Boreal-temperate Ecotone Research Articles

First evidence of nitrogen fixation associated with bryophytes from coastal Wabanaki–Acadian forests

Associations between bryophytes and dinitrogen (N2)-fixing bacteria are a significant source of exogenous N in unmanaged boreal and possibly temperate ecosystems. However, the extent to which biological N2-fixation (BNF) applies to the boreal–temperate ecotone remains elusive. The current focus on common species limits our understanding of BNF at the community level. Our objective was to characterize the presence of cyanobacteria and BNF activity associated with bryophytes in the coastal forests of Fundy National Park (New Brunswick, Canada). In 2021, we harvested three liverwort and 11 moss species from two sites (71 samples) and measured environmental covariates (e.g., canopy composition and soil pH). We used stable isotope incubations with 15N2 gas in growth chambers to quantify potential BNF activity and used phycocyanin extractions as a cyanobacteria abundance proxy. Many species presented detectable BNF rates, which were similar to or higher than those of well-studied feather mosses. These included species that have rarely been found to contribute to BNF. While cyanobacteria were present on most samples, we found no positive association between abundance and BNF. Our findings are among the first records for bryophyte-associated BNF in the boreal–temperate ecotone of eastern Canada and offer insights into the potential role of this process in N cycling in coastal conifer-dominated forests.

Climate change–induced stress disrupts ectomycorrhizal interaction networks at the boreal–temperate ecotone

The interaction networks formed by ectomycorrhizal fungi (EMF) and their tree hosts, which are important to both forest recruitment and ecosystem carbon and nutrient retention, may be particularly susceptible to climate change at the boreal-temperate forest ecotone where environmental conditions are changing rapidly. Here, we quantified the compositional and functional trait responses of EMF communities and their interaction networks with two boreal (Pinus banksiana and Betula papyrifera) and two temperate (Pinus strobus and Quercus macrocarpa) hosts to a factorial combination of experimentally elevated temperatures and reduced rainfall in a long-term open-air field experiment. The study was conducted at the B4WarmED (Boreal Forest Warming at an Ecotone in Danger) experiment in Minnesota, USA, where infrared lamps and buried heating cables elevate temperatures (ambient, +3.1 °C) and rain-out shelters reduce growing season precipitation (ambient, ~30% reduction). EMF communities were characterized and interaction networks inferred from metabarcoding of fungal-colonized root tips. Warming and rainfall reduction significantly altered EMF community composition, leading to an increase in the relative abundance of EMF with contact-short distance exploration types. These compositional changes, which likely limited the capacity for mycelial connections between trees, corresponded with shifts from highly redundant EMF interaction networks under ambient conditions to less redundant (more specialized) networks. Further, the observed changes in EMF communities and interaction networks were correlated with changes in soil moisture and host photosynthesis. Collectively, these results indicate that the projected changes in climate will likely lead to significant shifts in the traits, structure, and integrity of EMF communities as well as their interaction networks in forest ecosystems at the boreal-temperate ecotone.

Large fires or small fires, will they differ in affecting shifts in species composition and distributions under climate change?

Climate change is expected to increase fire activity, which has the potential to accelerate climate-induced shifts in species composition and distribution in the boreal-temperate ecotone. Wildfire can kill resident trees, and thus provide establishment opportunities for migrating tree species. However, the role of fire size and its interactions with tree species with varied life-history attributes in driving climate-induced shifts is not understood. Future fire regimes could be characterized by many small fires or a few large fires. Large and small fires create and regulate distinct burn patterns, which may influence tree-species responses and post-fire successional trajectories. Here we investigated the effects of future fire-regime variability on the boreal-temperate ecotone of northeastern China under climate change using a coupled forest dynamic model (LANDIS PRO) and ecosystem process model (LINKAGES). We simulated fire regimes using the LANDIS PRO fire module. We designed two fire scenarios (frequent, small fires and infrequent, large fires) to represent different fire regimes in terms of fire size. Results showed fire-catalyzed, climate-induced transitions from boreal to pioneer and temperate forest communities. Frequent, small fires resulted in 13% and 23% higher increases in pioneer and temperate species respectively, relative to infrequent, large fires. Therefore, species composition shifts were faster following frequent, small fires than infrequent, large fires. The results can help policymakers and forest managers determine tradeoffs among strategies to mitigate or adapt to climate change under altered fire regimes.

