Taller and larger: shifts in Arctic tundra leaf traits after 16 years of experimental warming

Understanding plant trait responses to elevated temperatures in the Arctic is critical in light of recent and continuing climate change, especially because these traits act as key mechanisms in climate‐vegetation feedbacks. Since 1992, we have artificially warmed three plant communities at Alexandra Fiord, Nunavut, Canada (79°N). In each of the communities, we used open‐top chambers (OTCs) to passively warm vegetation by 1–2 °C. In the summer of 2008, we investigated the intraspecific trait responses of five key species to 16 years of continuous warming. We examined eight traits that quantify different aspects of plant performance: leaf size, specific leaf area (SLA), leaf dry matter content (LDMC), plant height, leaf carbon concentration, leaf nitrogen concentration, leaf carbon isotope discrimination (LCID), and leaf δ¹⁵N. Long‐term artificial warming affected five traits, including at least one trait in every species studied. The evergreen shrub Cassiope tetragona responded most frequently (increased leaf size and plant height/decreased SLA, leaf carbon concentration, and LCID), followed by the deciduous shrub Salix arctica (increased leaf size and plant height/decreased SLA) and the evergreen shrub Dryas integrifolia (increased leaf size and plant height/decreased LCID), the forb Oxyria digyna (increased leaf size and plant height), and the sedge Eriophorum angustifolium spp. triste (decreased leaf carbon concentration). Warming did not affect δ¹⁵N, leaf nitrogen concentration, or LDMC. Overall, growth traits were more sensitive to warming than leaf chemistry traits. Notably, we found that responses to warming were sustained, even after many years of treatment. Our work suggests that tundra plants in the High Arctic will show a multifaceted response to warming, often including taller shoots with larger leaves.     

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.     

Thermal acclimation of leaf respiration of tropical trees and lianas: response to experimental canopy warming,and consequences for tropical forest carbon balance

Climate warming is expected to increase respiration rates of tropical forest trees and lianas, which may negatively affect the carbon balance of tropical forests. Thermal acclimation could mitigate the expected respiration increase, but the thermal acclimation potential of tropical forests remains largely unknown. In a tropical forest in Panama, we experimentally increased nighttime temperatures of upper canopy leaves of three tree and two liana species by on average 3  ° C for 1 week, and quantified temperature responses of leaf dark respiration. Respiration at 25  ° C (R₂₅) decreased with increasing leaf temperature, but acclimation did not result in perfect homeostasis of respiration across temperatures. In contrast, Q₁₀ of treatment and control leaves exhibited similarly high values (range 2.5–3.0) without evidence of acclimation. The decrease in R₂₅ was not caused by respiratory substrate depletion, as warming did not reduce leaf carbohydrate concentration. To evaluate the wider implications of our experimental results, we simulated the carbon cycle of tropical latitudes (24 ° S–24 ° N) from 2000 to 2100 using a dynamic global vegetation model (LM3VN) modified to account for acclimation. Acclimation reduced the degree to which respiration increases with climate warming in the model relative to a no‐acclimation scenario, leading to 21% greater increase in net primary productivity and 18% greater increase in biomass carbon storage over the 21st century. We conclude that leaf respiration of tropical forest plants can acclimate to nighttime warming, thereby reducing the magnitude of the positive feedback between climate change and the carbon cycle.     

Contrasting responses of steppe Stipa ssp. to warming and precipitation variability

Climate change, characterized by warming and precipitation variability, restricted the growth of plants in arid and semiarid areas, and various functional traits are impacted differently. Comparing responses of functional traits to warming and precipitation variability and determining critical water threshold of dominate steppe grasses from Inner Mongolia facilitates the identification and monitoring of water stress effects. A combination of warming (ambient temperature, +1.5°C and +2.0°C) and varying precipitation (?30%, ?15%, ambient, +15%, and +30%) manipulation experiments were performed on four Stipa species (S. baicalensis, S. bungeana, S. grandis, and S. breviflora) from Inner Mongolia steppe. The results showed that the functional traits of the four grasses differed in their responses to precipitation, but they shared common sensitive traits (root/shoot ratio, R/S, and specific leaf area; SLA) under ambient temperature condition. Warming increased the response of the four grasses to changing precipitation, and these differences in functional traits resulted in changes to their total biomass, with leaf area, SLA, and R/S making the largest contributions. Critical water thresholds of the four grasses were identified, and warming led to their higher optimum precipitation requirements. The four steppe grasses were able to adapt better to mild drought (summer precipitation decreased by 12%–28%) when warming 1.5°C rather than 2.0°C. These results indicated that if the Paris Agreement to limit global warming to 1.5°C will be accomplished, this will increase the probability for sustained viability of the Stipa steppes in the next 50–100 years.     

