Cultivation of Non‐timber Forest Products Alters Understory Light Availability in a Humid Tropical Forest in Mexico1

The planting of non‐timber forest products (NTFPs) in the understory of tropical forests is promoted in many regions as a strategy to conserve forested lands and meet the economic needs of rural communities. While the forest canopy is left intact in most understory plantations, much of the midstory and understory vegetation is removed in order to increase light availability for cultivated species. We assessed the extent to which the removal of vegetation in understory plantations of Chamaedorea hooperiana Hodel (Arecaceae) alters understory light conditions. We also examined how any changes in light availability may be reflected by changes in the composition of canopy tree seedlings regenerating in understory plantations. We employed a blocked design consisting of four C. hooperiana plantation sites; each site was paired with an adjacent, unmanaged forest site. Hemispherical canopy photographs were taken and canopy tree seedlings were identified and measured within 12 3 × 2 m randomly placed plots in each site for a total of 96 plots (4 blocks × 2 sites × 12 plots). Plantation management did not affect canopy openness or direct light availability but understory plantations had a higher frequency of plots with greater total and diffuse light availability than unmanaged forest. Comparisons of canopy tree seedling composition between understory plantations and unmanaged forest sites were less conclusive but suggest that management practices have the potential to increase the proportion of shade‐intolerant species of tree seedlings establishing in plantations. Given the importance of advanced regeneration in gap‐phase forest dynamics, these changes may have implications for future patterns of succession in the areas of forest where NTFPs are cultivated.     

Forest phenology and a warmer climate – growing season extension in relation to climatic provenance

Predicting forest responses to warming climates relies on assumptions about niche and temperature sensitivity that remain largely untested. Observational studies have related current and historical temperatures to phenological shifts, but experimental evidence is sparse, particularly for autumn responses. A 4 year field experiment exposed four deciduous forest species from contrasting climates (Liquidambar styraciflua, Quercus rubra, Populus grandidentata, and Betula alleghaniensis) to air temperatures 2 and 4 °C above ambient controls, using temperature‐controlled open top chambers. Impacts of year‐round warming on bud burst (BB), senescence, and abscission were evaluated in relation to thermal provenance. Leaves emerged earlier in all species by an average of 4–9 days at +2 °C and 6–14 days at +4 °C. Magnitude of advance varied with species and year, but was larger for the first 2 °C increment than for the second. Effect of warming increased with early BB, favoring Liquidambar, but even BB of northern species advanced, despite temperatures exceeding those of the realized niche. Treatment differences in BB were inadequately explained by temperature sums alone. In autumn, chlorophyll was retained an average of 4 and 7 days longer in +2 and +4 °C treatments, respectively, and abscission delayed by 8 and 13 days. Growing seasons in the warmer atmospheres averaged 5–18 days (E2) and 6–28 days (E4) longer, according to species, with the least impact in Quercus. Results are compared with a 16 years record of canopy onset and offset in a nearby upland deciduous forest, where BB showed similar responsiveness to spring temperatures (2–4 days °C^?1). Offset dates in the stand tracked August–September temperatures, except when late summer drought caused premature senescence. The common garden‐like experiment provides evidence that warming alone extends the growing season, at both ends, even if stand‐level impacts may be complicated by variation in other environmental factors.     

Effects of growth temperature and winter duration on leaf phenology of a spring ephemeral (Gagea lutea) and a summergreen forb (Maianthemum dilatatum)

Effects of growth temperature and winter duration on leaf longevity were compared between a spring ephemeral, Gagea lutea, and a forest summergreen forb, Maianthemum dilatatum. The plants were grown at day/night temperatures of 25/20°C and 15/10°C after a chilling treatment for variable periods at 2°C. The temperature regime of 25/20°C was much higher than the mean air temperatures for both species in their native habitats. Warm temperature of 25/20°C and/or long chilling treatment shortened leaf longevity in G. lutea, but not in M. dilatatum. The response of G. lutea was consistent with that reported for other spring ephemerals. Air temperature increases as the vegetative season progresses. The decrease in leaf longevity in G. lutea under warm temperature condition ensures leaf senescence in summer, an unfavorable season for its growth. This also implies that early leaf senescence could occur in years with early summers. Warm spring temperatures have been shown to accelerate the leafing-out of forest trees. The decrease in leaf longevity due to warm temperature helps synchronize the period of leaf senescence roughly with the time of the forest canopy leaf-out. Prolonged winter due to late snowmelt has been shown to shorten the vegetative period for spring ephemerals. The decrease in leaf longevity due to long chilling treatment would correspond with this shortened vegetative period.     

