Homoeostatic maintenance of nonstructural carbohydrates during the 2015–2016 El Niño drought across a tropical forest precipitation gradient

Nonstructural carbohydrates (NSCs) are essential for maintenance of plant metabolism and may be sensitive to short‐ and long‐term climatic variation. NSC variation in moist tropical forests has rarely been studied, so regulation of NSCs in these systems is poorly understood. We measured foliar and branch NSC content in 23 tree species at three sites located across a large precipitation gradient in Panama during the 2015–2016 El Niño to examine how short‐ and long‐term climatic variation impact carbohydrate dynamics. There was no significant difference in total NSCs as the drought progressed (leaf P = 0.32, branch P = 0.30) nor across the rainfall gradient (leaf P = 0.91, branch P = 0.96). Foliar soluble sugars decreased while starch increased over the duration of the dry period, suggesting greater partitioning of NSCs to storage than metabolism or transport as drought progressed. There was a large variation across species at all sites, but total foliar NSCs were positively correlated with leaf mass per area, whereas branch sugars were positively related to leaf temperature and negatively correlated with daily photosynthesis and wood density. The NSC homoeostasis across a wide range of conditions suggests that NSCs are an allocation priority in moist tropical forests.     

Causes of reduced leaf‐level photosynthesis during strong El Niño drought in a Central Amazon forest

Sustained drought and concomitant high temperature may reduce photosynthesis and cause tree mortality. Possible causes of reduced photosynthesis include stomatal closure and biochemical inhibition, but their relative roles are unknown in Amazon trees during strong drought events. We assessed the effects of the recent (2015) strong El Niño drought on leaf‐level photosynthesis of Central Amazon trees via these two mechanisms. Through four seasons of 2015, we measured leaf gas exchange, chlorophyll a fluorescence parameters, chlorophyll concentration, and nutrient content in leaves of 57 upper canopy and understory trees of a lowland terra firme forest on well‐drained infertile oxisol. Photosynthesis decreased 28% in the upper canopy and 17% in understory trees during the extreme dry season of 2015, relative to other 2015 seasons and was also lower than the climatically normal dry season of the following non‐El Niño year. Photosynthesis reduction under extreme drought and high temperature in the 2015 dry season was related only to stomatal closure in both upper canopy and understory trees, and not to chlorophyll a fluorescence parameters, chlorophyll, or leaf nutrient concentration. The distinction is important because stomatal closure is a transient regulatory response that can reverse when water becomes available, whereas the other responses reflect more permanent changes or damage to the photosynthetic apparatus. Photosynthesis decrease due to stomatal closure during the 2015 extreme dry season was followed 2 months later by an increase in photosynthesis as rains returned, indicating a margin of resilience to one‐off extreme climatic events in Amazonian forests.     

Potential impacts of a future persistent El Niño or La Niña on three subspecies of Australian butterflies

One of the major uncertainties of 21st century climate change is the potential for shifts to the intensity and frequency of the El Niño Southern Oscillation (ENSO) cycle. Although this phenomenon is known to have dramatic impacts on ecosystems regionally and globally, the biological consequences of climate change‐driven shifts in future ENSO events have been unexplored. Here, we investigate the potential impacts that a persistent El Niño, La Niña, or ‘Neutral' phase may have on species distributions. Using MaxEnt, we model the distribution of climatically suitable habitat for three northeast Australian butterfly subspecies (Doleschallia bisaltide australis, Hypolimnas alimena lamina, and Mycalesis terminus terminus) across the three ENSO phases. We find that the spatial extent and quality of habitat are lowest under conditions that would characterize a persistent El Niño (hot/dry). In contrast, suitable habitat is broadest under the warm/wet conditions associated with La Niña. Statistical analyses of the difference between pair‐wise combinations of suitability maps using Hellinger distance showed that projections for each subspecies and ENSO phase combination were significantly different from other combinations. The resilience of these, and other, butterfly (sub)species to changes in ENSO will be influenced by fluctuations in the strength of these events, availability of refugia, and life‐history characteristics. However, the population dynamics of wet‐ and dry‐season phenotypes of M. t. terminus and physiological limitations to high temperatures suggest that this subspecies, in particular, may have limited resilience should the strength and frequency of El Niño events increase.     