Thermal limits of leaf metabolism across biomes

High‐temperature tolerance in plants is important in a warming world, with extreme heat waves predicted to increase in frequency and duration, potentially leading to lethal heating of leaves. Global patterns of high‐temperature tolerance are documented in animals, but generally not in plants, limiting our ability to assess risks associated with climate warming. To assess whether there are global patterns in high‐temperature tolerance of leaf metabolism, we quantified T_crit (high temperature where minimal chlorophyll a fluorescence rises rapidly and thus photosystem II is disrupted) and T_max (temperature where leaf respiration in darkness is maximal, beyond which respiratory function rapidly declines) in upper canopy leaves of 218 plant species spanning seven biomes. Mean site‐based T_crit values ranged from 41.5 °C in the Alaskan arctic to 50.8 °C in lowland tropical rainforests of Peruvian Amazon. For T_max, the equivalent values were 51.0 and 60.6 °C in the Arctic and Amazon, respectively. T_crit and T_max followed similar biogeographic patterns, increasing linearly (?8 °C) from polar to equatorial regions. Such increases in high‐temperature tolerance are much less than expected based on the 20 °C span in high‐temperature extremes across the globe. Moreover, with only modest high‐temperature tolerance despite high summer temperature extremes, species in mid‐latitude (~20–50°) regions have the narrowest thermal safety margins in upper canopy leaves; these regions are at the greatest risk of damage due to extreme heat‐wave events, especially under conditions when leaf temperatures are further elevated by a lack of transpirational cooling. Using predicted heat‐wave events for 2050 and accounting for possible thermal acclimation of T_crit and T_max, we also found that these safety margins could shrink in a warmer world, as rising temperatures are likely to exceed thermal tolerance limits. Thus, increasing numbers of species in many biomes may be at risk as heat‐wave events become more severe with climate change.     

Responses of canopy duration to temperature changes in four temperate tree species: relative contributions of spring and autumn leaf phenology

While changes in spring phenological events due to global warming have been widely documented, changes in autumn phenology, and therefore in growing season length, are less studied and poorly understood. However, it may be helpful to assess the potential lengthening of the growing season under climate warming in order to determine its further impact on forest productivity and C balance. The present study aimed to: (1) characterise the sensitivity of leaf phenological events to temperature, and (2) quantify the relative contributions of leaf unfolding and senescence to the extension of canopy duration with increasing temperature, in four deciduous tree species (Acer pseudoplatanus, Fagus sylvatica, Fraxinus excelsior and Quercus petraea). For 3 consecutive years, we monitored the spring and autumn phenology of 41 populations at elevations ranging from 100 to 1,600 m. Overall, we found significant altitudinal trends in leaf phenology and species-specific differences in temperature sensitivity. With increasing temperature, we recorded an advance in flushing from 1.9 ± 0.3 to 6.6 ± 0.4 days °C⁻¹ (mean ± SD) and a 0 to 5.6 ± 0.6 days °C⁻¹ delay in leaf senescence. Together both changes resulted in a 6.9 ± 1.0 to 13.0 ± 0.7 days °C⁻¹ lengthening of canopy duration depending on species. For three of the four studied species, advances in flushing were the main factor responsible for lengthening canopy duration with increasing temperature, leading to a potentially larger gain in solar radiation than delays in leaf senescence. In contrast, for beech, we found a higher sensitivity to temperature in leaf senescence than in flushing, resulting in an equivalent contribution in solar radiation gain. These results suggest that climate warming will alter the C uptake period and forest productivity by lengthening canopy duration. Moreover, the between-species differences in phenological responses to temperature evidenced here could affect biotic interactions under climate warming.     