Stand-Level Variation Drives Canopy Water Storage by Non-vascular Epiphytes Across a Temperate-Boreal Ecotone

Epiphytes, including bryophytes and lichens, can significantly change the water interception and storage capacities of forest canopies. However, despite some understanding of this role, empirical evaluations of canopy and bole community water storage capacity by epiphytes are still quite limited. Epiphyte communities are shaped by both microclimate and host plant identity, and so the canopy and bole community storage capacity might also be expected to vary across similar spatial scales. We estimated canopy and bole community cover and biomass of bryophytes and lichens from ground-based surveys across a temperate-boreal ecotone in continental North America (Minnesota). Multiple forest types were studied at each site, to separate stand level and latitudinal effects. Biomass was converted into potential canopy and bole community storage on the basis of water-holding capacity measurements of dominant taxa. Bole biomass and potential water storage was a much larger contributor than outer canopy. Biomass and water storage capacity varied greatly, ranging from 9 to >900kg ha–1 and 0.003 to 0.38 mm, respectively. These values are lower than most reported results for temperate forests, which have emphasized coastal and old-growth forests. Variation was greatest within sites and appeared to reflect the strong effects of host tree identity on epiphyte communities, with conifer-dominated plots hosting more lichen-dominated epiphyte communities with lower potential water storage capacity. These results point to the challenges of estimating and incorporating epiphyte contributions to canopy hydrology from stand metrics. Further work is also needed to improve estimates of canopy epiphytes, including crustose lichens.

Forest structure, not climate, is the primary driver of functional diversity in northeastern North America

Functional diversity (FD), represented by plant traits, is fundamentally linked to an ecosystem's capacity to respond to environmental change. Yet, little is known about the spatial distribution of FD and its drivers. These knowledge gaps prevent the development of FD-based forest management approaches to increase the trait diversity insurance (i.e., the response diversity) against future environmental fluctuations and disturbances. Our study helps fill these knowledge gaps by (i) mapping the current FD distribution, (ii) and analyzing FD drivers across northeastern North America. Following the stress-dominance hypothesis, we expected a strong environmental filtering effect on FD. Moreover, we expected abundant species to determine the bulk of FD distributions as suggested by the mass-ratio hypothesis. We combined a literature and database review of 44 traits for 43 tree species with terrestrial inventory data of 48,426 plots spanning an environmental gradient from northern boreal to temperate biomes. We evaluated the statistical influence of 25 covariates related to forest structure, climate, topography, soils, and stewardship on FD by employing an ensemble approach consisting of 90 non-parametric models. Temperate forests and the boreal-temperate ecotone east and northeast of the Great Lakes were identified as FD hotspots. Environmental filtering by climate was of secondary importance, with forest structure explaining most of the FD distribution of tree species in northeastern North America. Thus, our study provides only partial support for the stress-dominance hypothesis. Species abundance weightings altered trait diversity distributions and drivers only marginally, supporting the mass-ratio hypothesis. Our results suggest that forest management could increase FD without requiring knowledge of functional ecology by fostering stand structural complexity instead. Further, mixing species from different functional groups identified in this study can enhance the trait diversity insurance of forests to an uncertain future.

Short- and long-term responses of photosynthetic capacity to temperature in four boreal tree species in a free-air warming and rainfall manipulation experiment.

