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Over the past several decades, various trends in vegetation productivity, from increases to decreases, have been observed throughout Arctic-Boreal ecosystems. While some of this variation can be explained by recent climate warming and increased disturbance, very little is known about the impacts of permafrost thaw on productivity across diverse vegetation communities. Active layer thickness data from 135 permafrost monitoring sites along a 10° latitudinal transect of the Northwest Territories, Canada, paired with a Landsat time series of normalized difference vegetation index from 1984 to 2019, were used to quantify the impacts of changing permafrost conditions on vegetation productivity. We found that active layer thickness contributed to the observed variation in vegetation productivity in recent decades in the northwestern Arctic-Boreal, with the highest rates of greening occurring at sites where the near-surface permafrost recently had thawed. However, the greening associated with permafrost thaw was not sustained after prolonged periods of thaw and appeared to diminish after the thaw front extended outside the plants' rooting zone. Highest rates of greening were found at the mid-transect sites, between 62.4° N and 65.2° N, suggesting that more southernly sites may have already surpassed the period of beneficial permafrost thaw, while more northern sites may have yet to reach a level of thaw that supports enhanced vegetation productivity. These results indicate that the response of vegetation productivity to permafrost thaw is highly dependent on the extent of active layer thickening and that increases in productivity may not continue in the coming decades., (© 2023 His Majesty the King in Right of Canada and The Authors. Global Change Biology published by John Wiley & Sons Ltd. Reproduced with the permission of the Minister of Natural Resources Canada.)

Most research on boreal populations of woodland caribou (Rangifer tarandus caribou) has been conducted in areas of high anthropogenic disturbance. However, a large portion of the species' range overlaps relatively pristine areas primarily affected by natural disturbances, such as wildfire. Climate-driven habitat change is a key concern for the conservation of boreal-dependent species, where management decisions have yet to consider knowledge from multiple ecological domains integrated into a cohesive and spatially explicit forecast of species-specific habitat and demography. We used a novel ecological forecasting framework to provide climate-sensitive projections of habitat and demography for five boreal caribou monitoring areas within the Northwest Territories (NWT), Canada, over 90 years. Importantly, we quantify uncertainty around forecasted mean values. Our results suggest habitat suitability may increase in central and southwest regions of the NWT's Taiga Plains ecozone but decrease in southern and northwestern regions driven by conversion of coniferous to deciduous forests. We do not project that boreal caribou population growth rates will change despite forecasted changes to habitat suitability. Our results emphasize the importance of efforts to protect and restore northern boreal caribou habitat despite climate uncertainty while highlighting expected spatial variations that are important considerations for local people who rely on them. An ability to reproduce previous work, and critical thought when incorporating sources of uncertainty, will be important to refine forecasts, derive management decisions, and improve conservation efficacy for northern species at risk., (© 2023 His Majesty the King in Right of Canada and The Authors. Ecological Applications published by Wiley Periodicals LLC on behalf of The Ecological Society of America. Reproduced with the permission of the Minister of Natural Resources.)

Resilience of plant communities to disturbance is supported by multiple mechanisms, including ecological legacies affecting propagule availability, species' environmental tolerances, and biotic interactions. Understanding the relative importance of these mechanisms for plant community resilience supports predictions of where and how resilience will be altered with disturbance. We tested mechanisms underlying resilience of forests dominated by black spruce ( Picea mariana ) to fire disturbance across a heterogeneous forest landscape in the Northwest Territories, Canada. We combined surveys of naturally regenerating seedlings at 219 burned plots with experimental manipulations of ecological legacies via seed addition of four tree species and vertebrate exclosures to limit granivory and herbivory at 30 plots varying in moisture and fire severity. Black spruce recovery was greatest where it dominated pre-fire, at wet sites with deep residual soil organic layers, and fire conditions of low soil or canopy combustion and longer return intervals. Experimental addition of seed indicated all species were seed-limited, emphasizing the importance of propagule legacies. Black spruce and birch ( Betula papyrifera ) recruitment were enhanced with vertebrate exclusion. Our combination of observational and experimental studies demonstrates black spruce is vulnerable to effects of increased fire activity that erode ecological legacies. Moreover, black spruce relies on wet areas with deep soil organic layers where other species are less competitive. However, other species can colonize these areas if enough seed is available or soil moisture is altered by climate change. Testing mechanisms underlying species' resilience to disturbance aids predictions of where vegetation will transform with effects of climate change., Supplementary Information: The online version contains supplementary material available at 10.1007/s10021-022-00772-7., (© The Author(s) 2022.)