Quantifying drylands' drought resistance and recovery: the importance of drought intensity,dominant life history and grazing regime

Projected global change will increase the level of land‐use and environmental stressors such as drought and grazing, particularly in drylands. Still, combined effects of drought and grazing on plant production are poorly understood, thus hampering adequate projections and development of mitigation strategies. We used a large, cross‐continental database consisting of 174 long‐term datasets from >30 dryland regions to quantify ecosystem responses to drought and grazing with the ultimate goal to increase functional understanding in these responses. Two key aspects of ecosystem stability, resistance to and recovery after a drought, were evaluated based on standardized and normalized aboveground net primary production (ANPP) data. Drought intensity was quantified using the standardized precipitation index. We tested effects of drought intensity, grazing regime (grazed, ungrazed), biome (grassland, shrubland, savanna) or dominant life history (annual, perennial) of the herbaceous layer to assess the relative importance of these factors for ecosystem stability, and to identify predictable relationships between drought intensity and ecosystem resistance and recovery. We found that both components of ecosystem stability were better explained by dominant herbaceous life history than by biome. Increasing drought intensity (quasi‐) linearly reduced ecosystem resistance. Even though annual and perennial systems showed the same response rate to increasing drought intensity, they differed in their general magnitude of resistance, with annual systems being ca. 27% less resistant. In contrast, systems with an herbaceous layer dominated by annuals had substantially higher postdrought recovery, particularly when grazed. Combined effects of drought and grazing were not merely additive but modulated by dominant life history of the herbaceous layer. To the best of our knowledge, our study established the first predictive, cross‐continental model between drought intensity and drought‐related relative losses in ANPP, and suggests that systems with an herbaceous layer dominated by annuals are more prone to ecosystem degradation under future global change regimes.     

Seasonal variability of forest sensitivity to heat and drought stresses: A synthesis based on carbon fluxes from North American forest ecosystems

Climate extremes such as heat waves and droughts are projected to occur more frequently with increasing temperature and an intensified hydrological cycle. It is important to understand and quantify how forest carbon fluxes respond to heat and drought stress. In this study, we developed a series of daily indices of sensitivity to heat and drought stress as indicated by air temperature (T_a) and evaporative fraction (EF). Using normalized daily carbon fluxes from the FLUXNET Network for 34 forest sites in North America, the seasonal pattern of sensitivities of net ecosystem productivity (NEP), gross ecosystem productivity (GEP) and ecosystem respiration (RE) in response to T_a and EF anomalies were compared for different forest types. The results showed that warm temperatures in spring had a positive effect on NEP in conifer forests but a negative impact in deciduous forests. GEP in conifer forests increased with higher temperature anomalies in spring but decreased in summer. The drought‐induced decrease in NEP, which mostly occurred in the deciduous forests, was mostly driven by the reduction in GEP. In conifer forests, drought had a similar dampening effect on both GEP and RE, therefore leading to a neutral NEP response. The NEP sensitivity to T_a anomalies increased with increasing mean annual temperature. Drier sites were less sensitive to drought stress in summer. Natural forests with older stand age tended to be more resilient to the climate stresses compared to managed younger forests. The results of the Classification and Regression Tree analysis showed that seasons and ecosystem productivity were the most powerful variables in explaining the variation of forest sensitivity to heat and drought stress. Our results implied that the magnitude and direction of carbon flux changes in response to climate extremes are highly dependent on the seasonal dynamics of forests and the timing of the climate extremes.     

