Species‐specific adaptations explain resilience of herbaceous understorey to increased precipitation variability in a Mediterranean oak woodland

To date, the implications of the predicted greater intra‐annual variability and extremes in precipitation on ecosystem functioning have received little attention. This study presents results on leaf‐level physiological responses of five species covering the functional groups grasses, forbs, and legumes in the understorey of a Mediterranean oak woodland, with increasing precipitation variability, without altering total annual precipitation inputs. Although extending the dry period between precipitation events from 3 to 6 weeks led to increased soil moisture deficit, overall treatment effects on photosynthetic performance were not observed in the studied species. This resilience to prolonged water stress was explained by different physiological and morphological strategies to withstand periods below the wilting point, that is, isohydric behavior in Agrostis, Rumex, and Tuberaria, leaf succulence in Rumex, and taproots in Tolpis. In addition, quick recovery upon irrigation events and species‐specific adaptations of water‐use efficiency with longer dry periods and larger precipitation events contributed to the observed resilience in productivity of the annual plant community. Although none of the species exhibited a change in cover with increasing precipitation variability, leaf physiology of the legume Ornithopus exhibited signs of sensitivity to moisture deficit, which may have implications for the agricultural practice of seeding legume‐rich mixtures in Mediterranean grassland‐type systems. This highlights the need for long‐term precipitation manipulation experiments to capture possible directional changes in species composition and seed bank development, which can subsequently affect ecosystem state and functioning.     

Climatic and biotic extreme events moderate long‐term responses of above‐ and belowground sub‐Arctic heathland communities to climate change

Climate change impacts are not uniform across the Arctic region because interacting factors causes large variations in local ecosystem change. Extreme climatic events and population cycles of herbivores occur simultaneously against a background of gradual climate warming trends and can redirect ecosystem change along routes that are difficult to predict. Here, we present the results from sub‐Arctic heath vegetation and its belowground micro‐arthropod community in response to the two main drivers of vegetation damage in this region: extreme winter warming events and subsequent outbreaks of the defoliating autumnal moth caterpillar (Epirrita autumnata). Evergreen dwarf shrub biomass decreased (30%) following extreme winter warming events and again by moth caterpillar grazing. Deciduous shrubs that were previously exposed to an extreme winter warming event were not affected by the moth caterpillar grazing, while those that were not exposed to warming events (control plots) showed reduced (23%) biomass from grazing. Cryptogam cover increased irrespective of grazing or winter warming events. Micro‐arthropods declined (46%) following winter warming but did not respond to changes in plant community. Extreme winter warming and caterpillar grazing suppressed the CO₂ fluxes of the ecosystem. Evergreen dwarf shrubs are disadvantaged in a future sub‐Arctic with more stochastic climatic and biotic events. Given that summer warming may further benefit deciduous over evergreen shrubs, event and trend climate change may both act against evergreen shrubs and the ecosystem functions they provide. This is of particular concern given that Arctic heath vegetation is typically dominated by evergreen shrubs. Other components of the vegetation showed variable responses to abiotic and biotic events, and their interaction indicates that sub‐Arctic vegetation response to multiple pressures is not easy to predict from single‐factor responses. Therefore, while biotic and climatic events may have clear impacts, more work is needed to understand their net effect on Arctic ecosystems.     

A genome‐wide search for local adaptation in a terrestrial‐breeding frog reveals vulnerability to climate change

Terrestrial‐breeding amphibians are likely to be vulnerable to warming and drying climates, as their embryos require consistent moisture for successful development. Adaptation to environmental change will depend on sufficient genetic variation existing within or between connected populations. Here, we use Single Nucleotide Polymorphism (SNP) data to investigate genome‐wide patterns in genetic diversity, gene flow and local adaptation in a terrestrial‐breeding frog (Pseudophryne guentheri) subject to a rapidly drying climate and recent habitat fragmentation. The species was sampled across 12 central and range‐edge populations (192 samples), and strong genetic structure was apparent, as were high inbreeding coefficients. Populations showed differences in genetic diversity, and one population lost significant genetic diversity in a decade. More than 500 SNP loci were putatively under directional selection, and 413 of these loci were correlated with environmental variables such as temperature, rainfall, evaporation and soil moisture. One locus showed homology to a gene involved in the activation of maturation in Xenopus oocytes, which may facilitate rapid development of embryos in drier climates. The low genetic diversity, strong population structuring and presence of local adaptation revealed in this study shows why management strategies such as targeted gene flow may be necessary to assist isolated populations to adapt to future climates.     

Compensating for climate change–induced cue‐environment mismatches: evidence for contemporary evolution of a photoperiodic reaction norm in Colias butterflies

Anthropogenic climate change alters seasonal conditions without altering photoperiod and can thus create a cue‐environment mismatch for organisms that use photoperiod as a cue for seasonal plasticity. We investigated whether evolution of the photoperiodic reaction norm has compensated for this mismatch in Colias eurytheme. This butterfly’s wing melanization has a thermoregulatory function and changes seasonally. In 1971, Hoffmann quantified how larval photoperiod determines adult wing melanization. We recreated his experiment 47 years later using a contemporary population. Comparing our results to his, we found decreased melanization at short photoperiods but no change in melanization at long photoperiods, which is consistent with the greater increase in spring than summer temperatures recorded for this region. Our study shows that evolution can help correct cue‐environment mismatches but not in the same way under all conditions. Studies of contemporary evolution may miss important changes if they focus on only a limited range of conditions.     