Making predictions from ecological models-and comparing them to data-offers a coherent approach to evaluate model quality, regardless of model complexity or modelling paradigm. To date, our ability to use predictions for developing, validating, updating, integrating and applying models across scientific disciplines while influencing management decisions, policies, and the public has been hampered by disparate perspectives on prediction and inadequately integrated approaches. We present an updated foundation for Predictive Ecology based on seven principles applied to ecological modelling: make frequent Predictions, Evaluate models, make models Reusable, Freely accessible and Interoperable, built within Continuous workflows that are routinely Tested (PERFICT). We outline some benefits of working with these principles: accelerating science; linking with data science; and improving science-policy integration., (© 2022 Her Majesty the Queen in Right of Canada. Ecology Letters published by John Wiley & Sons Ltd. Reproduced with the permission of the Minister of Natural Resources Canada.)

Land-use change and climate change are recognized as two main drivers of the current biodiversity decline. Protected areas help safeguard the landscape from additional anthropogenic disturbances and, when properly designed, can help species cope with climate change impacts. When designed to protect the regional biodiversity rather than to conserve focal species or landscape elements, protected areas need to cover a representative sample of the regional biodiversity and be functionally connected, facilitating individual movements among protected areas in a network to maximize their effectiveness. We developed a methodology to define effective protected areas to implement in a regional network using ecological representativeness and functional connectivity as criteria. We illustrated this methodology in the Gaspésie region of Québec, Canada. We simulated movements for the endangered Atlantic-Gaspésie caribou population (Rangifer tarandus caribou), using an individual-based model, to determine functional connectivity based on this large mammal. We created multiple protected areas network scenarios and evaluated their ecological representativeness and functional connectivity for the current and future conditions. We selected a subset of the most effective network scenarios and extracted the protected areas included in them. There was a tradeoff between ecological representativeness and functional connectivity for the created networks. Only a few protected areas among those available were repeatedly chosen in the most effective networks. Protected areas maximizing both ecological representativeness and functional connectivity represented suitable areas to implement in an effective protected areas network. These areas ensured that a representative sample of the regional biodiversity was covered by the network, as well as maximizing the movement over time between and inside the protected areas for the focal population., Competing Interests: The authors have declared that no competing interests exist.

Boreal forest fires emit large amounts of carbon into the atmosphere primarily through the combustion of soil organic matter 1-3 . During each fire, a portion of this soil beneath the burned layer can escape combustion, leading to a net accumulation of carbon in forests over multiple fire events 4 . Climate warming and drying has led to more severe and frequent forest fires 5-7 , which threaten to shift the carbon balance of the boreal ecosystem from net accumulation to net loss 1 , resulting in a positive climate feedback 8 . This feedback will occur if organic-soil carbon that escaped burning in previous fires, termed 'legacy carbon', combusts. Here we use soil radiocarbon dating to quantitatively assess legacy carbon loss in the 2014 wildfires in the Northwest Territories of Canada 2 . We found no evidence for the combustion of legacy carbon in forests that were older than the historic fire-return interval of northwestern boreal forests 9 . In forests that were in dry landscapes and less than 60 years old at the time of the fire, legacy carbon that had escaped burning in the previous fire cycle was combusted. We estimate that 0.34 million hectares of young forests (<60 years) that burned in the 2014 fires could have experienced legacy carbon combustion. This implies a shift to a domain of carbon cycling in which these forests become a net source-instead of a sink-of carbon to the atmosphere over consecutive fires. As boreal wildfires continue to increase in size, frequency and intensity 7 , the area of young forests that experience legacy carbon combustion will probably increase and have a key role in shifting the boreal carbon balance.