High latitude forests cope with considerable variation in moisture and temperature at multiple temporal scales. To assess how their photosynthetic physiology responds to short- and long-term temperature variation, we measured photosynthetic capacity for four tree species growing in an open-air experiment in the boreal-temperate ecotone `Boreal Forest Warming at an Ecotone in Danger' (B4WarmED). The experiment factorially manipulated temperature above- and below-ground (ambient, +3.2°C) and summer rainfall (ambient, 40% removal). We measured A/Ci curves at 18, 25 and 32°C for individuals of two boreal (Pinus banksiana Lamb., Betula papyrifera Marsh.) and two temperate species (Pinus strobus L., Acer rubrum L.) experiencing the long-term warming and/or reduced-rainfall conditions induced by our experimental treatments. We calculated the apparent photosynthetic capacity descriptors VCmax,Ci and Jmax,Ci and their ratio for each measurement temperate. We hypothesized that (i) VCmax,Ci and Jmax,Ci would be down-regulated in plants experiencing longer term (e.g., weeks to months) warming and reduced rainfall (i.e., have lower values at a given measurement temperature), as is sometimes found in the literature, and that (ii) plants growing at warmer temperatures or from warmer ranges would show greater sensitivity (steeper slope) to short-term (minutes to hours) temperature variation. Neither hypothesis was supported as a general trend across the four species, as there was not a significant main effect (across species) of either warming or rainfall reduction on VCmax,Ci and Jmax,Ci. All species markedly increased VCmax,Ci and Jmax,Ci (and decreased their ratio) with short-term increases in temperature (i.e., contrasting values at 18, 25 and 32°C), and those responses were independent of long-term treatments and did not differ among species. The Jmax,Ci:VCmax,Ci ratio was, however, significantly lower across species in warmed and reduced rainfall treatments. Collectively, these results suggest that boreal trees possess considerable short-term plasticity that may allow homeostasis of VCmax,Ci and Jmax,Ci to a longer term temperature treatment. Our results also caution against extrapolating results obtained under controlled and markedly contrasting temperature treatments to responses of photosynthetic parameters to more modest temperature changes expected in the near-term with climate warming in field conditions.

Tree Encroachment Varies by Plant Community in a Large Boreal Peatland Complex in the Boreal-Temperate Ecotone of Northeastern USA

Large peatland complexes at the boreal-temperate ecotone are essential habitats for boreal species at their southern range limits where they are threatened by tree encroachment accelerated by climate change and nitrogen deposition. To inform vascular plant and biodiversity conservation, we studied tree encroachment patterns in a large (> 400 ha) boreal peatland complex in the northeastern United States across vegetation types and environmental gradients. We characterized vascular plant composition, environmental drivers and tree demography on 50 plots (each 25 m2). We used non-metric multidimensional scaling (NMS) to identify two main drivers of vascular plant composition in the herbaceous layer—pH and tree canopy openness—that described three broad plant community types (open bog, forested bog, and fen). Tree demography suggested that woody encroachment (i.e., tree seedling recruitment) varied across these community types; open bog was colonized by Picea mariana seedlings, while forested bog and fen (dominated by evergreen conifers, Picea mariana and Thuja occidentalis, respectively) were colonized by deciduous tree species (Acer rubrum and Betula alleghaniensis). Our findings provide early warning signs of vegetation change in boreal peatlands near their southern range limits caused by the encroachment of temperate tree species into forested peatlands and expanding tree cover in open bogs.

Moderate disturbances accelerate forest transition dynamics under climate change in the temperate-boreal ecotone of eastern North America.

