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Photosynthetic acclimation to both warming and elevated CO2 of boreal trees remains a key uncertainty in modelling the response of photosynthesis to future climates. We investigated the impact of increased growth temperature and elevated CO2 on photosynthetic capacity (Vcmax and Jmax) in mature trees of two North American boreal conifers, tamarack and black spruce. We show that Vcmax and Jmax at a standard temperature of 25°C did not change with warming, while Vcmax and Jmax at their thermal optima (Topt) and growth temperature (Tg) increased. Moreover, Vcmax and Jmax at either 25°C, Topt or Tg decreased with elevated CO2. The Jmax/Vcmax ratio decreased with warming when assessed at both Topt and Tg but did not significantly vary at 25°C. The Jmax/Vcmax increased with elevated CO2 at either reference temperature. We found no significant interaction between warming and elevated CO2 on all traits. If this lack of interaction between warming and elevated CO2 on the Vcmax, Jmax and Jmax/Vcmax ratio is a general trend, it would have significant implications for improving photosynthesis representation in vegetation models. However, future research is required to investigate the widespread nature of this response in a larger number of species and biomes.

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Tree mortality due to global change-including range expansion of invasive pests and pathogens-is a paramount threat to forest ecosystems. Oak forests are among the most prevalent and valuable ecosystems both ecologically and economically in the United States. There is increasing interest in monitoring oak decline and death due to both drought and the oak wilt pathogen (Bretziella fagacearum). We combined anatomical and ecophysiological measurements with spectroscopy at leaf, canopy, and airborne levels to enable differentiation of oak wilt and drought, and detection prior to visible symptom appearance. We performed an outdoor potted experiment with Quercus rubra saplings subjected to drought stress and/or artificially inoculated with the pathogen. Models developed from spectral reflectance accurately predicted ecophysiological indicators of oak wilt and drought decline in both potted and field experiments with naturally grown saplings. Both oak wilt and drought resulted in blocked water transport through xylem conduits. However, oak wilt impaired conduits in localized regions of the xylem due to formation of tyloses instead of emboli. The localized tylose formation resulted in more variable canopy photosynthesis and water content in diseased trees than drought-stressed ones. Reflectance signatures of plant photosynthesis, water content, and cellular damage detected oak wilt and drought 12 d before visual symptoms appeared. Our results show that leaf spectral reflectance models predict ecophysiological processes relevant to detection and differentiation of disease and drought. Coupling spectral models that detect physiological change with spatial information enhances capacity to differentiate plant stress types such as oak wilt and drought.

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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.

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Warming shifts the thermal optimum of net photosynthesis (ToptA) to higher temperatures. However, our knowledge of this shift is mainly derived from seedlings grown in greenhouses under ambient atmospheric carbon dioxide (CO2) conditions. It is unclear whether shifts in ToptA of field-grown trees will keep pace with the temperatures predicted for the 21st century under elevated atmospheric CO2 concentrations. Here, using a whole-ecosystem warming controlled experiment under either ambient or elevated CO2 levels, we show that ToptA of mature boreal conifers increased with warming. However, shifts in ToptA did not keep pace with warming as ToptA only increased by 0.26-0.35 °C per 1 °C of warming. Net photosynthetic rates estimated at the mean growth temperature increased with warming in elevated CO2 spruce, while remaining constant in ambient CO2 spruce and in both ambient CO2 and elevated CO2 tamarack with warming. Although shifts in ToptA of these two species are insufficient to keep pace with warming, these boreal conifers can thermally acclimate photosynthesis to maintain carbon uptake in future air temperatures.