Freeze-protection of overwintering monarch butterflies in Mexico: critical role of the forest as a blanket and an umbrella

Abstract. 1. At their high-altitude overwintering sites in Mexico, monarch butterflies frequently are subjected to sub-zero°C temperatures during December-March. Although monarchs have moderate supercooling ability, two ecological factors strongly influence their capacity to resist freezing: wetting and exposure to the clear night sky. 2. As shown in Fig. 2, 50% of a population of butterflies with water on their body surfaces freeze at warmer sub-zero temperatures (-4.2°C) compared to butterflies with no water on their bodies (-7.7°C). 100% mortality occurs, respectively, at ?7.7°C and ?15°C. 3. Comparative measurements of rainfall within a large overwintering colony in Mexico indicated that the intact canopy acts as an umbrella that reduces butterfly wetting during winter storms. 4. Variable experimental exposure of butterflies to the clear night sky indicated that openings in the forest canopy increases radiational cooling and causes monarch body temperatures to drop as much as 4°C below ambient air temperature. Monarchs under dense cover had body temperatures approximately the same as the ambient air temperature, but more exposed individuals had body temperatures below ambient in direct proportion to the degree of exposure. Consequently, forest thinning increases the probability that the butterflies will freeze to death. 5. Whereas both wetting and exposure are increased by disturbance of the forest canopy, the interaction of these two factors exacerbates freezing mortality during winter storms: 50% of dry and unexposed butterflies froze at ?8°C, whereas wetted and fully exposed butterflies froze at only ?0.5°C. 6. Butterflies inside and on the bottom of the fir bough clusters are better protected from wetting than those on the outside. This supports the hypothesis that the structure of the butterfly clusters has evolved through individual selection to avoid wetting. 7. The data strongly reinforce previous evidence that forest thinning should be totally prevented within and adjacent to the overwintering sites in order to minimize both wetting and exposure of the butterflies that synergistically increase winter mortality at the overwintering sites in Mexico.     

Experimental warming alters spring phenology of certain plant functional groups in an early successional forest community

Experimental study of the effects of projected climate change on plant phenology allows us to isolate effects of warming on life‐history events such as leaf out. We simulated a 2 °C temperature increase and 20% precipitation increase in a recently harvested temperate deciduous forest community in central Pennsylvania, USA, and observed the leaf out phenology of all species in 2009 and 2010. Over 130 plant species were monitored weekly in study plots, but due to high variability in species composition among plots, species were grouped into five functional groups: short forbs, tall forbs, shrubs, small trees, and large trees. Tall forbs and large trees, which usually emerge in the late spring, advanced leaf out 14–18 days in response to warming. Short forbs, shrubs, and small trees emerge early in spring and did not alter their phenology in response to warming or increased precipitation treatments. Earlier leaf out of tall forbs and large trees coincided with almost 3 weeks of increased community‐level leaf area index, indicating greater competition and a condensed spring green‐up period. While phenology of large trees and tall forbs appears to be strongly influenced by temperature‐based growth cues, our results suggest that photoperiod and chilling cues more strongly influence the leaf out of other functional groups. Reduced freeze events and warmer temperatures from predicted climate change will interact with nontemperature growth cues to have cascading consequences throughout the ecosystem.     

Tree species growth response to climate warming varies by forest canopy position in boreal and temperate forests

Reports of forest sensitivity to climate change are based largely on the study of overstory trees, which contribute significantly to forest growth and wood supply. However, juveniles in the understory are also critical to predict future forest dynamics and demographics, but their sensitivity to climate remains less known. In this study, we applied boosted regression tree analysis to compare the sensitivity of understory and overstory trees for the 10 most common tree species in eastern North America using growth information from an unprecedented network of nearly 1.5 million tree records from 20,174 widely distributed, permanent sample plots across Canada and the United States. Fitted models were then used to project the near-term (2041–2070) growth for each canopy and tree species. We observed an overall positive effect of warming on tree growth for both canopies and most species, leading to an average of 7.8%–12.2% projected growth gains with climate change under RCP 4.5 and 8.5. The magnitude of these gains peaked in colder, northern areas for both canopies, while growth declines are projected for overstory trees in warmer, southern regions. Relative to overstory trees, understory tree growth was less positively affected by warming in northern regions, while displaying more positive responses in southern areas, likely driven by the buffering effect of the canopy from warming and climate extremes. Observed differences in climatic sensitivity between canopy positions underscore the importance of accounting for differential growth responses to climate between forest strata in future studies to improve ecological forecasts. Furthermore, latitudinal variation in the differential sensitivity of forest strata to climate reported here may help refine our comprehension of species range shift and changes in suitable habitat under climate change.     