Future human land use and climate change may disrupt movement behaviors of terrestrial animals, thereby altering the ability of individuals to move across a landscape. Some of the expected changes result from processes whose effects will be difficult to alter, such as global climate change. We present a novel framework in which we use models to (1) identify the ecological changes from these difficult-to-alter processes, as well as (2) the potential conservation measures that are best able to compensate for these changes. We illustrated this framework with the case of an endangered caribou population in Québec, Canada. We coupled a spatially explicit individual-based movement model with a range of landscape scenarios to assess the impacts of varying degrees of climate change, and the ability of conservation actions to compensate for such impacts on caribou movement behaviors. We found that (1) climate change impacts reduced movement potential, and that (2) the complete restoration of secondary roads inside protected areas was able to fully offset this reduction, suggesting that road restoration would be an effective compensatory conservation action. By evaluating conservation actions via landscape use simulated by an individual-based model, we were able to identify compensatory conservation options for an endangered species facing climate change.

Climate warming and drying is associated with increased wildfire disturbance and the emergence of megafires in North American boreal forests. Changes to the fire regime are expected to strongly increase combustion emissions of carbon (C) which could alter regional C balance and positively feedback to climate warming. In order to accurately estimate C emissions and thereby better predict future climate feedbacks, there is a need to understand the major sources of heterogeneity that impact C emissions at different scales. Here, we examined 211 field plots in boreal forests dominated by black spruce (Picea mariana) or jack pine (Pinus banksiana) of the Northwest Territories (NWT), Canada after an unprecedentedly large area burned in 2014. We assessed both aboveground and soil organic layer (SOL) combustion, with the goal of determining the major drivers in total C emissions, as well as to develop a high spatial resolution model to scale emissions in a relatively understudied region of the boreal forest. On average, 3.35 kg C m -2 was combusted and almost 90% of this was from SOL combustion. Our results indicate that black spruce stands located at landscape positions with intermediate drainage contribute the most to C emissions. Indices associated with fire weather and date of burn did not impact emissions, which we attribute to the extreme fire weather over a short period of time. Using these results, we estimated a total of 94.3 Tg C emitted from 2.85 Mha of burned area across the entire 2014 NWT fire complex, which offsets almost 50% of mean annual net ecosystem production in terrestrial ecosystems of Canada. Our study also highlights the need for fine-scale estimates of burned area that represent small water bodies and regionally specific calibrations of combustion that account for spatial heterogeneity in order to accurately model emissions at the continental scale., (© 2018 John Wiley & Sons Ltd.)

Factors affecting wildland-fire size distribution include weather, fuels, and fire suppression activities. We present a novel application of survival analysis to quantify the effects of these factors on a sample of sizes of lightning-caused fires from Alberta, Canada. Two events were observed for each fire: the size at initial assessment (by the first fire fighters to arrive at the scene) and the size at "being held" (a state when no further increase in size is expected). We developed a statistical classifier to try to predict cases where there will be a growth in fire size (i.e., the size at "being held" exceeds the size at initial assessment). Logistic regression was preferred over two alternative classifiers, with covariates consistent with similar past analyses. We conducted survival analysis on the group of fires exhibiting a size increase. A screening process selected three covariates: an index of fire weather at the day the fire started, the fuel type burning at initial assessment, and a factor for the type and capabilities of the method of initial attack. The Cox proportional hazards model performed better than three accelerated failure time alternatives. Both fire weather and fuel type were highly significant, with effects consistent with known fire behaviour. The effects of initial attack method were not statistically significant, but did suggest a reverse causality that could arise if fire management agencies were to dispatch resources based on a-priori assessment of fire growth potentials. We discuss how a more sophisticated analysis of larger data sets could produce unbiased estimates of fire suppression effect under such circumstances.