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The linkage of stomatal behaviour with photosynthesis is critical to understanding water and carbon cycles under global change. The relationship of stomatal conductance (gs ) and CO2 assimilation (Anet ) across a range of environmental contexts, as represented in the model parameter (g1 ), has served as a proxy of the marginal water cost of carbon acquisition. We use g1 to assess species differences in stomatal behaviour to a decade of open-air experimental climate change manipulations, asking whether generalisable patterns exist across species and climate contexts. Anet -gs measurements (17 727) for 21 boreal and temperate tree species under ambient and +3.3°C warming, and ambient and ~40% summer rainfall reduction, provided >2700 estimates of g1 . Warming and/or reduced rainfall treatments both lowered g1 because those treatments resulted in lower soil moisture and because stomatal behaviour changed more in warming when soil moisture was low. Species tended to respond similarly, although, in species from warmer and drier habitats, g1 tended to be slightly higher and to be the least sensitive to the decrease in soil water. Overall, both warming and rainfall reduction consistently made stomatal behaviour more conservative in terms of water loss per unit carbon gain across 21 species and a decade of experimental observation.

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The sensitivity of forests to near-term warming and associated precipitation shifts remains uncertain1-9. Herein, using a 5-year open-air experiment in southern boreal forest, we show divergent responses to modest climate alteration among juveniles of nine co-occurring North American tree species. Warming alone (+1.6 °C or +3.1 °C above ambient temperature) or combined with reduced rainfall increased the juvenile mortality of all species, especially boreal conifers. Species differed in growth responses to warming, ranging from enhanced growth in Acer rubrum and Acer saccharum to severe growth reductions in Abies balsamea, Picea glauca and Pinus strobus. Moreover, treatment-induced changes in both photosynthesis and growth help explain treatment-driven changes in survival. Treatments in which species experienced conditions warmer or drier than at their range margins resulted in the most adverse impacts on growth and survival. Species abundant in southern boreal forests had the largest reductions in growth and survival due to climate manipulations. By contrast, temperate species that experienced little mortality and substantial growth enhancement in response to warming are rare throughout southern boreal forest and unlikely to rapidly expand their density and distribution. Therefore, projected climate change will probably cause regeneration failure of currently dominant southern boreal species and, coupled with their slow replacement by temperate species, lead to tree regeneration shortfalls with potential adverse impacts on the health, diversity and ecosystem services of regional forests.

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Plants often adjust their leaf mitochondrial ("dark") respiration (Rd ) measured at a standardized temperature such as 20°C (R20 ) downward after experiencing warmer temperatures and upward after experiencing cooler temperatures. These responses may help leaves maintain advantageous photosynthetic capacity and/or be a response to recent photosynthate accumulation, and can occur within days after a change in thermal regime. It is not clear, however, how the sensitivity and magnitude of this response change over time, or which time period prior to a given measurement best predicts R20 . Nor is it known whether nighttime, daytime, or 24-hour temperatures should be most influential. To address these issues, we used data from 1620 Rd temperature response curves of 10 temperate and boreal tree species in a long-term field experiment in Minnesota, USA to assess how the observed nearly complete acclimation of R20 was related to past temperatures during periods of differing lengths. We hypothesized that R20 would be best related to prior midday temperatures associated with both photosynthetic biochemistry and peak carbon uptake rates that drive carbohydrate accumulation. Inconsistent with this hypothesis, prior night temperatures were the best predictors of R20 for all species. We had also hypothesized that recent (prior 3-10 days) temperatures should best predict R20 because they likely have stronger residual impacts on leaf-level physiology than periods extending further back in time, whereas a prior 1- to 2-day period might be a span shorter than one to which photosynthetic capacity and Rd adjust. There was little to no support for this idea, as for angiosperms, long time windows (prior 30-60 nights) were the best predictors, while for gymnosperms both near-term (prior 3-8 nights for pines, prior 10-14 nights for spruce/fir) and longer-term periods (prior 45 nights) were the best predictors. The importance of nighttime temperatures, the relatively long "time-averaging" that best explained acclimation, and dual peaks of temporal acclimation responsiveness in some species were all results that were unanticipated.