Landscape dynamics in Mediterranean oak forests under global change: understanding the role of anthropogenic and environmental drivers across forest types

The Mediterranean region is projected to be extremely vulnerable to global change, which will affect the distribution of typical forest types such as native oak forests. However, our understanding of Mediterranean oak forest responses to future conditions is still very limited by the lack of knowledge on oak forest dynamics and species‐specific responses to multiple drivers. We compared the long‐term (1966–2006) forest persistence and land cover change among evergreen (cork oak and holm oak) and deciduous oak forests and evaluated the importance of anthropogenic and environmental drivers on observed changes for Portugal. We used National Forest Inventories to quantify the changes in oak forests and explored the drivers of change using multinomial logistic regression analysis and an information theoretical approach. We found distinct trends among oak forest types, reflecting the differences in oak economic value, protection status and management schemes: cork oak forests were the most persistent (62%), changing mostly to pines and eucalypt; holm oak forests were less persistent (53.2%), changing mostly to agriculture; and deciduous oak forests were the least persistent (45.7%), changing mostly to shrublands. Drivers of change had distinct importance across oak forest types, but drivers from anthropogenic origin (wildfires, population density, and land accessibility) were always among the most important. Climatic extremes were also important predictors of oak forest changes, namely extreme temperatures for evergreen oak forests and deficit of precipitation for deciduous oak forests. Our results indicate that under increasing human pressure and forecasted climate change, evergreen oak forests will continue declining and deciduous oak forests will be replaced by forests dominated by more xeric species. In the long run, multiple disturbances may change competitive dominance from oak forests to pyrophytic shrublands. A better understanding of forest dynamics and the inclusion of anthropogenic drivers on models of vegetation change will improve predicting the future of Mediterranean oak forests.     

Effects of extreme drought on specific leaf area of grassland species: A meta‐analysis of experimental studies in temperate and sub‐Mediterranean systems

Here, we conducted a meta‐analysis of experimental drought manipulation studies using rainout shelters in five sites of natural grassland ecosystems of Europe. The single studies assess the effects of extreme drought on the intraspecific variation of the specific leaf area (SLA), a proxy of plant growth. We evaluate and compare the effect size of the SLA response for the functional groups of forbs and grasses in temperate and sub‐Mediterranean systems. We hypothesized that the functional groups of grasses and forbs from temperate grassland systems have different strategies in short‐term drought response, measured as adjustment of SLA, with SLA‐reduction in grasses and SLA‐maintenance in forbs. Second, we hypothesized that grasses and forbs from sub‐Mediterranean systems do not differ in their drought response as both groups maintain their SLA. We found a significant decrease of SLA in grasses of the temperate systems in response to drought while SLA of forbs showed no significant response. Lower SLA is associated with enhanced water‐use efficiency under water stress and thus can be seen as a strategy of phenotypic adjustment. By contrast, in the sub‐Mediterranean systems, grasses significantly increased their SLA in the drought treatment. This result points towards a better growth performance of these grasses, which is most likely related to their strategy to allocate resources to belowground parts. The observed SLA reduction of forbs is most likely a direct drought response given that competitive effect of grasses is unlikely due to the scanty vegetation cover. We point out that phenotypic adjustment is an important driver of short‐term functional plant response to climatic extremes such as drought. Differential reactions of functional groups have to be interpreted against the background of the group's evolutionary configuration that can differ between climatic zones.     

Both life‐history plasticity and local adaptation will shape range‐wide responses to climate warming in the tundra plant Silene acaulis

Many predictions of how climate change will impact biodiversity have focused on range shifts using species‐wide climate tolerances, an approach that ignores the demographic mechanisms that enable species to attain broad geographic distributions. But these mechanisms matter, as responses to climate change could fundamentally differ depending on the contributions of life‐history plasticity vs. local adaptation to species‐wide climate tolerances. In particular, if local adaptation to climate is strong, populations across a species’ range—not only those at the trailing range edge—could decline sharply with global climate change. Indeed, faster rates of climate change in many high latitude regions could combine with local adaptation to generate sharper declines well away from trailing edges. Combining 15 years of demographic data from field populations across North America with growth chamber warming experiments, we show that growth and survival in a widespread tundra plant show compensatory responses to warming throughout the species’ latitudinal range, buffering overall performance across a range of temperatures. However, populations also differ in their temperature responses, consistent with adaptation to local climate, especially growing season temperature. In particular, warming begins to negatively impact plant growth at cooler temperatures for plants from colder, northern populations than for those from warmer, southern populations, both in the field and in growth chambers. Furthermore, the individuals and maternal families with the fastest growth also have the lowest water use efficiency at all temperatures, suggesting that a trade‐off between growth and water use efficiency could further constrain responses to forecasted warming and drying. Taken together, these results suggest that populations throughout species’ ranges could be at risk of decline with continued climate change, and that the focus on trailing edge populations risks overlooking the largest potential impacts of climate change on species’ abundance and distribution.     