Nematode community shifts in response to experimental warming and canopy conditions are associated with plant community changes in the temperate-boreal forest ecotone

Global climate warming is one of the key forces driving plant community shifts, such as range shifts of temperate species into boreal forests. As plant community shifts are slow to observe, ecotones, boundaries between two ecosystems, are target areas for providing early evidence of ecological responses to warming. The role of soil fauna is poorly explored in ecotones, although their positive and negative effects on plant species can influence plant community structure. We studied nematode communities in response to experimental warming (ambient, +1.7, +3.4 °C) in soils of closed and open canopy forest in the temperate-boreal ecotone of Minnesota, USA and calculated various established nematode indices. We estimated species-specific coverage of understory herbaceous and shrub plant species from the same experimental plots and tested if changes in the nematode community are associated with plant cover and composition. Individual nematode trophic groups did not differ among warming treatments, but the ratio between microbial-feeding and plant-feeding nematodes increased significantly and consistently with warming in both closed and open canopy areas and at both experimental field sites. The increase in this ratio was positively correlated with total cover of understory plant species, perhaps due to increased predation pressure on soil microorganisms causing higher nutrient availability for plants. Multivariate analyses revealed that temperature treatment, canopy conditions and nematode density consistently shaped understory plant communities across experimental sites. Our findings suggest that warming-induced changes in nematode community structure are associated with shifts in plant community composition and productivity in the temperate-boreal forest ecotones.     

Shifts in the temperature‐sensitive periods for spring phenology in European beech and pedunculate oak clones across latitudes and over recent decades

Spring phenology of temperate trees has advanced worldwide in response to global warming. However, increasing temperatures may not necessarily lead to further phenological advance, especially in the warmer latitudes because of insufficient chilling and/or shorter day length. Determining the start of the forcing phase, that is, when buds are able to respond to warmer temperatures in spring, is therefore crucial to predict how phenology will change in the future. In this study, we used 4,056 leaf‐out date observations during the period 1969–2017 for clones of European beech (Fagus sylvatica L.) and pedunculate oak (Quercus robur L.) planted in 63 sites covering a large latitudinal gradient (from Portugal ~41°N to Norway ~63°N) at the International Phenological Gardens in order to (a) evaluate how the sensitivity periods to forcing and chilling have changed with climate warming, and (b) test whether consistent patterns occur along biogeographical gradients, that is, from colder to warmer environments. Partial least squares regressions suggest that the length of the forcing period has been extended over the recent decades with climate warming in the colder latitudes but has been shortened in the warmer latitudes for both species, with a more pronounced shift for beech. We attribute the lengthening of the forcing period in the colder latitudes to earlier opportunities with temperatures that can promote bud development. In contrast, at warmer or oceanic climates, the beginning of the forcing period has been delayed, possibly due to insufficient chilling. However, in spite of a later beginning of the forcing period, spring phenology has continued to advance at these areas due to a faster satisfaction of heat requirements induced by climate warming. Overall, our results support that ongoing climate warming will have different effects on the spring phenology of forest trees across latitudes due to the interactions between chilling, forcing and photoperiod.     

Changes in the onset of spring growth in shrubland species in response to experimental warming along a north–south gradient in Europe

Aim To test whether the onset of spring growth in European shrublands is advanced in response to the warmer conditions projected for the next two decades by climate models, and, if there is a change, whether it differs across Europe. Location The studied sites spanned a broad north–south European gradient with average annual temperatures (8.2–15.6 °C) and precipitation (511–1427 mm). Methods ‘Bud break’ was monitored in eight shrub and grass species in six European sites under control and experimentally warmer conditions generated by automatic roofs covering vegetation during the night. Results Species responsive to increased temperatures were Vaccinium myrtillus and Empetrum nigrum in Wales, Deschampsia flexuosa in Denmark, Calluna vulgaris in Netherlands, Populus alba in Hungary and Erica multiflora in Spain. Although the acceleration of spring growth was the commonest response to warming treatments, the responses at each site were species specific and year dependent. Under experimental warming 25% of cases exhibited a significantly earlier onset of the growing season and 10% had a significantly delayed onset of vegetative growth. No geographical gradient was detected in the experimental warming effects. However, there was a trend towards a greater dominance of phenological advances with more intense the warming treatments. Above 0.8 °C warming, only advancements were recorded. Main conclusions Our results show that warmer temperatures projected for the next decades have substantial potential effects on the phenology of the spring growth of dominant species in different European shrublands, with a dominant trend towards advancements the more intense the warming is. However, our study also demonstrates the overall difficulties of applying simple predictive relationships to extrapolate the effects of global change on phenology. Various combinations of environmental factors occur concurrently at different European sites and the interactions between different drivers (e.g. water and chilling) can alter phenology significantly.     