Resistance of European tree species to drought stress in mixed versus pure forests: evidence of stress release by inter‐specific facilitation

While previous studies focused on tree growth in pure stands, we reveal that tree resistance and resilience to drought stress can be modified distinctly through species mixing. Our study is based on tree ring measurement on cores from increment boring of 559 trees of Norway spruce (Picea abies [L.] Karst.), European beech (Fagus sylvatica [L.]) and sessile oak (Quercus petraea (Matt.) Liebl.) in South Germany, with half sampled in pure, respectively, mixed stands. Indices for resistance, recovery and resilience were applied for quantifying the tree growth reaction on the episodic drought stress in 1976 and 2003. The following general reaction patterns were found. (i) In pure stands, spruce has the lowest resistance, but the quickest recovery; oak and beech were more resistant, but recover was much slower and they are less resilient. (ii) In mixture, spruce and oak perform as in pure stands, but beech was significantly more resistant and resilient than in monoculture. (iii) Especially when mixed with oak, beech is facilitated. We hypothesise that the revealed water stress release of beech emerges in mixture because of the asynchronous stress reaction pattern of beech and oak and a facilitation of beech by hydraulic lift of water by oak. This facilitation of beech in mixture with oak means a contribution to the frequently reported overyield of beech in mixed versus pure stands. We discuss the far‐reaching implications that these differences in stress response under intra‐ and inter‐specific environments have for forest ecosystem dynamics and management under climate change.     

Deadly combination of genes and drought: increased mortality of herbivore-resistant trees in a foundation species

Current climate models predict a shift to warmer, drier conditions in the southwestern US. While major shifts in plant distribution are expected to follow these climate changes, interactions among species and intraspecific genetic variation rarely have been incorporated into models of future plant distributions. We examined the drought‐related mortality of pinyon pine (Pinus edulis) in northern Arizona focusing on trees that showed genetically‐based resistance or susceptibility to a nonlethal herbivore, the shoot‐boring moth, Dioryctria albovittella. Because moth resistant trees have outperformed susceptible trees during 20 years of study, and herbivory has been shown to increase drought related mortality, we expected higher mortality rates in susceptible trees. However, our field observations and greenhouse experiments showed several unexpected patterns relevant to understanding the consequences of climate change: (1) The mortality of adult P. edulis resistant to the moth was three times higher than the mortality of trees susceptible to the moth. (2) Over a few years, differential mortality caused a shift in stand structure from resistant dominated to equality (3 : 1 resistant : susceptible to 1 : 1). (3) Adult moth resistant trees suffered significantly greater water stress than adult moth susceptible trees, suggesting that variation among the two groups in drought tolerance may be a mechanism for differential mortality. (4) When grown under drought conditions in the greenhouse, seedlings from resistant mothers died sooner than seedlings from susceptible mothers. These data support the hypothesis that drought can act as an agent of balancing selection and that drought resistance is a heritable trait. Taken together, our findings suggest that genetic variation in a population can be an important factor in determining its response to future climate change, and argue for the inclusion of genetics into models developed to understand the consequences of climate change.     

Soil microbial moisture dependences and responses to drying–rewetting: The legacy of 18 years drought

Climate change will alter precipitation patterns with consequences for soil C cycling. An understanding of how fluctuating soil moisture affects microbial processes is therefore critical to predict responses to future global change. We investigated how long‐term experimental field drought influences microbial tolerance to lower moisture levels (“resistance”) and ability to recover when rewetted after drought (“resilience”), using soils from a heathland which had been subjected to experimental precipitation reduction during the summer for 18 years. We tested whether drought could induce increased resistance, resilience, and changes in the balance between respiration and bacterial growth during perturbation events, by following a two‐tiered approach. We first evaluated the effects of the long‐term summer drought on microbial community functioning to drought and drying–rewetting (D/RW), and second tested the ability to alter resistance and resilience through additional perturbation cycles. A history of summer drought in the field selected for increased resilience but not resistance, suggesting that rewetting after drought, rather than low moisture levels during drought, was the selective pressure shaping the microbial community functions. Laboratory D/RW cycles also selected for communities with a higher resilience rather than increased resistance. The ratio of respiration to bacterial growth during D/RW perturbation was lower for the field drought‐exposed communities and decreased for both field treatments during the D/RW cycles. This suggests that cycles of D/RW also structure microbial communities to respond quickly and efficiently to rewetting after drought. Our findings imply that microbial communities can adapt to changing climatic conditions and that this might slow the rate of soil C loss predicted to be induced by future cyclic drought.     