Several temperate tree species are expected to migrate northward and colonize boreal forests in response to climate change. Tree migrations could lead to transitions in forest types, but these could be influenced by several non-climatic factors, such as disturbances and soil conditions. We analysed over 10,000 forest inventory plots, sampled from 1970 to 2018 in meridional Québec, Canada, to identify what environmental conditions promote or prevent regional-scale forest transitions. We used a continuous-time multi-state Markov model to quantify the probabilities of transitions between forest states (temperate, boreal, mixed, pioneer) as a function of climate (mean temperature and climate moisture index during the growing season), soil conditions (pH and drainage) and disturbances (severity levels of natural disturbances and logging). We further investigate how different disturbance types and severities impact forests' short-term transient dynamics and long-term equilibrium using properties of Markov transition matrices. The most common transitions observed during the study period were from mixed to temperate states, as well as from pioneer to boreal forests. In our study, transitions were mainly driven by natural and anthropogenic disturbances and secondarily by climate, whereas soil characteristics exerted relatively minor constraints. While major disturbances only promoted transitions to the pioneer state, moderate disturbances increased the probability of transition from mixed to temperate states. Long-term projections of our model under the current environmental conditions indicate that moderate disturbances would promote a northward shift of the temperate forest. Moreover, disturbances reduced turnover and convergence time for all transitions, thereby accelerating forest dynamics. Contrary to our expectation, mixed to temperate transitions were not driven by temperate tree recruitment but by mortality and growth. Overall, our results suggest that moderate disturbances could catalyse rapid forest transitions and accelerate broad-scale biome shifts.

Warming and disturbance alter soil microbiome diversity and function in a northern forest ecotone

The response to global change by soil microbes is set to affect important ecosystem processes. These impacts could be most immediate in transitional zones, such as the temperate-boreal forest ecotone, yet previous work in these forests has primarily focused on specific subsets of microbial taxa. Here, we examined how bacterial and fungal communities respond to simulated above- and below-ground warming under realistic field conditions in closed and open canopy treatments in Minnesota, USA. Our results show that warming and canopy disturbance shifted bacterial and fungal community structure as dominant bacterial and fungal groups differed in the direction and intensity of their responses. Ectomycorrhizal and saprotrophic fungal communities with greater connectivity (higher prevalence of strongly interconnected taxa based on pairwise co-occurrence relationships) were more resistant to compositional change. Warming effects on soil enzymes involved in the hydrolytic and oxidative liberation of carbon from plant cell walls and nutrients from organic matter were most strongly linked to fungal community responses, although community structure-function relationships differed between fungal guilds. Collectively, these findings indicate that warming and disturbance will influence the composition and function of microbial communities in the temperate-boreal ecotone, and fungal responses are particularly important to understand for predicting future ecosystem functioning.

Ectomycorrhizal fungal diversity predicted to substantially decline due to climate changes in North American Pinaceae forests

AbstractAimEctomycorrhizal fungi (ECMF) are partners in a globally distributed tree symbiosis implicated in most major ecosystem functions. However, resilience of ECMF to future climates is uncertain. We forecast these changes over the extent of North American Pinaceae forests.LocationAbout 68 sites from North American Pinaceae forests ranging from Florida to Ontario in the east and southern California to Alaska in the west.TaxonEctomycorrhizal fungi (Asco‐ and Basidiomycetes).MethodsWe characterized ECMF communities at each site using molecular methods and modelled climatic drivers of diversity and community composition with general additive, generalized dissimilarity models and Threshold Indicator Taxa ANalysis (TITAN). Next, we projected our models across the extent of North American Pinaceae forests and forecast ECMF responses to climate changes in these forests over the next 50 years.ResultsWe predict median declines in ECMF species richness as high as 26% in Pinaceae forests throughout a climate zone comprising more than 3.5 million square kilometres of North America (an area twice that of Alaska state). Mitigation of greenhouse gas emissions can reduce these declines, but not prevent them. The existence of multiple diversity optima along climate gradients suggest regionally divergent trajectories for North American ECMF, which is corroborated by corresponding ECMF community thresholds identified in TITAN models. Warming of forests along the boreal–temperate ecotone results in projected ECMF species loss and declines in the relative abundance of long‐distance foraging ECMF species, whereas warming of eastern temperate forests has the opposite effect.Main ConclusionsOur results reveal potentially unavoidable ECMF species‐losses over the next 50 years, which is likely to have profound (if yet unclear) effects on ECMF‐associated biogeochemical cycles.

