Extreme climate events limit northern range expansion of wild turkeys.

For species inhabiting areas at the limit of their environmental tolerance, extreme events often drive population persistence. However, because extreme events are uncommon, their effects on population dynamics of expanding species are poorly known. We examined how extreme climate events in winter and summer affected three populations of wild turkeys occupying a natural climate gradient at the northern edge of their range. First, we examined the mechanism by which vital rates affect the population growth rate. Second, we developed a climate-dependent structured population model. Finally, by linking this population model to IPCC-class climate projections, we projected wild turkey population abundance in response to the frequency of extreme snow events by 2100 for the northernmost population. We showed that the population dynamics of the three populations is driven through different pathways expected from the theory of invading population dynamics; that those populations were mainly limited by heavy snow that decreases winter survival by restraining food access; and that a population of immigrant is projected to decline at the northern species range. This study exemplifies how extreme events affect population dynamics and range expansion of temperate species at the northern edge of the distribution.

Cool as a moose: How can browsing counteract climate warming effects across boreal forest ecosystems?

Herbivory has potential to modify vegetation responses to climatic changes. However, climate and herbivory also affect each other, and rarely work in isolation from other ecological factors, such as plant-plant competition. Thus, it is challenging to predict the extent to which herbivory can counteract, amplify, or interact with climate impacts on ecosystems. Here, we investigate how moose modify climatic responses of boreal trees by using experimental exclosures on two continents and modeling complex causal pathways including several climatic factors, multiple tree species, competition, tree height, time, food availability, and herbivore presence, density, and browsing intensity. We show that moose can counteract, that is, "cool down" positive temperature responses of trees, but that this effect varies between species depending on moose foraging preferences. Growth of preferred deciduous trees was strongly affected by moose, whereas growth of less preferred conifers was mostly driven by climate and tree height. In addition, moose changed temperature responses of rowan in Norway and balsam fir in Canada, by making fir more responsive to temperature but decreasing the strength of the temperature response of rowan. Snow protected trees from browsing, and therefore moose "cooling power" might increase should a warming climate result in decreased snow cover. Furthermore, we found evidence of indirect effects of moose via plant-plant competition: By constraining growth of competing trees, moose can contribute positively to the growth of other trees. Our study shows that in boreal forests, herbivory cooling power is highly context dependent, and in order to understand its potential to prevent changes induced by warming climate, species differences, snow, competition, and climate effects on browsing need to be considered.

Phytochemicals Involved in Plant Resistance to Leporids and Cervids: a Systematic Review.

Non-nutritive phytochemicals (secondary metabolites and fibre) can influence plant resistance to herbivores and have ecological impacts on animal and plant population dynamics. A major hindrance to the ecological study of these phytochemicals is the uncertainty in the compounds one should measure, especially when limited by cost and expertise. With the underlying goal of identifying proxies of plant resistance to herbivores, we performed a systematic review of the effects of non-nutritive phytochemicals on consumption by leporids (rabbits and hares) and cervids (deer family). We identified 133 out of 1790 articles that fit our selection criteria (leporids = 33, cervids = 97, both herbivore types = 3). These articles cover 18 species of herbivores, on four continents. The most prevalent group of phytochemicals in the selected articles was phenolics, followed by terpenes for leporids and by fibre for cervids. In general, the results were variable but phenolic concentration seems linked with high resistance to both types of herbivores. Terpene concentration is also linked to high plant resistance; this relationship seems driven by total terpene content for cervids and specific terpenes for leporids. Tannins and fibre did not have a consistent positive effect on plant resistance. Because of the high variability in results reported and the synergistic effects of phytochemicals, we propose that the choice of chemical analyses must be tightly tailored to research objectives. While researchers pursuing ecological or evolutionary objectives should consider multiple specific analyses, researchers in applied studies could focus on a fewer number of specific analyses. An improved consideration of plant defence, based on meaningful chemical analyses, could improve studies of plant resistance and allow us to predict novel or changing plant-herbivore interactions.

Spatial correlations between browsing on balsam fir by white-tailed deer and the nutritional value of neighboring winter forage.