Experimental climate warming alters aspen and birch phytochemistry and performance traits for an outbreak insect herbivore

Climate change and insect outbreaks are key factors contributing to regional and global patterns of increased tree mortality. While links between these environmental stressors have been established, our understanding of the mechanisms by which elevated temperature may affect tree–insect interactions is limited. Using a forest warming mesocosm, we investigated the influence of elevated temperature on phytochemistry, tree resistance traits, and insect performance. Specifically, we examined warming effects on forest tent caterpillar (Malacosoma disstria) and host trees aspen (Populus tremuloides) and birch (Betula papyrifera). Trees were grown under one of three temperature treatments (ambient, +1.7 °C, +3.4 °C) in a multiyear open‐air warming experiment. In the third and fourth years of warming (2011, 2012), we assessed foliar nutrients and defense chemistry. Elevated temperatures altered foliar nitrogen, carbohydrates, lignin, and condensed tannins, with differences in responses between species and years. In 2012, we performed bioassays using a common environment approach to evaluate plant‐mediated indirect warming effects on larval performance. Warming resulted in decreased food conversion efficiency and increased consumption, ultimately with minimal effect on larval development and biomass. These changes suggest that insects exhibited compensatory feeding due to reduced host quality. Within the context of observed phytochemical variation, primary metabolites were stronger predictors of insect performance than secondary metabolites. Between‐year differences in phytochemical shifts corresponded with substantially different weather conditions during these two years. By sampling across years within an ecologically realistic and environmentally open setting, our study demonstrates that plant and insect responses to warming can be temporally variable and context dependent. Results indicate that elevated temperatures can alter phytochemistry, tree resistance traits, and herbivore feeding, but that annual weather variability may modulate warming effects leading to uncertain consequences for plant–insect interactions with projected climate change.     

Effects of thermal history on the responses to thermal stress of a large benthic foraminifera,Calcarina gaudichaudii

Marine ecosystems, particularly coastal environments, are rapidly changing due to anthropogenic impacts resulting in increased global climate change (ocean warming), ocean acidification, hypoxia, and eutrophication. On coral reefs, symbiont-bearing large benthic foraminifera (LBFs) can play a key role as reef constituents and carbonate producers, contributing up to 5% of reef-scale carbonate budgets. However, projected climate change, particularly ocean warming, has the potential to significantly alter the conditions in which marine organisms persist. While the response of LBFs to elevated thermal stress is well documented in laboratory studies, the potential influence of adaptation or acclimatization through prior environmental thermal history on this response remains largely unknown. In this study, specimens of Calcarina gaudichaudii, an LBF from the Penghu Islands, Taiwan, were collected from thermally variable intertidal and thermally stable subtidal (~ 6 m depth) environments representing thermal history. LBFs were then acclimated to laboratory conditions at ambient (25 °C) and elevated (28 °C) temperatures for three weeks, and subsequently exposed to control and heat stress treatments (25 °C, 28 °C, 30 °C, 33 °C) for an additional one week. Photosynthetic rates (determined through oxygen flux measurements) of C. gaudichaudii significantly decreased in specimens collected at subtidal depths acclimated at 25 °C when compared to those acclimated at 28 °C, whereas there was no effect of thermal history on respiration, indicating that symbiont and holobiont responses may differ in LBFs. Additionally, maximum photochemical efficiency (Fv/Fm) significantly decreased as a result of heat stress, although bleaching was not visually observed after one week. These results highlight the plastic responses of the algal microbiome and indicate that thermal history, acclimatization temperature, and heat stress interact to affect the physiological status of C. gaudichaudii. This study adds to the growing literature which highlights the larger implications of understanding thermal history as an important factor to consider to better understand how ecosystem processes (e.g., carbonate production) are altered on modern coral reefs.

     