The impact of global warming on germination and seedling emergence in Alliaria petiolata,a woodland species with dormancy loss dependent on low temperature

• The impact of global warming on seed dormancy loss and germination was investigated in Alliaria petiolata (garlic mustard), a common woodland/hedgerow plant in Eurasia, considered invasive in North America. Increased temperature may have serious implications, since seeds of this species germinate and emerge at low temperatures early in spring to establish and grow before canopy development of competing species.
• Dormancy was evaluated in seeds buried in field soils. Seedling emergence was also investigated in the field, and in a thermogradient tunnel under global warming scenarios representing predicted UK air temperatures through to 2080.
• Dormancy was simple, and its relief required the accumulation of low temperature chilling time. Under a global warming scenario, dormancy relief and seedling emergence declined and seed mortality increased as soil temperature increased along a thermal gradient. Seedling emergence advanced with soil temperature, peaking 8 days earlier under 2080 conditions.
• The results indicate that as mean temperature increases due to global warming, the chilling requirement for dormancy relief may not be fully satisfied, but seedling emergence will continue from low dormancy seeds in the population. Adaptation resulting from selection of this low dormancy proportion is likely to reduce the overall population chilling requirement. Seedling emergence is also likely to keep pace with the advancement of biological spring, enabling A. petiolata to maintain its strategy of establishment before the woodland canopy closes. However, this potential for adaptation may be countered by increased seed mortality in the seed bank as soils warm.

     

Understorey vegetation moderates climate in open forests: The role of the skirt-forming grass tree Xanthorrhoea semiplana F.Muell

Microsites are created by abiotic and biotic features of the landscape and may provide essential habitats for the persistence of biota. Forest canopies and understorey plants may moderate wind and solar radiation to create microclimatic conditions that differ considerably from regional climates. Skirt-forming plants, where senescent leaves create hut-like cavities around the stem, create microsites that are sheltered from ambient conditions and extreme weather events, constituting potential refuges for wildlife. We investigate day and night temperatures and humidity for four locations (grass tree cavities, soil, 20 cm above-ground, 1 m above-ground) in a South Australian forest with relatively open canopy of stringybark eucalypts (Eucalyptus baxteri, E. obliqua) and an understorey of skirt-forming grass trees (Xanthorrhoea semiplana) at 5, 10, 20, and 40 m from the forest edge. We also measured the percentage of canopy and understorey covers. Generally, temperature and humidity differed significantly between more sheltered (grass tree cavities, soil) and open-air microsites, with the former being cooler during the day and warmer and more humid during the night. Furthermore, our results suggest that canopy cover tends to decrease, and understorey cover tends to increase, the temperature of microsites. Distance to the edge was not significantly related to temperature for any microsite, suggesting that the edge effect did not extend beyond 10 m from the edge. Overall, grass trees influenced microclimatic conditions by forming a dense understorey and providing cavities that are relatively insulated. The capacity of grass tree cavities to buffer external conditions increased linearly with ambient temperatures, by 0.46°C per degree increase in maximum and 0.25°C per degree decrease in minimum temperatures, potentially offsetting climate warming and enabling persistence of fauna within their thermal limits. These climate moderation properties will make grass trees increasingly important refuges as extreme weather events become more common under anthropogenic climate change.     