Long‐term nitrogen additions increase likelihood of climate stress and affect recovery from wildfire in a lowland heath

Increases in the emissions and associated atmospheric deposition of nitrogen (N) have the potential to cause significant changes to the structure and function of N‐limited ecosystems. Here, we present the results of a long‐term (13 year) experiment assessing the impacts of N addition (30 kg ha^?1 yr^?1) on a UK lowland heathland under a wide range of environmental conditions, including the occurrence of prolonged natural drought episodes and a severe summer fire. Our findings indicate that elevated N deposition results in large, persistent effects on Calluna growth, phenology and chemistry, severe suppression of understorey lichen flora and changes in soil biogeochemistry. Growing season rainfall was found to be a strong driver of inter‐annual variation in Calluna growth and, although interactions between N and rainfall for shoot growth were not significant until the later phase of the experiment, N addition exacerbated the extent of drought injury to Calluna shoots following naturally occurring droughts in 2003 and 2009. Following a severe wildfire at the experimental site in 2006, heathland regeneration dynamics were significantly affected by N, with a greater abundance of pioneering moss species and suppression of the lichen flora in plots receiving N additions. Significant interactions between climate and N were also apparent post fire, with the characteristic stimulation in Calluna growth in +N plots suppressed during dry years. Carbon (C) and N budgets demonstrate large increases in both above‐ and below‐ground stocks of these elements in N‐treated plots prior to the fire, despite higher levels of soil microbial activity and organic matter turnover. Although much of the organic material was removed during the fire, pre‐existing treatment differences were still evident following the burn. Post fire accumulation of below‐ground C and N stocks was increased rapidly in N‐treated plots, highlighting the role of N deposition in ecosystem C sequestration.     

Tests of species‐specific models reveal the importance of drought in postglacial range shifts of a Mediterranean‐climate tree: insights from integrative distributional,demographic and coalescent modelling and ABC model selection

Past climate change has caused shifts in species distributions and undoubtedly impacted patterns of genetic variation, but the biological processes mediating responses to climate change, and their genetic signatures, are often poorly understood. We test six species‐specific biologically informed hypotheses about such processes in canyon live oak (Quercus chrysolepis) from the California Floristic Province. These hypotheses encompass the potential roles of climatic niche, niche multidimensionality, physiological trade‐offs in functional traits, and local‐scale factors (microsites and local adaptation within ecoregions) in structuring genetic variation. Specifically, we use ecological niche models (ENMs) to construct temporally dynamic landscapes where the processes invoked by each hypothesis are reflected by differences in local habitat suitabilities. These landscapes are used to simulate expected patterns of genetic variation under each model and evaluate the fit of empirical data from 13 microsatellite loci genotyped in 226 individuals from across the species range. Using approximate Bayesian computation (ABC), we obtain very strong support for two statistically indistinguishable models: a trade‐off model in which growth rate and drought tolerance drive habitat suitability and genetic structure, and a model based on the climatic niche estimated from a generic ENM, in which the variables found to make the most important contribution to the ENM have strong conceptual links to drought stress. The two most probable models for explaining the patterns of genetic variation thus share a common component, highlighting the potential importance of seasonal drought in driving historical range shifts in a temperate tree from a Mediterranean climate where summer drought is common.     

