources under different environmental conditions. For instance, elemental niches may differ in response to large-scale latitudinal temperature and precipitation regimes that occur between ecoregions and small-scale differences in nutrient dynamics based on species co-occurrences at a community level. At a species level, we compared foliar elemental niche hypervolumes for balsam fir (Abies balsamea (L.) Mill.) and white birch (Betula papyrifera Marshall) between a northern and southern ecoregion. At a community level, we grouped our focal species using plot data into conspecific (i.e., only one focal species is present) and heterospecific groups (i.e., both focal species are present) and compared their foliar elemental concentrations under these community conditions across, within, and between these ecoregions. Between ecoregions at the species and community level, we expected niche hypervolumes to be different and driven by regional biophysical effects on foliar N and P concentrations. At the community level, we expected niche hypervolume displacement and expansion patterns for fir and birch, respectively-patterns that reflect their resource strategy. At the species level, foliar elemental niche hypervolumes between ecoregions differed significantly for fir (F = 14.591, p-value = .001) and birch (F = 75.998, p-value = .001) with higher foliar N and P in the northern ecoregion. At the community level, across ecoregions, the foliar elemental niche hypervolume of birch differed significantly between heterospecific and conspecific groups (F = 4.075, p-value = .021) but not for fir. However, both species displayed niche expansion patterns, indicated by niche hypervolume increases of 35.49% for fir and 68.92% for birch. Within the northern ecoregion, heterospecific conditions elicited niche expansion responses, indicated by niche hypervolume increases for fir of 29.04% and birch of 66.48%. In the southern ecoregion, we observed a contraction response for birch (niche hypervolume decreased by 3.66%) and no changes for fir niche hypervolume. Conspecific niche hypervolume comparisons between ecoregions yielded significant differences for fir and birch (F = 7.581, p-value = .005 and F = 8.038, p-value = .001) as did heterospecific comparisons (F = 6.943, p-value = .004, and F = 68.702, p-value = .001, respectively). Our results suggest species may exhibit biogeographical specific elemental niches-driven by biophysical differences such as those used to describe ecoregion characteristics. We also demonstrate how a species resource strategy may inform niche shift patterns in response to different community settings. Our study highlights how biogeographical differences may influence foliar elemental traits and how this may link to concepts of ecosystem and landscape functionality.

Ecological network complexity scales with area.

Larger geographical areas contain more species-an observation raised to a law in ecology. Less explored is whether biodiversity changes are accompanied by a modification of interaction networks. We use data from 32 spatial interaction networks from different ecosystems to analyse how network structure changes with area. We find that basic community structure descriptors (number of species, links and links per species) increase with area following a power law. Yet, the distribution of links per species varies little with area, indicating that the fundamental organization of interactions within networks is conserved. Our null model analyses suggest that the spatial scaling of network structure is determined by factors beyond species richness and the number of links. We demonstrate that biodiversity-area relationships can be extended from species counts to higher levels of network complexity. Therefore, the consequences of anthropogenic habitat destruction may extend from species loss to wider simplification of natural communities.

Incorporating abiotic controls on animal movements in metacommunities.

Local dynamics are influenced by regional processes. Meta-ecology, or the study of spatial flows of energy, materials, and species between local systems, is becoming increasingly concerned with accurate depictions of species movements and the impacts of this movement on landscape-level ecosystem function. Indeed, incorporating diverse types of movement is a major frontier in metacommunity theory. Here, we synthesize literature to demonstrate that the movement of organisms between patches is governed by the interplay between both a species' ability to move and the combined effects of landscape structure and physical flows (termed abiotic controls), which together we refer to as abiotic-dependent species connectivity. For example, two lakes that share geographic proximity may be inaccessible for mobile fish species because they lack a river connecting them (landscape structure), but wind currents may disperse insects between them (physical flows). Empirical evidence suggests that abiotic controls, such as ocean currents, lead to abiotic-dependent species connectivity and that, in nature, this type of connectivity is the rule rather than the exception. Based on this empirical evidence, we introduce a novel mathematical framework to demonstrate how species movement capabilities and abiotic conditions, can interact to influence metacommunity stability. We apply this framework to predict how incorporating abiotic-dependent species connectivity applies to classic empirical examples of aquatic, aquatic-terrestrial, and terrestrial experimental metacommunities. We demonstrate that incorporating abiotic-dependent species connectivity into metacommunity models can lead to a much broader range of dynamics than models previously predicted, including a wider range of metacommunity stability. Our framework fills critical gaps in our basic understanding of organismal movement across landscapes and provides testable predictions for how such common natural phenomena impact landscape-level ecosystem function. Finally, we present future perspectives for further development of meta-ecological theory from questions about fragmentation to ecosystems. Anthropogenic change is not only leading to habitat loss from the damming of rivers to denuding the landscape, but altering the physical flows that have historically connected communities. Thus, recognizing the importance of these processes in tandem with species' movement abilities is critical for predicting and preserving the structure and function of ecological communities.