Fire activity in North American forests is expected to increase substantially with climate change. This would represent a growing risk to human settlements and industrial infrastructure proximal to forests, and to the forest products industry. We modelled fire size distributions in southern Québec as functions of fire weather and land cover, thus explicitly integrating some of the biotic interactions and feedbacks in a forest-wildfire system. We found that, contrary to expectations, land-cover and not fire weather was the primary driver of fire size in our study region. Fires were highly selective on fuel-type under a wide range of fire weather conditions: specifically, deciduous forest, lakes and to a lesser extent recently burned areas decreased the expected fire size in their vicinity compared to conifer forest. This has large implications for fire risk management in that fuels management could reduce fire risk over the long term. Our results imply, for example, that if 30% of a conifer-dominated landscape were converted to hardwoods, the probability of a given fire, occurring in that landscape under mean fire weather conditions, exceeding 100,000 ha would be reduced by a factor of 21. A similarly marked but slightly smaller effect size would be expected under extreme fire weather conditions. We attribute the decrease in expected fire size that occurs in recently burned areas to fuel availability limitations on fires spread. Because regenerating burned conifer stands often pass through a deciduous stage, this would also act as a negative biotic feedback whereby the occurrence of fires limits the size of nearby future for some period of time. Our parameter estimates imply that changes in vegetation flammability or fuel availability after fires would tend to counteract shifts in the fire size distribution favoring larger fires that are expected under climate warming. Ecological forecasts from models neglecting these feedbacks may markedly overestimate the consequences of climate warming on fire activity, and could be misleading. Assessments of vulnerability to climate change, and subsequent adaptation strategies, are directly dependent on integrated ecological forecasts. Thus, we stress the need to explicitly incorporate land-cover's direct effects and feedbacks in simulation models of coupled climate-fire-fuels systems.

It is becoming clear that fires in boreal forests are not uniformly stand-replacing. On the contrary, marked variation in fire severity, measured as tree mortality, has been found both within and among individual fires. It is important to understand the conditions under which this variation can arise. We integrated forest sample plot data, tree allometries and historical forest fire records within a diameter class-structured model of 1.0 ha patches of mono-specific black spruce and jack pine stands in northern Québec, Canada. The model accounts for crown fire initiation and vertical spread into the canopy. It uses empirical relations between fire intensity, scorch height, the percent of crown scorched and tree mortality to simulate fire severity, specifically the percent reduction in patch basal area due to fire-caused mortality. A random forest and a regression tree analysis of a large random sample of simulated fires were used to test for an effect of fireline intensity, stand structure, species composition and pyrogeographic regions on resultant severity. Severity increased with intensity and was lower for jack pine stands. The proportion of simulated fires that burned at high severity (e.g. >75% reduction in patch basal area) was 0.80 for black spruce and 0.11 for jack pine. We identified thresholds in intensity below which there was a marked sensitivity of simulated fire severity to stand structure, and to interactions between intensity and structure. We found no evidence for a residual effect of pyrogeographic region on simulated severity, after the effects of stand structure and species composition were accounted for. The model presented here was able to produce variation in fire severity under a range of fire intensity conditions. This suggests that variation in stand structure is one of the factors causing the observed variation in boreal fire severity.