Water quality benefits of Acer rubrum and Betula alleghaniensis in the boreal-temperate ecotone on Eastern Cape Breton Island, Canada

Forest management objectives in public water supply areas are designed to optimize water quality. We employed a paired-tree sampling approach within mixed stands to test the effects of evergreen conifer versus deciduous angiosperm taxa on water chemistry beneath canopies (i.e., throughfall; TF) in a post-industrial region with Ca-impoverished acidic soils and pronounced coastal weather conditions. Reduced concentrations of protons and sea salt were two discernable benefits of deciduous angiosperm trees. These effects became more pronounced during fall recharge when tree physiological dormancy coincided with onshore storm events and increasing seasonal demand for industrial and residential coal combustion. Taxon differences however, did not exert any significant effect on base cation proportions in soil leachates and on soil cation exchange complexes. Thus, ecosystem-based management of stand-level tree diversity in public water supply areas should monitor pH and electrical conductivity of TF water to mitigate adverse water quality effects of persistent acidifying air pollution and the preponderance and high concentration of Na+ in coastal regions. Results largely corroborated previous observations of a decreasing trend in acidifying air pollution at the northeastern periphery of the Eastern Deciduous Forest biome, although elevated SO42−/Cl− – ratios above seawater levels were still noted in TF water chemistry. Electrical conductivity was a reliable predictor of Cl− concentration in TF water regardless of taxon, phenology, and stand location (R2 = 0.99).

Surprising lack of sensitivity of biochemical limitation of photosynthesis of nine tree species to open-air experimental warming and reduced rainfall in a southern boreal forest.

Photosynthetic biochemical limitation parameters (i.e., Vcmax , Jmax and Jmax :Vcmax ratio) are sensitive to temperature and water availability, but whether these parameters in cold climate species at biome ecotones are positively or negatively influenced by projected changes in global temperature and water availability remains uncertain. Prior exploration of this question has largely involved greenhouse based short-term manipulative studies with mixed results in terms of direction and magnitude of responses. To address this question in a more realistic context, we examined the effects of increased temperature and rainfall reduction on the biochemical limitations of photosynthesis using a long-term chamber-less manipulative experiment located in northern Minnesota, USA. Nine tree species from the boreal-temperate ecotone were grown in natural neighborhoods under ambient and elevated (+3.4°C) growing season temperatures and ambient or reduced (≈40% of rainfall removed) summer rainfall. Apparent rubisco carboxylation and RuBP regeneration standardized to 25°C (Vcmax25°C and Jmax25°C , respectively) were estimated based on ACi curves measured in situ over three growing seasons. Our primary objective was to test whether species would downregulate Vcmax25°C and Jmax25°C in response to warming and reduced rainfall, with such responses expected to be greatest in species with the coldest and most humid native ranges, respectively. These hypotheses were not supported, as there were no overall main treatment effects on Vcmax25°C or Jmax25°C (p>.14). However, Jmax :Vcmax ratio decreased significantly with warming (p=.0178), whereas interactions between warming and rainfall reduction on the Jmax25°C to Vcmax25°C ratio were not significant. The insensitivity of photosynthetic parameters to warming contrasts with many prior studies done under larger temperature differentials and often fixed daytime temperatures. In sum, plants growing in relatively realistic conditions under naturally varying temperatures and soil moisture levels were remarkably insensitive in terms of their Jmax25°C and Vcmax25°C when grown at elevated temperatures, reduced rainfall, or both combined.