川西林线交错带岷江冷杉幼苗异龄叶形态对长期模拟增温的响应

全球气候变暖对高纬度、高海拔地区的植物形态产生强烈影响。川西林线交错带是青藏高原东部高寒生态系统的重要组成部分,对全球变化极度敏感。以川西林线交错带岷江冷杉(Abies faxoniana)幼苗异龄叶为对象,采用原位开顶式生长室(Open-top chamber,OTCs)模拟增温,研究了长期模拟增温下岷江冷杉幼苗异龄叶片叶长、叶厚等叶形态的响应,采用表型可塑性指数和变异系数对叶形态的可塑性进行分析。结果表明:增温限制岷江冷杉幼苗叶片的增长、增宽和叶面积、体积的扩大,使叶长、叶宽、叶面积、叶体积分别较对照减小12.77%、11.86%、17.76%、11.49%;增温促进叶片厚度的增加,较对照增加7.27%;除叶长宽比外,增温对其余叶形态均产生显著影响(P0.05)。叶形态对模拟增温的响应具有显著的年龄差异(P0.05)。温度、叶龄的交互效应对叶长和叶面积影响显著(P0.05),对叶宽和叶厚影响不显著(P0.05)。两种表型可塑性分析结果表明,除1 a叶叶长外,增温不同程度增大各叶形态可塑性。长期增温使冷杉幼苗叶片有旱生倾向且形态值更发散。研究提供了岷江冷杉幼苗叶片对长期增温的差异性响应证据,为评估青藏高原东缘优势植物对响应气候变暖提供了基础数据和理论参考。     

Contrasting acclimation abilities of two dominant boreal conifers to elevated CO2 and temperature

High latitude forests will experience large changes in temperature and CO₂ concentrations this century. We evaluated the effects of future climate conditions on 2 dominant boreal tree species, Pinus sylvestris L. and Picea abies (L.) H. Karst, exposing seedlings to 3 seasons of ambient (430 ppm) or elevated CO₂ (750 ppm) and ambient temperatures, a + 4 °C warming or a + 8 °C warming. Pinus sylvestris responded positively to warming: seedlings developed a larger canopy, maintained high net CO₂ assimilation rates (A_net), and acclimated dark respiration (R_dark). In contrast, carbon fluxes in Picea abies were negatively impacted by warming: maximum rates of A_net decreased, electron transport was redirected to alternative electron acceptors, and thermal acclimation of R_dark was weak. Elevated CO₂ tended to exacerbate these effects in warm‐grown Picea abies, and by the end of the experiment Picea abies from the +8 °C, high CO₂ treatment produced fewer buds than they had 3 years earlier. Treatments had little effect on leaf and wood anatomy. Our results highlight that species within the same plant functional type may show opposite responses to warming and imply that Picea abies may be particularly vulnerable to warming due to low plasticity in photosynthetic and respiratory metabolism.     

The relative impacts of daytime and night-time warming on photosynthetic capacity in Populus deltoides

In order to investigate the relative impacts of increases in day and night temperature on tree carbon relations, we measured night‐time respiration and daytime photosynthesis of leaves in canopies of 4‐m‐tall cottonwood (Populus deltoides Bartr. ex Marsh) trees experiencing three daytime temperatures (25, 28 or 31 °C) and either (i) a constant nocturnal temperature of 20 °C or (ii) increasing nocturnal temperatures (15, 20 or 25 °C). In the first (day warming only) experiment, rates of night‐time leaf dark respiration (R_dark) remained constant and leaves displayed a modest increase (11%) in light‐saturated photosynthetic capacity (A_max) during the day (1000–1300 h) over the 6 °C range. In the second (dual night and day warming) experiment, R_dark increased by 77% when nocturnal temperatures were increased from 15 °C (0·36 µmol m^?2 s^?1) to 25 °C (0·64 µmol m^?2 s^?1). A_max responded positively to the additional nocturnal warming, and increased by 38 and 64% in the 20/28 and 25/31 °C treatments, respectively, compared with the 15/25 °C treatment. These increases in photosynthetic capacity were associated with strong increases in the maximum carboxylation rate of rubisco (V_cmax) and ribulose‐1,5‐bisphosphate (RuBP) regeneration capacity mediated by maximum electron transport rate (J_max). Leaf soluble sugar and starch concentration, measured at sunrise, declined significantly as nocturnal temperature increased. The nocturnal temperature manipulation resulted in a significant inverse relationship between A_max and pre‐dawn leaf carbohydrate status. Independent measurements of the temperature response of photosynthesis indicated that the optimum temperature (T_opt) acclimated fully to the 6 °C range of temperature imposed in the daytime warming. Our findings are consistent with the hypothesis that elevated night‐time temperature increases photosynthetic capacity during the following light period through a respiratory‐driven reduction in leaf carbohydrate concentration. These responses indicate that predicted increases in night‐time minimum temperatures may have a significant influence on net plant carbon uptake.     