Seed germination responses to seasonal temperature and drought stress are species‐specific but not related to seed size in a desert steppe: Implications for effect of climate change on community structure

Investigating how seed germination of multiple species in an ecosystem responds to environmental conditions is crucial for understanding the mechanisms for community structure and biodiversity maintenance. However, knowledge of seed germination response of species to environmental conditions is still scarce at the community level. We hypothesized that responses of seed germination to environmental conditions differ among species at the community level, and that germination response is not correlated with seed size. To test this hypothesis, we determined the response of seed germination of 20 common species in the Siziwang Desert Steppe, China, to seasonal temperature regimes (representing April, May, June, and July) and drought stress (0, ?0.003, ?0.027, ?0.155, and ?0.87 MPa). Seed germination percentage increased with increasing temperature regime, but Allium ramosum, Allium tenuissimum, Artemisia annua, Artemisia mongolica, Artemisia scoparia, Artemisia sieversiana, Bassia dasyphylla, Kochia prastrata, and Neopallasia pectinata germinated to >60% in the lowest temperature regime (April). Germination decreased with increasing water stress, but Allium ramosum, Artemisia annua, Artemisia scoparia, Bassia dasyphylla, Heteropappus altaicus, Kochia prastrata, Neopallasia pectinata, and Potentilla tanacetifolia germinated to near 60% at ?0.87 MPa. Among these eight species, germination of six was tolerant to both temperature and water stress. Mean germination percentage in the four temperature regimes and the five water potentials was not significantly correlated with seed mass or seed area, which were highly correlated. Our results suggest that the species‐specific germination responses to environmental conditions are important in structuring the desert steppe community and have implications for predicting community structure under climate change. Thus, the predicted warmer and dryer climate will favor germination of drought‐tolerant species, resulting in altered proportions of germinants of different species and subsequently change in community composition of the desert steppe.     

Response of plant species richness and primary productivity in shrublands along a north–south gradient in Europe to seven years of experimental warming and drought: reductions in primary productivity in the heat and drought year of 2003

We used a nonintrusive field experiment carried out at six sites – Wales (UK), Denmark (DK), the Netherlands (NL), Hungary (HU), Sardinia (Italy – IT), and Catalonia (Spain – SP) – along a climatic and latitudinal gradient to examine the response of plant species richness and primary productivity to warming and drought in shrubland ecosystems. The warming treatment raised the plot daily temperature by ca. 1 °C, while the drought treatment led to a reduction in soil moisture at the peak of the growing season that ranged from 26% at the SP site to 82% in the NL site. During the 7 years the experiment lasted (1999–2005), we used the pin‐point method to measure the species composition of plant communities and plant biomass, litterfall, and shoot growth of the dominant plant species at each site. A significantly lower increase in the number of species pin‐pointed per transect was found in the drought plots at the SP site, where the plant community was still in a process of recovering from a forest fire in 1994. No changes in species richness were found at the other sites, which were at a more mature and stable state of succession and, thus less liable to recruitment of new species. The relationship between annual biomass accumulation and temperature of the growing season was positive at the coldest site and negative at the warmest site. The warming treatment tended to increase the aboveground net primary productivity (ANPP) at the northern sites. The relationship between annual biomass accumulation and soil moisture during the growing season was not significant at the wettest sites, but was positive at the driest sites. The drought treatment tended to reduce the ANPP in the NL, HU, IT, and SP sites. The responses to warming were very strongly related to the Gaussen aridity index (stronger responses the lower the aridity), whereas the responses to drought were not. Changes in the annual aboveground biomass accumulation, litterfall, and, thus, the ANPP, mirrored the interannual variation in climate conditions: the most outstanding change was a decrease in biomass accumulation and an increase in litterfall at most sites during the abnormally hot year of 2003. Species richness also tended to decrease in 2003 at all sites except the cold and wet UK site. Species‐specific responses to warming were found in shoot growth: at the SP site, Globularia alypum was not affected, while the other dominant species, Erica multiflora, grew 30% more; at the UK site, Calluna vulgaris tended to grow more in the warming plots, while Empetrum nigrum tended to grow less. Drought treatment decreased plant growth in several studied species, although there were some species such as Pinus halepensis at the SP site or C. vulgaris at the UK site that were not affected. The magnitude of responses to warming and drought thus depended greatly on the differences between sites, years, and species and these multiple plant responses may be expected to have consequences at ecosystem and community level. Decreases in biodiversity and the increase in E. multiflora growth at the SP site as a response to warming challenge the assumption that sensitivity to warming may be less well developed at more southerly latitudes; likewise, the fact that one of the studied shrublands presented negative ANPP as a response to the 2003 heat wave also challenges the hypothesis that future climate warming will lead to an enhancement of plant growth and carbon sequestration in temperate ecosystems. Extreme events may thus change the general trend of increased productivity in response to warming in the colder sites.     