Photosynthetic and Respiratory Responses of Two Bog Shrub Species to Whole Ecosystem Warming and Elevated CO2 at the Boreal-Temperate Ecotone

Peatlands within the boreal-temperate ecotone contain the majority of terrestrial carbon in this region, and there is concern over the fate of such carbon stores in the face of global environmental changes. The Spruce and Peatland Response Under Changing Environments (SPRUCE) facility aims to advance the understanding of how such peatlands may respond to such changes, using a combination of whole ecosystem warming (WEW; +0, 2.25, 4.5, 6.75 and 9 °C) and elevated CO2 (eCO2; +500 ppm) treatments in an intact bog ecosystem. We examined photosynthetic and respiration responses in leaves of two ericaceous shrub species--leatherleaf (Chamaedaphne calyculata (L.) Moench) and bog Labrador tea (Rhododendron groenlandicum (Oeder) Kron & Judd)--to the first year of combined eCO2 and WEW treatments at SPRUCE. We surveyed the leaf N content per area (Narea), net photosynthesis (AST) and respiration (RD25) at 25°C and 400 ppm CO2 and net photosynthesis at mean growing conditions (AGR) of newly emerged, mature and overwintered leaves. We also measured leaf non-structural carbohydrate content (NSC) in mature leaves. The effects of WEW and eCO2 varied by season and species, highlighting the need to accommodate such variability in modeling this system. In mature leaves, we did not observe a response to either treatment of AST or RD25 in R. groenlandicum, but we did observe a 50% decrease in AST of C. calyculata with eCO2. In mature leaves under eCO2, neither species had increased AGR and both had increases in NSC, indicating acclimation of photosynthesis to eCO2 may be related to source-sink imbalances of carbohydrates. Thus, productivity gains of shrubs under eCO2 may be lower than previously predicted for this site by models not accounting for such acclimation. In newly emerged leaves, AST increased with WEW in R. groenlandicum, but not C. calyculata. Overwintered leaves exhibited a decrease in RD25 for R. groenlandicum and in AST for C. calyculata with increasing WEW, as well as an increase of AGR with eCO2 in both species. Responses in newly emerged and overwintered leaves may reflect physiological acclimation or phenological changes in response to treatments.

Temporal changes in avian community composition in lowland conifer habitats at the southern edge of the boreal zone in the Adirondack Park, NY

Climate change represents one of the most significant threats to human and wildlife communities on the planet. Populations at range margins or transitions between biomes can be particularly instructive for observing changes in biological communities that may be driven by climate change. Avian communities in lowland boreal habitats in the Adirondack Park, located at the North American boreal-temperate ecotone, have been the focus of long-term monitoring efforts since 2007. By documenting long-term changes in community structure and composition, such datasets provide an opportunity to understand how boreal species are responding differently to climate change, and which habitat characteristics may be best able to retain boreal avian communities. We examined three specific questions in order to address how well current biological communities in Adirondack boreal wetland habitats are being maintained in a changing climate: (1) how do trends in occupancy vary across species, and what guilds or characteristics are associated with increasing or decreasing occupancy? (2) how is avian community composition changing differently across sites, and (3) what distinguishes sites which are retaining boreal birds to a higher degree than other sites? Our analysis revealed that (1) boreal species appear to exhibit the largest changes in occupancy among our study locations as compared to the larger avian community, (2) dynamics of community change are not uniform across sites and habitat structure may play an important role in driving observed changes, and (3) the particular characteristics of large open peatlands may allow them to serve as refugia for boreal species in the context of climate change.