Rapid loss of an ecosystem engineer: Sphagnum decline in an experimentally warmed bog

Sphagnum mosses are keystone components of peatland ecosystems. They facilitate the accumulation of carbon in peat deposits, but climate change is predicted to expose peatland ecosystem to sustained and unprecedented warming leading to a significant release of carbon to the atmosphere. Sphagnum responses to climate change, and their interaction with other components of the ecosystem, will determine the future trajectory of carbon fluxes in peatlands. We measured the growth and productivity of Sphagnum in an ombrotrophic bog in northern Minnesota, where ten 12.8‐m‐diameter plots were exposed to a range of whole‐ecosystem (air and soil) warming treatments (+0 to +9°C) in ambient or elevated (+500 ppm) CO₂. The experiment is unique in its spatial and temporal scale, a focus on response surface analysis encompassing the range of elevated temperature predicted to occur this century, and consideration of an effect of co‐occurring CO₂ altering the temperature response surface. In the second year of warming, dry matter increment of Sphagnum increased with modest warming to a maximum at 5°C above ambient and decreased with additional warming. Sphagnum cover declined from close to 100% of the ground area to <50% in the warmest enclosures. After three years of warming, annual Sphagnum productivity declined linearly with increasing temperature (13–29 g C/m² per °C warming) due to widespread desiccation and loss of Sphagnum. Productivity was less in elevated CO₂ enclosures, which we attribute to increased shading by shrubs. Sphagnum desiccation and growth responses were associated with the effects of warming on hydrology. The rapid decline of the Sphagnum community with sustained warming, which appears to be irreversible, can be expected to have many follow‐on consequences to the structure and function of this and similar ecosystems, with significant feedbacks to the global carbon cycle and climate change.     

Warming water and leaf litter quality but not plant origin drive decomposition and fungal diversity in an experiment

The fungi associated with leaf litter play a key role in decomposition and can be affected both by the warming water and the invasion of non-native species in riparian vegetation. Warming water and invasion of non-native riparian species on stream fungal communities have been studied mainly in temperate ecosystems. We tested the effects of warming water and non-native plant Psidium guajava on leaf litter decomposition, conidia density, species richness and beta diversity of tropical stream fungi. Thus, we carried out an experiment using the current mean temperature of streams from northwestern Paraná in South Brazil (22 °C) and two temperatures above the current mean temperature (26 °C and 29 °C). We also used the leaves of a non-native plant (P. guajava), and two native plants (one of similar nutritional quality, and the other of higher nutritional quality than the non-native species) occurring in Neotropical streams riparian vegetation. Warming water accelerated leaf litter decomposition and reduced conidia density and fungal richness in native and non-native plants. However, species composition and beta diversity were not affected by water temperature. Our study showed that warming affects the fungi of streams, the main microorganisms responsible for decomposition and that the nutritional quality of the leaves may be more important than the origin of riparian plant species. Despite this, further investigations should be conducted on the interaction of P. guajava with the flow of nutrients in these environments and how it can affect other ecosystem processes and the food chain. Efforts to study the effects of water warming and biological invasion on the attributes and distribution of fungi in streams are vital, making them a tool for the conservation of riparian ecosystems.     

Contrasting acclimation responses to elevated CO2 and warming between an evergreen and a deciduous boreal conifer

Rising atmospheric carbon dioxide (CO₂) concentrations may warm northern latitudes up to 8°C by the end of the century. Boreal forests play a large role in the global carbon cycle, and the responses of northern trees to climate change will thus impact the trajectory of future CO₂ increases. We grew two North American boreal tree species at a range of future climate conditions to assess how growth and carbon fluxes were altered by high CO₂ and warming. Black spruce (Picea mariana, an evergreen conifer) and tamarack (Larix laricina, a deciduous conifer) were grown under ambient (407 ppm) or elevated CO₂ (750 ppm) and either ambient temperatures, a 4°C warming, or an 8°C warming. In both species, the thermal optimum of net photosynthesis (T_optA) increased and maximum photosynthetic rates declined in warm‐grown seedlings, but the strength of these changes varied between species. Photosynthetic capacity (maximum rates of Rubisco carboxylation, V_cmax, and of electron transport, J_max) was reduced in warm‐grown seedlings, correlating with reductions in leaf N and chlorophyll concentrations. Warming increased the activation energy for V_cmax and J_max (E_aV and E_aJ, respectively) and the thermal optimum for J_max. In both species, the T_optA was positively correlated with both E_aV and E_aJ, but negatively correlated with the ratio of J_max/V_cmax. Respiration acclimated to elevated temperatures, but there were no treatment effects on the Q₁₀ of respiration (the increase in respiration for a 10°C increase in leaf temperature). A warming of 4°C increased biomass in tamarack, while warming reduced biomass in spruce. We show that climate change is likely to negatively affect photosynthesis and growth in black spruce more than in tamarack, and that parameters used to model photosynthesis in dynamic global vegetation models (E_aV and E_aJ) show no response to elevated CO₂.     