Investigating the abundance and flight period of bark beetles (Coleoptera: Curculionidae: Scolytinae) over elevational gradients in Sitka spruce forests

1. A warming climate, as predicted under current climate change projections, is likely to influence the population dynamics of many forest insect species. Numerous bark beetle species in both Europe and North America have already responded to a warming climate by significantly expanding their geographical ranges.
2. The aim of the current study was to investigate how populations of bark beetles within stands of Sitka spruce, a widely planted non-native commercial plantation tree species in the U.K., were likely to respond to a warming climate. Experimental plots were established in stands of Sitka spruce over elevational gradients in two commercial forest plantations, and the abundance and emergence times of key bark beetle species were assessed over a 3-year period using flight interception traps. The air temperature difference between the lowest and highest experimental plot in each forest was consistently >1°C throughout the 3-year period.
3. In general, the abundance of the most dominant bark beetle species (e.g. Trypodendron, Dryocoetes, Hylastes spp.) was higher, and emergence times tended to be earlier in the year at the lower elevation plots, where temperatures were higher, although not all bark beetle species responded in the same manner.
4. The results of the study indicated that, under the projected future climate warming scenarios, monoculture Sitka spruce stands at low elevations may potentially be more vulnerable to significant outbreak events from existing or invasive bark beetle species. Hence, consideration of establishing more resilient forests of Sitka spruce by diversifying the species composition and structure of Sitka spruce stands is discussed.

     

Tropical understory herbaceous community responds more strongly to hurricane disturbance than to experimental warming

The effects of climate change on tropical forests may have global consequences due to the forests’ high biodiversity and major role in the global carbon cycle. In this study, we document the effects of experimental warming on the abundance and composition of a tropical forest floor herbaceous plant community in the Luquillo Experimental Forest, Puerto Rico. This study was conducted within Tropical Responses to Altered Climate Experiment (TRACE) plots, which use infrared heaters under free‐air, open‐field conditions, to warm understory vegetation and soils + 4°C above nearby control plots. Hurricanes Irma and María damaged the heating infrastructure in the second year of warming, therefore, the study included one pretreatment year, one year of warming, and one year of hurricane response with no warming. We measured percent leaf cover of individual herbaceous species, fern population dynamics, and species richness and diversity within three warmed and three control plots. Results showed that one year of experimental warming did not significantly affect the cover of individual herbaceous species, fern population dynamics, species richness, or species diversity. In contrast, herbaceous cover increased from 20% to 70%, bare ground decreased from 70% to 6%, and species composition shifted pre to posthurricane. The negligible effects of warming may have been due to the short duration of the warming treatment or an understory that is somewhat resistant to higher temperatures. Our results suggest that climate extremes that are predicted to increase with climate change, such as hurricanes and droughts, may cause more abrupt changes in tropical forest understories than longer‐term sustained warming.     

Different initial responses of the canopy herbivory rate in mature oak trees to experimental soil and branch warming in a soil‐freezing area

Plants and insects comprise more than 50% of known species on earth, and their interactions are of major importance in most natural ecosystems. To understand the mechanisms by which global warming affects plant–insect interactions in the canopy of mature cool‐temperate forests with a freeze–thaw cycle, we examined changes in the herbivory rate and leaf traits in oak Quercus crispula. From 2007 to 2009, we experimentally increased the temperature of the surrounding soil and canopy branches of mature oak trees by approximately 5°C using electric heating cables. Soil warming decreased the rate of herbivory in the canopy, whereas branch warming had no effect. The magnitude of the effect of soil warming on canopy herbivory varied. For the first year, the decrease was 32%, but this doubled (63%) in the third year. Branch warming did not affect canopy leaf traits; however, soil warming decreased the leaf nutritional quality by decreasing N and increasing the carbon:nitrogen (CN) ratio for three years. Additionally, soil warming increased total phenolics in the third year. Stepwise multiple regression models showed that among the leaf traits that were changed by soil warming, N explained the variation in herbivory for the first and second years, whereas total phenolics explained it for the third year. Our experimental results demonstrate that soil warming drives the rate of herbivory in the canopy of mature oak trees, and the magnitude of the soil warming effect was gradually enhanced during the initial three years. This suggests the importance of belowground temperature elevation in predicting the effect of global warming on plant–insect interactions in a forest canopy.     