Disturbances amplify tree community responses to climate change in the temperate–boreal ecotone

AbstractAimClimate change causes major shifts in species distributions, reshuffling community composition and favouring warm‐adapted species (“thermophilization”). The tree community response is likely to be affected by major disturbances, such as fire and harvest. Here, we quantify the relative contributions of climate change and disturbances to temporal shifts in tree composition over the last decades and evaluate whether disturbances accelerate community thermophilization.LocationQuébec, Canada.Time period1970–2016.Taxa studiedTrees.MethodsUsing 6,281 forest inventory plots, we quantified temporal changes in species composition between a historical (1970–1980) and a contemporary period (2000–2016) by measuring temporal β‐diversity, gains and losses. The effects of climate and disturbances on temporal β‐diversity were quantified using multiple regressions and variation partitioning. We compared how community indices of species temperature preference (CTI) and shade tolerance (CSI) changed for forests that experienced different levels of disturbance. We quantified the contribution of species gains and losses to change in CTI.ResultsTemporal β‐diversity was mainly driven by disturbances, with historical harvesting as the most important predictor. Despite the prevailing influence of disturbances, we revealed a significant thermophilization (ΔCTI = +.03 °C/decade) throughout forests in Québec. However, this shift in community composition was weakly explained by climate change and considerably slower than the rate of warming (+.14 °C/decade). Importantly, thermophilization was amplified by moderate disturbances (+.044 °C/decade), almost a threefold increase compared with minor disturbances (+.015 °C/decade). The gains and losses of a few tree species contributed to this community‐level shift.ConclusionsOur study provides evidence that disturbances can strongly modify tree community responses to climate change. Moderate disturbances, such as harvesting, might reduce competition and facilitate gains of warm‐adapted species, which then accelerate thermophilization of tree communities under climate change. Although accelerated by disturbances, community thermophilization was driven by the gains and losses of a small number of species, notably gains of maples.

Climate change will affect the ability of forest management to reduce gaps between current and presettlement forest composition in southeastern Canada

Forest landscapes at the boreal–temperate ecotone have been extensively altered. Reducing the gap between current and presettlement forest conditions through ecosystem-based forest management (EBFM) is thought to enhance ecological integrity. However, climate change may interfere with this goal and make these targets unrealistic. We evaluated the impacts of climate change on the ability of EBFM to reduce discrepancies between current and presettlement forest conditions in southeastern Canada. We used early-land-survey data as well as projections from a forest landscape model (LANDIS-II) under four climate change scenarios and four management scenarios to evaluate future discrepancies between presettlement forest conditions and future forest landscapes. By triggering swift declines in most late-succession boreal conifer species biomass, climate change would greatly reduce the ability of forest management to reduce the gap with presettlement forest composition, especially under severe anthropogenic climate forcing. Scenarios assuming extensive clearcutting also favor aggressive competitor species that have already increased with high historical harvest levels (e.g., poplars, maples). EBFM would still be the “less bad” forest harvesting strategy in order to mitigate composition discrepancies with the presettlement forests, though it is likely to fail under severe climate forcing. In this latter case, one might thus question the relevancy of using presettlement forest composition as a target for restoring degraded forest landscapes. As such, we advocate that managers should relax the centrality of the reference condition and focus on functional restoration rather than aiming at reducing the gaps with presettlement forest composition per se.

Biodiversity bottleneck: seedling establishment under changing climatic conditions at the boreal–temperate ecotone

Climate change is increasing global temperatures, severe rainfall events, and the occurrence and severity of drought. Changes in global climate may have negative consequences for particular plant species and for biodiversity overall. In the short term, altered temperature and precipitation regimes may have the most severe effects on plant species near their range limits and in the earliest stages of plant development. To address these issues, we assessed seedling emergence, early survival, and growth of 18 boreal, temperate, and exotic woody species at the boreal–temperate forest ecotone in central Minnesota. We experimentally warmed forest plots to mimic projected warming by the end of the twenty-first century (+ 1.7 °C and + 3.4 °C). We also experimentally removed summer rainfall (~ 42% reduction) to simulate drought conditions in this region. We found that emergence and survival of boreal and exotic species was lower in experimentally warmed plots. This was exacerbated by drought. Temperate species emergence and survival was largely unaffected by climate manipulations (on average). Conversely, temperate seedling growth was greater in warmer conditions, but only when paired with drought. We found that overall seedling species richness was reduced by warming, mostly due to lower boreal and exotic species emergence and survival (conifers were also strongly negatively affected across species-range groups). If temperate seedling emergence and survival does not compensate for loss of boreal species, these forests may experience loss of biodiversity (and associated ecosystem functions) in the future.