Global synthesis of the temperature sensitivity of leaf litter breakdown in streams and rivers

Streams and rivers are important conduits of terrestrially derived carbon (C) to atmospheric and marine reservoirs. Leaf litter breakdown rates are expected to increase as water temperatures rise in response to climate change. The magnitude of increase in breakdown rates is uncertain, given differences in litter quality and microbial and detritivore community responses to temperature, factors that can influence the apparent temperature sensitivity of breakdown and the relative proportion of C lost to the atmosphere vs. stored or transported downstream. Here, we synthesized 1025 records of litter breakdown in streams and rivers to quantify its temperature sensitivity, as measured by the activation energy (E_a, in eV). Temperature sensitivity of litter breakdown varied among twelve plant genera for which E_a could be calculated. Higher values of E_a were correlated with lower‐quality litter, but these correlations were influenced by a single, N‐fixing genus (Alnus). E_a values converged when genera were classified into three breakdown rate categories, potentially due to continual water availability in streams and rivers modulating the influence of leaf chemistry on breakdown. Across all data representing 85 plant genera, the E_a was 0.34 ± 0.04 eV, or approximately half the value (0.65 eV) predicted by metabolic theory. Our results indicate that average breakdown rates may increase by 5–21% with a 1–4 °C rise in water temperature, rather than a 10–45% increase expected, according to metabolic theory. Differential warming of tropical and temperate biomes could result in a similar proportional increase in breakdown rates, despite variation in E_a values for these regions (0.75 ± 0.13 eV and 0.27 ± 0.05 eV, respectively). The relative proportions of gaseous C loss and organic matter transport downstream should not change with rising temperature given that E_a values for breakdown mediated by microbes alone and microbes plus detritivores were similar at the global scale.     

Combined effects of elevated temperatures and reduced leaf litter quality on the life-history parameters of a saprophagous macroarthropod

Because soil macroinvertebrates strongly modify decomposition processes, it is important to know how their abundance will respond to global change. We investigated in laboratory microcosms, the effects of elevated temperatures and reduced leaf litter quality on the life‐history traits of a saprophagous macroarthropod (development time, growth, survival and reproduction). Millipedes (Polydesmus angustus) from an Atlantic temperate forest were reared throughout their life cycle (≥16 months) under two temperature regimes differing on average by 3.3 °C; in a factorial design, they were fed either on Atlantic leaf litter or on Mediterranean leaf litter with a higher C : N ratio; humidity was consistently high. The components of the population growth rate (r) were affected positively by the temperature rise and negatively by the switch from Atlantic to Mediterranean leaf litter. When both treatments were combined, litter effects offset temperature effects. These results show that the short‐term response of saprophagous macroarthropods to warming is positive but depends on the availability of high‐quality litter, which is difficult to predict in the global change context. In a parallel experiment, conspecific millipedes from a Mediterranean population, which have evolved for a long time in a warmer climate and on poor‐quality litter, were reared at elevated temperatures on Mediterranean leaf litter. All components of r were higher than in the Atlantic population under the same conditions. This suggests that in the longer term, macroarthropods can overcome detrimental trophic interactions. Based on our study and the literature, we conclude that for decades the positive effects of warming on saprophagous macrofauna should exceed the negative effects of changes in litter quality. The abundance of those organisms in temperate forests could increase, which is confirmed by latitudinal patterns in Europe. Studies aimed at predicting the impacts of global change on decomposition will need to consider interactions with soil macroinvertebrates.     