Consumer-mediated recycling and cascading trophic interactions.

Cascading trophic interactions mediated by consumers are complex phenomena, which encompass many direct and indirect effects. Nonetheless, most experiments and theory on the topic focus uniquely on the indirect, positive effects of predators on producers via regulation of herbivores. Empirical research in aquatic ecosystems, however, demonstrate that the indirect, positive effects of consumer-mediated recycling on primary producer stocks may be larger than the effects of herbivore regulation, particularly when predators have access to alternative prey. We derive an ecosystem model with both recipient- and donor-controlled trophic relationships to test the conditions of four hypotheses generated from recent empirical work on the role of consumer-mediated recycling in cascading trophic interactions. Our model predicts that predator regulation of herbivores will have larger, positive effects on producers than consumer-mediated recycling in most cases but that consumer-mediated recycling does generally have a positive effect on producer stocks. We demonstrate that herbivore recycling will have larger effects on producer biomass than predator recycling when turnover rates and recycling efficiencies are high and predators prefer local prey. In addition, predictions suggest that consumer-mediated recycling has the largest effects on primary producers when predators prefer allochthonous prey and predator attack rates are high. Finally, our model predicts that consumer-mediated recycling effects may not be largest when external nutrient loading is low. Our model predictions highlight predator and prey feeding relationships, turnover rates, and external nutrient loading rates as key determinants of the strength of cascading trophic interactions. We show that existing hypotheses from specific empirical systems do not occur under all conditions, which further exacerbates the need to consider a broad suite of mechanisms when investigating trophic cascades.

Herbivore Impacts on Carbon Cycling in Boreal Forests.

Large herbivores can have substantial effects on carbon (C) cycling, yet these animals are often overlooked in C budgets. Zoogeochemical effects may be particularly important in boreal forests, where diverse human activities are facilitating the expansion of large herbivore populations. Here, we argue that considering trophic dynamics is necessary to understand spatiotemporal variability in boreal forest C budgets. We propose a research agenda to scale local studies to landscape extents to measure the zoogeochemical impacts of large herbivores on boreal forest C cycling. Distributed networks of exclosure experiments, empirical studies across gradients in large herbivore abundance, multiscale models using herbivore distribution data, and remote sensing paired with empirical data will provide comprehensive accounting of C source-sink dynamics in boreal forests.

Quantity-quality trade-offs revealed using a multiscale test of herbivore resource selection on elemental landscapes.

Herbivores consider the variation of forage qualities (nutritional content and digestibility) as well as quantities (biomass) when foraging. Such selection patterns may change based on the scale of foraging, particularly in the case of ungulates that forage at many scales.To test selection for quality and quantity in free-ranging herbivores across scales, however, we must first develop landscape-wide quantitative estimates of both forage quantity and quality. Stoichiometric distribution models (StDMs) bring opportunity to address this because they predict the elemental measures and stoichiometry of resources at landscape extents.Here, we use StDMs to predict elemental measures of understory white birch quality (% nitrogen) and quantity (g carbon/m2) across two boreal landscapes. We analyzed global positioning system (GPS) collared moose (n = 14) selection for forage quantity and quality at the landscape, home range, and patch extents using both individual and pooled resource selection analyses. We predicted that as the scale of resource selection decreased from the landscape to the patch, selection for white birch quantity would decrease and selection for quality would increase.Counter to our prediction, pooled-models showed selection for our estimates of quantity and quality to be neutral with low explanatory power and no scalar trends. At the individual-level, however, we found evidence for quality and quantity trade-offs, most notably at the home-range scale where resource selection models explain the largest amount of variation in selection. Furthermore, individuals did not follow the same trade-off tactic, with some preferring forage quantity over quality and vice versa.Such individual trade-offs show that moose may be flexible in attaining a limiting nutrient. Our findings suggest that herbivores may respond to forage elemental compositions and quantities, giving tools like StDMs merit toward animal ecology applications. The integration of StDMs and animal movement data represents a promising avenue for progress in the field of zoogeochemistry.