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Global change is shifting disturbance regimes that may rapidly change ecosystems, sometimes causing ecosystems to shift between states. Interactions between disturbances such as fire and disease could have especially severe effects, but experimental tests of multi-decadal changes in disturbance regimes are rare. Here, we surveyed vegetation for 35 years in a 54-year fire frequency experiment in a temperate oak savanna-forest ecotone that experienced a recent outbreak of oak wilt. Different fire regimes determined whether plots were savanna or forest by regulating tree abundance (r2  = 0.70), but disease rapidly reversed the effect of fire exclusion, increasing mortality by 765% in unburned forests, but causing relatively minor changes in frequently burned savannas. Model simulations demonstrated that disease caused unburned forests to transition towards a unique woodland that was prone to transition to savanna if fire was reintroduced. Consequently, disease-fire interactions could shift ecosystem resilience and biome boundaries as pathogen distributions change.

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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.

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BACKGROUND AND AIMS: Warmer temperatures and altered precipitation patterns are expected to continue to occur as the climate changes. How these changes will impact the flowering phenology of herbaceous perennials in northern forests is poorly understood but could have consequences for forest functioning and species interactions. Here, we examine the flowering phenology responses of five herbaceous perennials to experimental warming and reduced summer rainfall over 3 years. METHODS: This study is part of the B4WarmED experiment located at two sites in northern Minnesota, USA. Three levels of warming (ambient, +1.6 °C and +3.1 °C) were crossed with two rainfall manipulations (ambient and 27 % reduced growing season rainfall). KEY RESULTS: We observed species-specific responses to the experimental treatments. Warming alone advanced flowering for four species. Most notably, the two autumn blooming species showed the strongest advance of flowering to warming. Reduced rainfall alone advanced flowering for one autumn blooming species and delayed flowering for the other, with no significant impact on the three early blooming species. Only one species, Solidago spp., showed an interactive response to warming and rainfall manipulation by advancing in +1.6 °C warming (regardless of rainfall manipulation) but not advancing in the warmest, driest treatment. Species-specific responses led to changes in temporal overlap between species. Most notably, the two autumn blooming species diverged significantly in their flowering timing. In ambient conditions, these two species flowered within the same week. In the warmest, driest treatment, flowering occurred over a month apart. CONCLUSIONS: Herbaceous species may differ in how they respond to future climate conditions. Changes to phenology may lead to fewer resources for insects or a mismatch between plants and pollinators.

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Changes in plant phenology associated with climate change have been observed globally. What is poorly known is whether and how phenological responses to climate warming will differ from year to year, season to season, habitat to habitat, or species to species. Here, we present 5 y of phenological responses to experimental warming for 10 subboreal tree species. Research took place in the open-air B4WarmED experiment in Minnesota. The design is a two habitat (understory and open) × three warming treatments (ambient, +1.7 °C, +3.4 °C) factorial at two sites. Phenology was measured twice weekly during the growing seasons of 2009 through 2013. We found significant interannual variation in the effect of warming and differences among species in response to warming that relate to geographic origin and plant functional group. Moreover, responses to experimental temperature variation were similar to responses to natural temperature variation. Warming advanced the date of budburst more in early compared to late springs, suggesting that to simulate interannual variability in climate sensitivity of phenology, models should employ process-based or continuous development approaches. Differences among species in timing of budburst were also greater in early compared to late springs. Our results suggest that climate change-which will make most springs relatively "early"-could lead to a future with more variable phenology among years and among species, with consequences including greater risk of inappropriately early leafing and altered interactions among species.