Associational effects, that is, the influence of neighboring plants on herbivory suffered by a plant, are an outcome of forage selection. Although forage selection is a hierarchical process, few studies have investigated associational effects at multiple spatial scales. Because the nutritional quality of plants can be spatially structured, it might differently influence associational effects across multiple scales. Our objective was to determine the radius of influence of neighbor density and nutritional quality on balsam fir (Abies balsamea) herbivory by white-tailed deer (Odocoileus virginianus) in winter. We quantified browsing rates on fir and the density and quality of neighboring trees in a series of 10-year-old cutovers on Anticosti Island (Canada). We used cross-correlations to investigate relationships between browsing rates and the density and nutritional quality of neighboring trees at distances up to 1,000 m. Balsam fir and white spruce (Picea glauca) fiber content and dry matter in vitro true digestibility were correlated with fir browsing rate at the finest extra-patch scale (across distance of up to 50 m) and between cutover areas (300-400 m). These correlations suggest associational effects, that is, low nutritional quality of neighbors reduces the likelihood of fir herbivory (associational defense). Our results may indicate associational effects mediated by intraspecific variation in plant quality and suggest that these effects could occur at scales from tens to hundreds of meters. Understanding associational effects could inform strategies for restoration or conservation; for example, planting of fir among existing natural regeneration could be concentrated in areas of low nutritional quality.

Experimental warming alters migratory caribou forage quality.

Global declines in caribou and reindeer (Rangifer) populations have drawn attention to the myriad of stressors that these Arctic and boreal forest herbivores currently face. Arctic warming has resulted in increased tundra shrub growth and therefore Rangifer forage quantity. However, its effects on forage quality have not yet been addressed although they may be critical to Rangifer body condition and fecundity. We investigated the impact of 8 yrs of summer warming on the quality of forage available to the Bathurst caribou herd using experimental greenhouses (n = 5) located in mesic birch hummock tundra in the central Canadian Low Arctic. Leaf forage quality and digestibility characteristics associated with nutrients (nitrogen and phosphorus), phenolics, and fiber were measured on the deciduous shrub Betula glandulosa (an important Rangifer diet component) at six time points through the growing season, and on five other very common vascular plant and lichen species in late summer. Experimental warming reduced B. glandulosa leaf nitrogen concentrations by ~10% in both late June and mid-July, but not afterwards. It also reduced late summer forage quality of the graminoid Eriophorum vaginatum by increasing phenolic concentrations 38%. Warming had mixed effects on forage quality of the lichen Cetraria cucullata in that it increased nutrient concentrations and tended to decrease fiber contents, but it also increased phenolics. Altogether, these warming-induced changes in forage quality over the growing season, and response differences among species, highlight the importance of Rangifer adaptability in diet selection. Furthermore, the early season reduction in B. glandulosa nitrogen content is a particular concern given the importance of this time for calf growth. Overall, our demonstration of the potential for significant warming impacts on forage quality at critical times for these animals underscores the importance of effective Rangifer range conservation to ensure sufficient appropriate habitat to support adaptability in forage selection in a rapidly changing environment.

Combining near infrared spectra of feces and geostatistics to generate forage nutritional quality maps across landscapes.

An important asset for the management of wild ungulates is recognizing the spatial distribution of forage quality across heterogeneous landscapes. To do so typically requires knowledge of which plant species are eaten, in what abundance they are eaten, and what their nutritional quality might be. Acquiring such data, however, may be difficult and time consuming. Here, we are proposing a rapid and cost-effective forage quality monitoring tool that combines near infrared (NIR) spectra of fecal samples and easily obtained data on plant community composition. Our approach rests on the premise that NIR spectra of fecal samples collected within low population density exclosures reflect the optimal forage quality of a given landscape. Forage quality can thus be based on the Mahalanobis distance of fecal spectral scans across the landscape relative to fecal spectral scans inside exclosures (referred to as DISTEX). The Gi* spatial autocorrelation statistic can then be applied among neighboring DISTEX values to detect and map "hot spots" and "cold spots" of nutritional quality over the landscape. We tested our approach in a heterogeneous boreal landscape on Anticosti Island (Québec, Canada), where white-tailed deer (Odocoileus virginianus) populations over the landscape have ranged from 20 to 50 individuals/km2 for at least 80 years, resulting in a loss of most palatable and nutritious plant species. Our results suggest that hot spots of forage quality occur when old-growth balsam fir stands comprise >39.8% of 300 ha neighborhoods, whereas cold spots occur in laggs (i.e., transition zones from forest to peatland). In terms of ground-level indicator plant species, the presence of Canada bunchberry (Cornus canadensis) was highly correlated with hot spots, whereas tamarack (Larix laricina) was highly correlated with cold spots. Mean DISTEX values were positively and significantly correlated with the neutral detergent fiber and acid detergent lignin contents of feces. While our approach would need more independent field trials before it is fully validated, its low cost and ease of execution should make it a valuable tool for advancing both the basic and applied ecology of large herbivores.

Isotopic Differences between Forage Consumed by a Large Herbivore in Open, Closed, and Coastal Habitats: New Evidence from a Boreal Study System.