Assessing spatial variation in waterfowl harvest probabilities from banding data is challenging because reporting and recovery probabilities have distinct spatial patterns that covary temporally with harvesting regulations, hunter effort, and reporting methods. We analyzed direct band recovery data from American black ducks banded on the Canadian breeding grounds from 1970 through 2010. Data were registered to a 1-degree grid and analyzed using hierarchical logistic regression models with spatially correlated errors to estimate the annual probabilities of band recovery and the proportion of individuals recovered in Canada. Probability of harvest was estimated from these values, in combination with independent estimates of reporting probabilities in Canada and the USA. Model covariates included estimates of hunting effort and factors for harvest regulation and band reporting methods. Both the band recovery processes and the proportion of individuals recovered in Canada had significant spatial structure. Recovery probabilities were highest in southern Ontario, along the Saint Lawrence River in Quebec, and in Nova Scotia. Black ducks breeding in Nova Scotia and southern Quebec were harvested predominantly in Canada. Recovery probabilities for juveniles were correlated with hunter effort, while the adult recoveries were weakly correlated with the implementation of stricter harvest regulations in the early 1980s. Mean harvest probability decreased in the northern portion of the survey area but remained stable or even increased in the south. Harvest probabilities for juveniles in 2010 exceeded 20% in southern Quebec and the Atlantic provinces. Our results demonstrate fine-scale variation in harvest probabilities for black duck on the Canadian breeding ground. In particular, harvest probabilities should be closely monitored along the Saint Lawrence River system and in the Atlantic provinces to avoid overexploitation.

For climate change projections to be useful, the magnitude of change must be understood relative to the magnitude of uncertainty in model predictions. We quantified the signal-to-noise ratio in projected distributional responses of boreal birds to climate change, and compared sources of uncertainty. Boosted regression tree models of abundance were generated for 80 boreal-breeding bird species using a comprehensive data set of standardized avian point counts (349,629 surveys at 122,202 unique locations) and 4-km climate, land use, and topographic data. For projected changes in abundance, we calculated signal-to-noise ratios and examined variance components related to choice of global climate model (GCM) and two sources of species distribution model (SDM) uncertainty: sampling error and variable selection. We also evaluated spatial, temporal, and interspecific variation in these sources of uncertainty. The mean signal-to-noise ratio across species increased over time to 2.87 by the end of the 21st century, with the signal greater than the noise for 88% of species. Across species, climate change represented the largest component (0.44) of variance in projected abundance change. Among sources of uncertainty evaluated, choice of GCM (mean variance component = 0.17) was most important for 66% of species, sampling error (mean= 0.12) for 29% of species, and variable selection (mean =0.05) for 5% of species. Increasing the number of GCMs from four to 19 had minor effects on these results. The range of projected changes and uncertainty characteristics across species differed markedly, reinforcing the individuality of species' responses to climate change and the challenges of one-size-fits-all approaches to climate change adaptation. We discuss the usefulness of different conservation approaches depending on the strength of the climate change signal relative to the noise, as well as the dominant source of prediction uncertainty.

Many animal species exhibit broad-scale latitudinal or longitudinal gradients in their response to biotic and abiotic components of their habitat. Although knowing the underlying mechanism of these patterns can be critical to the development of sound measures for the preservation or recovery of endangered species, few studies have yet identified which processes drive the existence of geographical gradients in habitat selection. Using extensive spatial data of broad latitudinal and longitudinal extent, we tested three hypotheses that could explain the presence of geographical gradients in landscape selection of the endangered boreal woodland caribou (Rangifer tarandus caribou) during winter in Eastern Canadian boreal forests: 1) climate-driven selection, which postulates that geographic gradients are surrogates for climatic gradients; 2) road-driven selection, which proposes that boreal caribou adjust their selection for certain habitat classes as a function of proximity to roads; and 3) an additive effect of both roads and climate. Our data strongly supported road-driven selection over climate influences. Thus, direct human alteration of landscapes drives boreal caribou distribution and should likely remain so until the climate changes sufficiently from present conditions. Boreal caribou avoided logged areas two-fold more strongly than burnt areas. Limiting the spread of road networks and accounting for the uneven impact of logging compared to wildfire should therefore be integral parts of any habitat management plan and conservation measures within the range of the endangered boreal caribou. The use of hierarchical spatial models allowed us to explore the distribution of spatially-structured errors in our models, which in turn provided valuable insights for generating alternative hypotheses about processes responsible for boreal caribou distribution.