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Hyperspectral reflectance tools have been used to detect multiple pathogens in agricultural settings and single sources of infection or broad declines in forest stands. However, differentiation of any one disease from other sources of tree stress is integral for stand and landscape-level applications in mixed species systems. We tested the ability of spectral models to differentiate oak wilt, a fatal disease in oaks caused by Bretziella fagacearum ``Bretz'', from among other mechanisms of decline. We subjected greenhouse-grown oak seedlings (Quercus ellipsoidalis ``E.J. Hill'' and Quercus macrocarpa ``Michx.'') to chronic drought or inoculation with the oak wilt fungus or bur oak blight fungus (Tubakia iowensis ``T.C. Harr. & D. McNew''). We measured leaf and canopy spectroscopic reflectance (400-2400 nm) and instantaneous photosynthetic and stomatal conductance rates, then used partial least-squares discriminant analysis to predict treatment from hyperspectral data. We detected oak wilt before symptom appearance, and classified the disease with high accuracy in symptomatic leaves. Classification accuracy from spectra increased with declines in photosynthetic function in oak wilt-inoculated plants. Wavelengths diagnostic of oak wilt were only found in non-visible spectral regions and are associated with water status, non-structural carbohydrates and photosynthetic mechanisms. We show that hyperspectral models can differentiate oak wilt from other causes of tree decline and that detection is correlated with biological mechanisms of oak wilt infection and disease progression. We also show that within the canopy, symptom heterogeneity can reduce detection, but that symptomatic leaves and tree canopies are suitable for highly accurate diagnosis. Remote application of hyperspectral tools can be used for specific detection of disease across a multi-species forest stand exhibiting multiple stress symptoms.

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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.

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Widespread changes in arctic and boreal Normalized Difference Vegetation Index (NDVI) values captured by satellite platforms indicate that northern ecosystems are experiencing rapid ecological change in response to climate warming. Increasing temperatures and altered hydrology are driving shifts in ecosystem biophysical properties that, observed by satellites, manifest as long-term changes in regional NDVI. In an effort to examine the underlying ecological drivers of these changes, we used field-scale remote sensing of NDVI to track peatland vegetation in experiments that manipulated hydrology, temperature, and carbon dioxide (CO2 ) levels. In addition to NDVI, we measured percent cover by species and leaf area index (LAI). We monitored two peatland types broadly representative of the boreal region. One site was a rich fen located near Fairbanks, Alaska, at the Alaska Peatland Experiment (APEX), and the second site was a nutrient-poor bog located in Northern Minnesota within the Spruce and Peatland Responses Under Changing Environments (SPRUCE) experiment. We found that NDVI decreased with long-term reductions in soil moisture at the APEX site, coincident with a decrease in photosynthetic leaf area and the relative abundance of sedges. We observed increasing NDVI with elevated temperature at the SPRUCE site, associated with an increase in the relative abundance of shrubs and a decrease in forb cover. Warming treatments at the SPRUCE site also led to increases in the LAI of the shrub layer. We found no strong effects of elevated CO2 on community composition. Our findings support recent studies suggesting that changes in NDVI observed from satellite platforms may be the result of changes in community composition and ecosystem structure in response to climate warming.

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Climate warming will influence photosynthesis via thermal effects and by altering soil moisture1-11. Both effects may be important for the vast areas of global forests that fluctuate between periods when cool temperatures limit photosynthesis and periods when soil moisture may be limiting to carbon gain4-6,9-11. Here we show that the effects of climate warming flip from positive to negative as southern boreal forests transition from rainy to modestly dry periods during the growing season. In a three-year open-air warming experiment with juveniles of 11 temperate and boreal tree species, an increase of 3.4 °C in temperature increased light-saturated net photosynthesis and leaf diffusive conductance on average on the one-third of days with the wettest soils. In all 11 species, leaf diffusive conductance and, as a result, light-saturated net photosynthesis decreased during dry spells, and did so more sharply in warmed plants than in plants at ambient temperatures. Consequently, across the 11 species, warming reduced light-saturated net photosynthesis on the two-thirds of days with driest soils. Thus, low soil moisture may reduce, or even reverse, the potential benefits of climate warming on photosynthesis in mesic, seasonally cold environments, both during drought and in regularly occurring, modestly dry periods during the growing season.