Design and performance of combined infrared canopy and belowground warming in the B4WarmED (Boreal Forest Warming at an Ecotone in Danger) experiment

Conducting manipulative climate change experiments in complex vegetation is challenging, given considerable temporal and spatial heterogeneity. One specific challenge involves warming of both plants and soils to depth. We describe the design and performance of an open‐air warming experiment called Boreal Forest Warming at an Ecotone in Danger (B4WarmED) that addresses the potential for projected climate warming to alter tree function, species composition, and ecosystem processes at the boreal‐temperate ecotone. The experiment includes two forested sites in northern Minnesota, USA, with plots in both open (recently clear‐cut) and closed canopy habitats, where seedlings of 11 tree species were planted into native ground vegetation. Treatments include three target levels of plant canopy and soil warming (ambient, +1.7 °C, +3.4 °C). Warming was achieved by independent feedback control of voltage input to aboveground infrared heaters and belowground buried resistance heating cables in each of 72‐7.0 m² plots. The treatments emulated patterns of observed diurnal, seasonal, and annual temperatures but with superimposed warming. For the 2009 to 2011 field seasons, we achieved temperature elevations near our targets with growing season overall mean differences (?T_below) of +1.84 °C and +3.66 °C at 10 cm soil depth and (?T^above) of +1.82 °C and +3.45 °C for the plant canopies. We also achieved measured soil warming to at least 1 m depth. Aboveground treatment stability and control were better during nighttime than daytime and in closed vs. open canopy sites in part due to calmer conditions. Heating efficacy in open canopy areas was reduced with increasing canopy complexity and size. Results of this study suggest the warming approach is scalable: it should work well in small‐statured vegetation such as grasslands, desert, agricultural crops, and tree saplings (<5 m tall).     

Thermal plasticity of photosynthesis: the role of acclimation in forest responses to a warming climate

The increasing air temperatures central to climate change predictions have the potential to alter forest ecosystem function and structure by exceeding temperatures optimal for carbon gain. Such changes are projected to threaten survival of sensitive species, leading to local extinctions, range migrations, and altered forest composition. This study investigated photosynthetic sensitivity to temperature and the potential for acclimation in relation to the climatic provenance of five species of deciduous trees, Liquidambar styraciflua, Quercus rubra, Quercus falcata, Betula alleghaniensis, and Populus grandidentata. Open‐top chambers supplied three levels of warming (+0, +2, and +4 °C above ambient) over 3 years, tracking natural temperature variability. Optimal temperature for CO₂ assimilation was strongly correlated with daytime temperature in all treatments, but assimilation rates at those optima were comparable. Adjustment of thermal optima was confirmed in all species, whether temperatures varied with season or treatment, and regardless of climate in the species' range or provenance of the plant material. Temperature optima from 17° to 34° were observed. Across species, acclimation potentials varied from 0.55 °C to 1.07 °C per degree change in daytime temperature. Responses to the temperature manipulation were not different from the seasonal acclimation observed in mature indigenous trees, suggesting that photosynthetic responses should not be modeled using static temperature functions, but should incorporate an adjustment to account for acclimation. The high degree of homeostasis observed indicates that direct impacts of climatic warming on forest productivity, species survival, and range limits may be less than predicted by existing models.     

MICROCLIMATIC EFFECTS ON WATER RELATIONS,LEAF TEMPERATURES,AND THE DISTRIBUTION OF HERACLEUM LANATUM AT HIGH ELEVATIONS

The small-scale distribution of an understory herb, Heracleum lanatum, was evaluated in terms of leaf temperature and water relations limitations due to a large leaf size (630 cm²). Diurnal variations in transpiration (4 to 60 mg m⁻² s⁻¹) were influenced by fluctuations in solar irradiance, wind speed, leaf temperature and stomatal conductance. Computer simulations indicated that leaf temperatures in a forest clearing would be > 12 C above air temperature, with maximum transpiration rates of 140 mg m⁻² s⁻¹, and daily water loss to be over 200% greater than values at natural understory locations. Simulations of nocturnal temperature relations indicated ~100 W m ⁻² less incident longwave irradiance in the forest clearing as compared to the understory (560 vs. 660 W m⁻² at 400 hr). This difference led to predicted leaf temperatures being as low as 6 C below air temperature in the forest clearing while measured leaf temperatures in the forest understory were within 1.5 C of air temperature throughout the night. Furthermore, minimum air temperatures were at or below 6 C on 36% of the nights during the summer growth period indicating that in open areas leaves of H. lanatum would frequently be below 0 C and subject to possible freeze damage. Heracleum lanatum may be more abundant in the shaded understory of the subalpine forest because exposure in open environments would result in high leaf temperatures and increased transpirational water loss during the day, as well as low leaf temperatures with the possibility of freeze damage at night.     