Predictions of future fire activity over Canada's boreal forests have primarily been generated from climate data following assumptions that direct effects of weather will stand alone in contributing to changes in burning. However, this assumption needs explicit testing. First, areas recently burned can be less likely to burn again in the near term, and this endogenous regulation suggests the potential for self-limiting, negative biotic feedback to regional climate-driven increases in fire. Second, forest harvest is ongoing, and resulting changes in vegetation structure have been shown to affect fire activity. Consequently, we tested the assumption that fire activity will be driven by changes in fire weather without regulation by biotic feedback or regional harvest-driven changes in vegetation structure in the mixedwood boreal forest of Alberta, Canada, using a simulation experiment that includes the interaction of fire, stand dynamics, climate change, and clear cut harvest management. We found that climate change projected with fire weather indices calculated from the Canadian Regional Climate Model increased fire activity, as expected, and our simulations established evidence that the magnitude of regional increase in fire was sufficient to generate negative feedback to subsequent fire activity. We illustrate a 39% (1.39-fold) increase in fire initiation and 47% (1.47-fold) increase in area burned when climate and stand dynamics were included in simulations, yet 48% (1.48-fold) and 61% (1.61-fold) increases, respectively, when climate was considered alone. Thus, although biotic feedbacks reduced burned area estimates in important ways, they were secondary to the direct effect of climate on fire. We then show that ongoing harvest management in this region changed landscape composition in a way that led to reduced fire activity, even in the context of climate change. Although forest harvesting resulted in decreased regional fire activity when compared to unharvested conditions, forest composition and age structure was shifted substantially, illustrating a trade-off between management goals to minimize fire and conservation goals to emulate natural disturbance.

Systematic conservation plans have only recently considered the dynamic nature of ecosystems. Methods have been developed to incorporate climate change, population dynamics, and uncertainty in reserve design, but few studies have examined how to account for natural disturbance. Considering natural disturbance in reserve design may be especially important for the world's remaining intact areas, which still experience active natural disturbance regimes. We developed a spatially explicit, dynamic simulation model, CONSERV, which simulates patch dynamics and fire, and used it to evaluate the efficacy of hypothetical reserve networks in northern Canada. We designed six networks based on conventional reserve design methods, with different conservation targets for woodland caribou habitat, high-quality wetlands, vegetation, water bodies, and relative connectedness. We input the six reserve networks into CONSERV and tracked the ability of each to maintain initial conservation targets through time under an active natural disturbance regime. None of the reserve networks maintained all initial targets, and some over-represented certain features, suggesting that both effectiveness and efficiency of reserve design could be improved through use of spatially explicit dynamic simulation during the planning process. Spatial simulation models of landscape dynamics are commonly used in natural resource management, but we provide the first illustration of their potential use for reserve design. Spatial simulation models could be used iteratively to evaluate competing reserve designs and select targets that have a higher likelihood of being maintained through time. Such models could be combined with dynamic planning techniques to develop a general theory for reserve design in an uncertain world.

Lightning fire is the dominant natural disturbance of the western mixedwood boreal forest of North America. We quantified the independent effects of weather and forest composition on lightning fire initiation (a detected and recorded fire start) patterns in Alberta, Canada, to demonstrate how these biotic and abiotic components contribute to ecosystem dynamics in the mixedwood boreal forest. We used logistic regression to describe variation in annual initiation occurrence among 10,000-ha landscape units (voxels) covering a 9 million-ha study region over 11 years. At a voxel scale, forest composition explained more variation in annual initiation than did weather indices. Initiations occurred more frequently in landscapes with more conifer fuels (Picea spp.), and less in aspen-dominated (Populus spp.) ones. Initiations were less frequent in landscapes that had recently burned. Variation in initiation was also influenced by joint weather-lightning indices, but to a lesser degree. For each voxel, these indices quantified the number of days in the fire season when moisture levels were low and lightning was detected. Regional indices of fire weather severity explained substantial interannual variation of initiation, and the effect of forest composition was stronger in years with more severe fire weather. Our study is a conclusive demonstration of biotic and abiotic regulation of lightning fire initiation in the mixedwood boreal forest. The independent effects of forest composition emphasize that vegetation feedbacks strongly regulate disturbance dynamics in the region.