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PREMISE OF THE STUDY: Changes to plant phenology have been linked to warmer temperatures caused by climate change. Despite the importance of the groundlayer to community and forest dynamics, few warming experiments have focused on herbaceous plant and shrub phenology. METHODS: Using a field study in Minnesota, United States, we investigated phenological responses of 16 species to warming over five growing seasons (2009-2013) at two sites, under two canopy covers, and in three levels of simultaneous above- and belowground warming: ambient temperature, ambient +1.7°C and ambient +3.4°C. We tested whether warming led to earlier phenology throughout the growing season and whether responses varied among species and years and depended on canopy cover. KEY RESULTS: Warming extended the growing season between 11-30 days, primarily through earlier leaf unfolding. Leaf senescence was delayed for about half of the species. Warming advanced flowering across species, especially those flowering in August, with modest impacts on fruit maturation for two species. Importantly, warming caused more than half of the species to either converge or diverge phenologically in relation to each other, suggesting that future warmed climate conditions will alter phenological relationships of the groundlayer. Warm springs elicited a stronger advance of leaf unfolding compared to cool spring years. Several species advanced leaf unfolding (in response to warming) more in the closed canopy compared to the open. CONCLUSIONS: Climate warming will extend the growing season of groundlayer species in the boreal-temperate forest ecotone and alter the synchrony of their phenology.

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The phenology of trees is highly susceptible to changing global temperatures. Leaf budburst advances with increasing spring temperatures, but can also be delayed when warmer winters reduce chilling exposure. Results from long-term observations show that increasing temperatures have triggered advanced budburst in the past decades, but some studies also show that budburst advance has slowed recently. Here, we conducted an experiment with five temperate deciduous tree species (Acer rubrum L., Larix laricina (Du Rois) K. Koch, Populus tremuloides Michx., Quercus ellipsoidalis E. j. Hill, Betula papyrifera Marsh.) and one invasive species (Rhamnus cathartica L.) in Minnesota, USA, to assess the impact of chilling on the timing of leaf budburst. We collected twigs over two winter seasons (2011/2012 and 2012/2013) on a biweekly basis and exposed them to spring-like temperatures of 21 °C/16 °C day and night, long day photoperiod (16 h). We found a significant relationship of advanced budbreak with increased chilling for all species tested (P < 0.001) and significant differences in the timing to budburst among all species (P < 0.001). Acer rubrum responded strongly to chilling, showing a very steep linear decline in days to budburst with increased exposure to chilling. On the other end of the spectrum, L. laricina responded least to increases in chilling. These results suggest that rising global temperatures will likely have diverse impacts on tree species with potential implications for species interactions such as competition.

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Over the last two decades, empirical work has established that higher biodiversity can lead to greater primary productivity; however, the importance of different aspects of biodiversity in contributing to such relationships is rarely elucidated. We assessed the relative importance of species richness, phylogenetic diversity, functional diversity, and identity of neighbors for stem growth 3 yr after seedling establishment in a tree diversity experiment in eastern Minnesota. Generally, we found that community-weighted means of key functional traits (including mycorrhizal association, leaf nitrogen and calcium, and waterlogging tolerance) as well as species richness were strong, independent predictors of stem biomass growth. More phylogenetically diverse communities did not consistently produce more biomass than expected, and the trait values or diversity of individual functional traits better predicted biomass production than did a multidimensional functional diversity metric. Furthermore, functional traits and species richness best predicted growth at the whole-plot level (12 m2 ), whereas neighborhood composition best predicted growth at the focal tree level (0.25 m2 ). The observed effects of biodiversity on growth appear strongly driven by positive complementary effects rather than by species-specific selection effects, suggesting that synergistic species' interactions rather than the influence of a few important species may drive overyielding.

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Plant diversity experiments generally find that increased diversity causes increased productivity; however, primary productivity is typically measured in the presence of a diverse food web, including pathogens, mutualists and herbivores. If food web impacts on productivity vary with plant diversity, as predicted by both theoretical and empirical studies, estimates of the effect of plant diversity on productivity may be biased. We experimentally removed arthropods, foliar fungi and soil fungi from the longest-running plant diversity experiment. We found that fungi and arthropods removed a constant, large proportion of biomass leading to a greater reduction of total biomass in high diversity plots. As a result, the effect of diversity on measured plant productivity was much higher in the absence of fungi and arthropods. Thus, diversity increases productivity more than reported in previous studies that did not control for the effects of heterotrophic consumption.

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