Short photoperiod reduces the temperature sensitivity of leaf‐out in saplings of Fagus sylvatica but not in horse chestnut

Leaf phenology is one of the most reliable bioindicators of ongoing global warming in temperate and boreal zones because it is highly sensitive to temperature variation. A large number of studies have reported advanced spring leaf‐out due to global warming, yet the temperature sensitivity of leaf‐out has significantly decreased in temperate deciduous tree species over the past three decades. One of the possible mechanisms is that photoperiod is limiting further advance to protect the leaves against potential damaging frosts. However, the “photoperiod limitation” hypothesis remains poorly investigated and experimentally tested. Here, we conducted a photoperiod‐ and temperature‐manipulation experiment in climate chambers on two common deciduous species in Europe: Fagus sylvatica (European beech, a typically late flushing species) and Aesculus hippocastanum (horse chestnut, a typically early flushing species). In agreement with previous studies, we found that the warming significantly advanced the leaf‐out dates by 4.3 and 3.7 days/°C for beech and horse chestnut saplings, respectively. However, shorter photoperiod significantly reduced the temperature sensitivity of beech only (3.0 days/°C) by substantially increasing the heat requirement to avoid leafing‐out too early. Interestingly, the photoperiod limitation only occurs below a certain daylength (photoperiod threshold) when the warming increased above 4°C for beech trees. In contrast, for chestnut, no photoperiod threshold was found even when the ambient air temperature was warmed by 5°C. Given the species‐specific photoperiod effect on leaf phenology, the sequence of the leaf‐out timing among forest tree species may change under future climate warming conditions. Nonphotoperiodic species may benefit from warmer springs by starting the growing season earlier than photoperiodic sensitive species, modifying forest ecosystem structure and functions, but this photoperiod limitation needs to be further investigated experimentally in numerous species.     

The endemic kelp Lessonia corrugata is being pushed above its thermal limits in an ocean warming hotspot

Kelps are in global decline due to climate change, which includes ocean warming. To identify vulnerable species, we need to identify their tolerances to increasing temperatures and determine whether tolerances are altered by co-occurring drivers such as inorganic nutrient levels. This is particularly important for those species with restricted distributions, which may already be experiencing thermal stress. To identify thermal tolerance of the range-restricted kelp Lessonia corrugata, we conducted a laboratory experiment on juvenile sporophytes to measure performance (growth, photosynthesis) across its thermal range (4–22°C). We determined the upper thermal limit for growth and photosynthesis to be ~22–23°C, with a thermal optimum of ~16°C. To determine if elevated inorganic nitrogen availability could enhance thermal tolerance, we compared the performance of juveniles under low (4.5 μmol · d⁻¹) and high (90 μmol · d⁻¹) nitrate conditions at and above the thermal optimum (16–23.5°C). Nitrate enrichment did not enhance thermal performance at temperatures above the optimum but did lead to elevated growth rates at the thermal optimum. Our results indicate L. corrugata is likely to be extremely susceptible to moderate ocean warming and marine heatwaves. Peak sea surface temperatures during summer in eastern and northeastern Tasmania can reach up to 20–21°C, and climate projections suggest that L. corrugata's thermal limit will be regularly exceeded by 2050 as southeastern Australia is a global ocean-warming hotspot. By identifying the upper thermal limit of L. corrugata, we have taken a critical step in predicting the future of the species in a warming climate.     

Density-dependent responses of Picea purpurea seedlings for plant growth and resource allocation under elevated temperature

This study was conducted to determine whether plants in the presence or absence of competition differ in their responses to warming, and whether density modifies the effect of warming. Picea purourea seedlings were grown under ambient and warming (ambient +2.2 °C) conditions in climate control chambers with two different planting densities. After 4 years, seedlings were harvested and measured for height, stem diameter, leaf area, structural biomass, carbon, nitrogen, chlorophyll and carbohydrate levels of needles, branches, stem and roots. At low density, warming increased height, stem diameter, total leaf area biomass production and carbohydrate concentration per seedling, while it decreased C/N ratio for all plant parts, but did not affect chlorophyll content. By contrast, at high density, although warming increased biomass and total leaf area, it did not affect plant height and stem diameter. At the same time, it had different effects on chlorophyll content, C/N ratio and carbohydrate levels among plant parts. On the other hand, high density limited plant growth and altered resource allocation pattern. Our study demonstrates that planting densities decreased the temperature-induced growth enhancement of P. purpurea seedlings and the effects of warming on resource allocation not only showed density-dependence, but also vary with tissue age classes and root diameter; the responses of plants to elevated temperature, acquired from plants growing as individuals, may not be applicable to plants grown under intraspecific competition as typically found in the field.