Reconciling estimates of the contemporary North American carbon balance among terrestrial biosphere models,atmospheric inversions,and a new approach for estimating net ecosystem exchange from inventory‐based data

We develop an approach for estimating net ecosystem exchange (NEE) using inventory‐based information over North America (NA) for a recent 7‐year period (ca. 2000–2006). The approach notably retains information on the spatial distribution of NEE, or the vertical exchange between land and atmosphere of all non‐fossil fuel sources and sinks of CO₂, while accounting for lateral transfers of forest and crop products as well as their eventual emissions. The total NEE estimate of a ?327 ± 252 TgC yr^?1 sink for NA was driven primarily by CO₂ uptake in the Forest Lands sector (?248 TgC yr^?1), largely in the Northwest and Southeast regions of the US, and in the Crop Lands sector (?297 TgC yr^?1), predominantly in the Midwest US states. These sinks are counteracted by the carbon source estimated for the Other Lands sector (+218 TgC yr^?1), where much of the forest and crop products are assumed to be returned to the atmosphere (through livestock and human consumption). The ecosystems of Mexico are estimated to be a small net source (+18 TgC yr^?1) due to land use change between 1993 and 2002. We compare these inventory‐based estimates with results from a suite of terrestrial biosphere and atmospheric inversion models, where the mean continental‐scale NEE estimate for each ensemble is ?511 TgC yr^?1 and ?931 TgC yr^?1, respectively. In the modeling approaches, all sectors, including Other Lands, were generally estimated to be a carbon sink, driven in part by assumed CO₂ fertilization and/or lack of consideration of carbon sources from disturbances and product emissions. Additional fluxes not measured by the inventories, although highly uncertain, could add an additional ?239 TgC yr^?1 to the inventory‐based NA sink estimate, thus suggesting some convergence with the modeling approaches.     

Nitrogen deposition outweighs climatic variability in driving annual growth rate of canopy beech trees: Evidence from long‐term growth reconstruction across a geographic gradient

In this study, we investigated the role of climatic variability and atmospheric nitrogen deposition in driving long‐term tree growth in canopy beech trees along a geographic gradient in the montane belt of the Italian peninsula, from the Alps to the southern Apennines. We sampled dominant trees at different developmental stages (from young to mature tree cohorts, with tree ages spanning from 35 to 160 years) and used stem analysis to infer historic reconstruction of tree volume and dominant height. Annual growth volume (G_V) and height (G_H) variability were related to annual variability in model simulated atmospheric nitrogen deposition and site‐specific climatic variables, (i.e. mean annual temperature, total annual precipitation, mean growing period temperature, total growing period precipitation, and standard precipitation evapotranspiration index) and atmospheric CO₂ concentration, including tree cambial age among growth predictors. Generalized additive models (GAM), linear mixed‐effects models (LMM), and Bayesian regression models (BRM) were independently employed to assess explanatory variables. The main results from our study were as follows: (i) tree age was the main explanatory variable for long‐term growth variability; (ii) GAM, LMM, and BRM results consistently indicated climatic variables and CO₂ effects on G_V and G_H were weak, therefore evidence of recent climatic variability influence on beech annual growth rates was limited in the montane belt of the Italian peninsula; (iii) instead, significant positive nitrogen deposition (N_dep) effects were repeatedly observed in G_V and G_H; the positive effects of N_dep on canopy height growth rates, which tended to level off at N_dep values greater than approximately 1.0 g m^?2 y^?1, were interpreted as positive impacts on forest stand above‐ground net productivity at the selected study sites.