Infrared heater system for warming tropical forest understory plants and soils

The response of tropical forests to global warming is one of the largest uncertainties in predicting the future carbon balance of Earth. To determine the likely effects of elevated temperatures on tropical forest understory plants and soils, as well as other ecosystems, an infrared (IR) heater system was developed to provide in situ warming for the Tropical Responses to Altered Climate Experiment (TRACE) in the Luquillo Experimental Forest in Puerto Rico. Three replicate heated 4‐m‐diameter plots were warmed to maintain a 4°C increase in understory vegetation compared to three unheated control plots, as sensed by IR thermometers. The equipment was larger than any used previously and was subjected to challenges different from those of many temperate ecosystem warming systems, including frequent power surges and outages, high humidity, heavy rains, hurricanes, saturated clayey soils, and steep slopes. The system was able to maintain the target 4.0°C increase in hourly average vegetation temperatures to within ± 0.1°C. The vegetation was heterogeneous and on a 21° slope, which decreased uniformity of the warming treatment on the plots; yet, the green leaves were fairly uniformly warmed, and there was little difference among 0–10 cm depth soil temperatures at the plot centers, edges, and midway between. Soil temperatures at the 40–50 cm depth increased about 3°C compared to the controls after a month of warming. As expected, the soil in the heated plots dried faster than that of the control plots, but the average soil moisture remained adequate for the plants. The TRACE heating system produced an adequately uniform warming precisely controlled down to at least 50‐cm soil depth, thereby creating a treatment that allows for assessing mechanistic responses of tropical plants and soil to warming, with applicability to other ecosystems. No physical obstacles to scaling the approach to taller vegetation (i.e., trees) and larger plots were observed.     

Predicting xylem phenology in black spruce under climate warming

In the next century, the boreal ecosystems are projected to experience greater rates of warming than most other regions of the world. As the boreal forest constitutes a reservoir of trees of huge ecological importance and only partially known economic potential, any possible climate‐related change in plant growth and dynamics has to be promptly predicted and evaluated. A model for assessing xylem phenology in black spruce [Picea mariana (Mill.) B.S.P.] using daily temperatures and thermal thresholds was defined and applied to predict changes in onset, ending and duration of xylem growth under different warming scenarios with temperatures rising by up to 3 °C. This was achieved by collecting and analyzing a dataset obtained from a 7‐year monitoring of cambium phenology and wood formation on a weekly time‐scale in trees growing in four sites at different latitudes and altitudes in the Saguenay‐Lac‐Saint‐Jean region (Quebec, Canada). The onset of xylem growth occurred between mid‐May and early June while the end ranged between mid‐September and early October, resulting in a growing season of 101–141 days. The model predicted longer duration of xylem growth at higher temperatures, with an increase of 8–11 days/ °C, because of an earlier onset and later ending of growth. With an increase of 3 °C in the mean temperature during the year, the duration of xylem growth changed on average from 125 to 160 days. The predicted changes in cambial phenology could significantly affect future wood production of the boreal ecosystems.     

Trees tolerate an extreme heatwave via sustained transpirational cooling and increased leaf thermal tolerance

Heatwaves are likely to increase in frequency and intensity with climate change, which may impair tree function and forest C uptake. However, we have little information regarding the impact of extreme heatwaves on the physiological performance of large trees in the field. Here, we grew Eucalyptus parramattensis trees for 1 year with experimental warming (+3°C) in a field setting, until they were greater than 6 m tall. We withheld irrigation for 1 month to dry the surface soils and then implemented an extreme heatwave treatment of 4 consecutive days with air temperatures exceeding 43°C, while monitoring whole‐canopy exchange of CO₂ and H₂O, leaf temperatures, leaf thermal tolerance, and leaf and branch hydraulic status. The heatwave reduced midday canopy photosynthesis to near zero but transpiration persisted, maintaining canopy cooling. A standard photosynthetic model was unable to capture the observed decoupling between photosynthesis and transpiration at high temperatures, suggesting that climate models may underestimate a moderating feedback of vegetation on heatwave intensity. The heatwave also triggered a rapid increase in leaf thermal tolerance, such that leaf temperatures observed during the heatwave were maintained within the thermal limits of leaf function. All responses were equivalent for trees with a prior history of ambient and warmed (+3°C) temperatures, indicating that climate warming conferred no added tolerance of heatwaves expected in the future. This coordinated physiological response utilizing latent cooling and adjustment of thermal thresholds has implications for tree tolerance of future climate extremes as well as model predictions of future heatwave intensity at landscape and global scales.