Identifying environmental drivers of greenhouse gas emissions under warming and reduced rainfall in boreal–temperate forests

Abstract Atmospheric concentrations of carbon dioxide (CO2), methane (CH4) and nitrous oxide (N2O) are predicted to increase as a consequence of fossil fuel emissions and the impact on biosphere–atmosphere interactions. Forest ecosystems in general, and forest soils in particular, can be sinks or sources for CO2, CH4, and N2O. Environmental studies traditionally target soil temperature and moisture as the main predictors of soil greenhouse gas (GHG) flux from different ecosystems; however, these emissions are primarily biologically driven. Thus, little is known about the degree of regulation by soil biotic vs. abiotic factors on GHG emissions, particularly under predicted increase in global temperatures, and changes in intensity and frequency of precipitation events. Here we measured net CO2, CH4 and N2O fluxes after 5 years of experimental warming (+3.4°C), and 2 years of ≈45% summer rainfall reduction, in two forest sites in a boreal–temperate ecotone under different habitat conditions (closed or open canopy) in Minnesota, USA. We evaluated the importance of microbial gene abundance and climo‐edaphic factors (soil texture, canopy, seasonality, climate, and soil physicochemical properties) driving GHG emissions. We found that changes in CO2 fluxes were predominantly determined abiotically by temperature and moisture, after accounting for bacterial abundance. Methane fluxes on the other hand, were determined both abiotically, by gas diffusivity (via soil texture) and microbially, by methanotroph pmoA gene abundance, whereas, N2O emissions showed only a strong biotic regulation via ammonia‐oxidizing bacteria amoA gene abundance. Warming did not significantly alter CO2 and CH4 fluxes after 5 years of manipulation, while N2O emissions were greater with warming under open canopy. Our findings provide evidence that soil GHG emissions result from multiple direct and indirect interactions of microbial and abiotic drivers. Overall, this study highlights the need to include both microbial and climo‐edaphic properties in predictive models in order to provide improved mechanistic understanding for the development of future mitigation strategies. A plain language summary is available for this article.

Where does the carbon go? Thermal acclimation of respiration and increased photosynthesis in trees at the temperate-boreal ecotone.

Where does the carbon go? Thermal acclimation of respiration and increased photosynthesis in trees at the temperate-boreal ecotone.

Ectomycorrhizal fungal response to warming is linked to poor host performance at the boreal-temperate ecotone.

Rising temperatures associated with climate change have been shown to negatively affect the photosynthetic rates of boreal forest tree saplings at their southern range limits. To quantify the responses of ectomycorrhizal (EM) fungal communities associated with poorly performing hosts, we sampled the roots of Betula papyrifera and Abies balsamea saplings growing in the B4Warmed (Boreal Forest Warming at an Ecotone in Danger) experiment. EM fungi on the root systems of both hosts were compared from ambient and +3.4°C air and soil warmed plots at two sites in northern Minnesota. EM fungal communities were assessed with high-throughput sequencing along with measures of plant photosynthesis, soil temperature, moisture, and nitrogen. Warming selectively altered EM fungal community composition at both the phylum and genus levels, but had no significant effect on EM fungal operational taxonomic unit (OTU) diversity. Notably, warming strongly favored EM Ascomycetes and EM fungi with short-contact hyphal exploration types. Declining host photosynthetic rates were also significantly inversely correlated with EM Ascomycete and EM short-contact exploration type abundance, which may reflect a shift to less carbon demanding fungi due to lower photosynthetic capacity. Given the variation in EM host responses to warming, both within and between ecosystems, better understanding the link between host performance and EM fungal community structure will to clarify how climate change effects cascade belowground.