Documenting habitat-related patterns in foraging behaviour at the individual level and over large temporal scales remains challenging for large herbivores. Stable isotope analysis could represent a valuable tool to quantify habitat-related foraging behaviour at the scale of individuals and over large temporal scales in forest dwelling large herbivores living in coastal environments, because the carbon (δ13C) or nitrogen (δ15N) isotopic signatures of forage can differ between open and closed habitats or between terrestrial and littoral forage, respectively. Here, we examined if we could detect isotopic differences between the different assemblages of forage taxa consumed by white-tailed deer that can be found in open, closed, supralittoral, and littoral habitats. We showed that δ13C of assemblages of forage taxa were 3.0 ‰ lower in closed than in open habitats, while δ15N were 2.0 ‰ and 7.4 ‰ higher in supralittoral and littoral habitats, respectively, than in terrestrial habitats. Stable isotope analysis may represent an additional technique for ecologists interested in quantifiying the consumption of terrestrial vs. marine autotrophs. Yet, given the relative isotopic proximity and the overlap between forage from open, closed, and supralittoral habitats, the next step would be to determine the potential to estimate their contribution to herbivore diet.

Forage-mediated density and climate effects on body mass in a temperate herbivore: a mechanistic approach.

The interplay between density and climate in shaping the dynamics of herbivore populations is widely acknowledged, and current research is fueled by the identification of mechanisms underlying their effects on individuals and populations. We assessed whether forage availability mediated the effects of density and winter climate on body mass of white-tailed deer (Odocoileus virginianus) yearlings by experimentally reducing deer density to 7.5 and 15 deer/km2 during eight growing seasons, and by using causal (graphical) hierarchical models and Bayesian hierarchical modeling to assess relationships. The abundance of preferred forage decreased with deer density and varied quadratically (positive parabola) with winter North Atlantic Oscillation (NAO), whereas the fall mass of yearlings increased with forage abundance and spring mass. Fall mass did not differ between experimentally reduced deer densities, yet experimental yearlings were 30% heavier than yearlings harvested at ambient densities. Hence, forage abundance simultaneously mediated the effects of density and climate on fall body mass, which was also influenced by carry-over effects of spring body mass. Our findings increase our ability to anticipate how temperate large herbivores will respond to ongoing changes in intrinsic (e.g., large-herbivore density) and extrinsic (e.g., climate) factors.

Functional responses and resilience of boreal forest ecosystem after reduction of deer density.

The functional trait-based approach is increasingly used to predict responses of ecological communities to disturbances, but most studies target a single taxonomic group. Here, we assessed the resilience of a forest ecosystem to an overabundant herbivore population by assessing changes in 19 functional traits for plant, 13 traits for ground beetle and 16 traits for songbird communities after six years of controlled browsing on Anticosti Island (Quebec, Canada). Our results indicated that plants were more responsive to 6 years of reduced browsing pressure than ground beetles and songbirds. However, co-inertia analysis revealed that ground beetle communities responded in a similar way than plant communities with stronger relationships between plant and ground beetle traits at reduced deer density, a pattern not detected between plant and songbird. High deer density favored plants species that reproduce vegetatively and with abiotic pollination and seed dispersal, traits implying little interaction with animal. On the other hand, traits found at reduced deer density mostly involved trophic interaction. For example, plants in this treatment had fleshy fruits and large seeds dispersed by birds or other animals whereas ground beetle species were carnivorous. Overall, our results suggest that plant communities recovered some functional components to overabundant herbivore populations, since most traits associated with undisturbed forests were reestablished after six years of deer reduction. The re-establishment of functional plant communities with traits involving trophic interaction induces changes in the ground-beetle trait community, but forest structure remains likely insufficiently heterogeneous to shift the songbird trait community within six years.

A large herbivore triggers alternative successional trajectories in the boreal forest.

Alternative successional trajectories (AST) may result in multiple climax states within an ecosystem when disturbances affect colonization history. In the boreal forest, ungulates have been proposed to drive AST because, under herbivore pressure, preferred species may go extinct and apparent competition may benefit browsing-resistant species. Over a 15-year period following logging, we tested whether deer herbivory altered plant species composition and whether the competitive advantage of resistant species was maintained following herbivore removal. We compared exclosures built immediately after logging with delayed exclosures built eight years later on Anticosti Island, Quebec, Canada. Although the palatable tree Betula papyrifera (paper birch) and some palatable herbs recovered in delayed exclosures, we observed legacies in both tree and herb cover. Woody regeneration in delayed exclosures was dominated by Picea glauca (white spruce), and Poaceae (grasses) were abundant in the field layer. Given that only early-successional species recovered, whereas late-successional broadleaf species and Abies balsamea (balsam fir) remained rare, succession may follow an AST after a limited browsing period during early succession.

Tolerance of an expanding subarctic shrub, Betula glandulosa, to simulated caribou browsing.

Densification of the shrub layer has been reported in many subarctic regions, raising questions about the implication for large herbivores and their resources. Shrubs can tolerate browsing and their level of tolerance could be affected by browsing and soils productivity, eventually modifying resource availability for the caribou. Our objective was to assess the compensatory growth potential of a subarctic shrub, Betula glandulosa Michx., in relation with caribou browsing and nutriment availability for the plants. We used a simulated browsing (0, 25 and 75% of available shoots) and nitrogen-fertilisation (0 and 10 g m(-2)) experiment to test two main hypotheses linking tolerance to resource availability, the Compensatory Continuum Hypothesis and the Growth Rate Hypothesis as well as the predictions from the Limiting Resource Model. We seek to explicitly integrate the relative browsing pressure in our predictions since the amount of tissues removed could affect the capacity of long-lived plants to compensate. Birches fully compensated for moderate browsing with an overall leaf biomass similar to unbrowsed birches but undercompensated under heavy browsing pressure. The main mechanism explaining compensation appears to be the conversion of short shoots into long shoots. The leaf area increased under heavy browsing pressure but only led to undercompensation. Fertilisation for two consecutive years did not influence the response of birch, thus we conclude that our results support the LRM hypothesis of equal tolerance under both high and low nitrogen availability. Our results highlight that the potential for compensatory growth in dwarf birch is surpassed under heavy browsing pressure independently of the fertilisation regime. In the context of the worldwide decline in caribou herds, the reduction in browsing pressure could act synergistically with global climate change to promote the current shrub expansion reported in subarctic regions.

A new way of assessing foraging behaviour at the individual level using faeces marking and satellite telemetry.

Heterogeneity in foraging behaviour can profoundly influence ecological processes shaping populations. To scale-up from individual foraging behaviour to processes occurring at the population scale, one needs to sample foraging behaviour at the individual level, and over large temporal scales or during critical seasons known to influence life-history traits. We developed an innovative technique to monitor foraging behaviour at the individual level in secretive species, a technique that can be ultimately used to investigate the links between foraging behaviour and life-history traits. First, the technique used a novel approach, namely the combination of telemetry tracking and biomarking of faeces with food dyes to locate fresh signs of presence left by individuals equipped with GPS collars. Second, the technique is based on the simultaneous or successive sampling of life-history traits and individual foraging behaviour, using tracks with high probabilities of recovery of dyed faeces. We first describe our methodological approach, using a case study of a large herbivore, and then provide recommendations and guidelines for its use. Sampling single snow tracks of individuals equipped with a GPS collar was a reliable way to assess individual winter foraging behaviour in a white-tailed deer (Odocoileus virginianus Zimmermann) population. During that period, the probability of recovery of dyed faeces within the range of the collar precision was very high for single snow tracks of equipped deer (97%). Our approach is well suited to study individual foraging behaviour, and could ultimately be used to investigate the interplay between intra-population heterogeneity in foraging behaviour, life-history traits, and demographic processes.

Divergent nonlinear responses of the boreal forest field layer along an experimental gradient of deer densities.

The early responses of the field layer to changes in biotic and abiotic conditions are key determinants of the future composition and structure of forests where sustained heavy browsing pressure has depauperated the shrub understory. We investigated the relationships between white-tailed deer density and field layer plant community dynamics in boreal forests managed for wildlife and timber production. We hypothesized that the growth and reproduction of field layer plants are either: (H(1)) directly proportional to deer density, or (H(2)) related to deer density through nonlinear relationships or (H(3)) through nonlinear relationships with thresholds. We tested these hypotheses using data from a controlled browsing experiment involving a gradient of deer densities (0, 7.5, 15, 27 and 56 deer km(-2)) in interaction with timber harvesting conducted on Anticosti Island, Canada. In recent clearcuts, the dominant responses of the field layer plants were exponential recovery in growth and reproduction with decreasing deer densities. The abundance of browse-tolerant species such as grasses was positively related to deer density, suggesting an apparent competitive gain. These results support the prediction from our second hypothesis, although the presence of ecological thresholds should not be ruled out. Rapid changes in the early successional stages have potentially long-term consequences on successional patterns through processes such as the modulation of germination and early establishment success of seedlings from later successional species. Quantitative data as those presented here are essential for the development of ecosystem management prescriptions. On Anticosti Island, reduction of local deer densities to levels <15-7.5 deer km(-2) in the first 3 years following timber harvesting appears to be compatible with the regeneration dynamics of this system although lower levels of deer densities may be required for the conservation of browse-sensitive plant species.