Douglas‐fir plantations in Europe: a retrospective test of assisted migration to address climate change

We evaluate genetic test plantations of North American Douglas‐fir provenances in Europe to quantify how tree populations respond when subjected to climate regime shifts, and we examined whether bioclimate envelope models developed for North America to guide assisted migration under climate change can retrospectively predict the success of these provenance transfers to Europe. The meta‐analysis is based on long‐term growth data of 2800 provenances transferred to 120 European test sites. The model was generally well suited to predict the best performing provenances along north–south gradients in Western Europe, but failed to predict superior performance of coastal North American populations under continental climate conditions in Eastern Europe. However, model projections appear appropriate when considering additional information regarding adaptation of Douglas‐fir provenances to withstand frost and drought, even though the model partially fails in a validation against growth traits alone. We conclude by applying the partially validated model to climate change scenarios for Europe, demonstrating that climate trends observed over the last three decades warrant changes to current use of Douglas‐fir provenances in plantation forestry throughout Western and Central Europe.     

Eco‐evolutionary responses of Bromus tectorum to climate change: implications for biological invasions

How plant populations, communities, and ecosystems respond to climate change is a critical focus in ecology today. The responses of introduced species may be especially rapid. Current models that incorporate temperature and precipitation suggest that future Bromus tectorum invasion risk is low for the Colorado Plateau. With a field warming experiment at two sites in southeastern Utah, we tested this prediction over 4 years, measuring B. tectorum phenology, biomass, and reproduction. In a complimentary greenhouse study, we assessed whether changes in field B. tectorum biomass and reproductive output influence offspring performance. We found that following a wet winter and early spring, the timing of spring growth initiation, flowering, and summer senescence all advanced in warmed plots at both field sites and the shift in phenology was progressively larger with greater warming. Earlier green‐up and development was associated with increases in B. tectorum biomass and reproductive output, likely due early spring growth, when soil moisture was not limiting, and a lengthened growing season. Seeds collected from plants grown in warmed plots had higher biomass and germination rates and lower mortality than seeds from ambient plots. However, in the following two dry years, we observed no differences in phenology between warmed and ambient plots. In addition, warming had a generally negative effect on B. tectorum biomass and reproduction in dry years and this negative effect was significant in the plots that received the highest warming treatment. In contrast to models that predict negative responses of B. tectorum to warmer climate on the Colorado Plateau, the effects of warming were more nuanced, relied on background climate, and differed between the two field sites. Our results highlight the importance of considering the interacting effects of temperature, precipitation, and site‐specific characteristics such as soil texture, on plant demography and have direct implications for B. tectorum invasion dynamics on the Colorado Plateau.     

Evaluating within‐population variability in behavior and demography for the adaptive potential of a dispersal‐limited species to climate change

Multiple pathways exist for species to respond to changing climates. However, responses of dispersal‐limited species will be more strongly tied to ability to adapt within existing populations as rates of environmental change will likely exceed movement rates. Here, we assess adaptive capacity in Plethodon cinereus, a dispersal‐limited woodland salamander. We quantify plasticity in behavior and variation in demography to observed variation in environmental variables over a 5‐year period. We found strong evidence that temperature and rainfall influence P. cinereus surface presence, indicating changes in climate are likely to affect seasonal activity patterns. We also found that warmer summer temperatures reduced individual growth rates into the autumn, which is likely to have negative demographic consequences. Reduced growth rates may delay reproductive maturity and lead to reductions in size‐specific fecundity, potentially reducing population‐level persistence. To better understand within‐population variability in responses, we examined differences between two common color morphs. Previous evidence suggests that the color polymorphism may be linked to physiological differences in heat and moisture tolerance. We found only moderate support for morph‐specific differences for the relationship between individual growth and temperature. Measuring environmental sensitivity to climatic variability is the first step in predicting species' responses to climate change. Our results suggest phenological shifts and changes in growth rates are likely responses under scenarios where further warming occurs, and we discuss possible adaptive strategies for resulting selective pressures.     

Tissue chemistry and morphology affect root decomposition of perennial bioenergy grasses on sandy soil in a sub‐tropical environment

Second‐generation biofuels and bio‐based products derived from lignocellulosic biomass are likely to replace current fuels derived from simple sugars and starch because of greater yield potential and less competition with food production. Besides the high aboveground biomass production, these bioenergy grasses also exhibit extensive root systems. The decomposition of root biomass greatly influences nutrient cycling and microbial activity and subsequent accumulation of carbon (C) in the soil. The objective of this research was thus to characterize root morphological and chemical differences in six perennial grass species in order to better understand root decomposition and belowground C cycling of these bioenergy cropping systems. Giant reed (Arundo donax), elephantgrass (Pennisetum purpureum), energycane (Saccharum spp.), sugarcane (Saccharum spp.), sweetcane (Saccharum arundinaceum), and giant miscanthus (Miscanthus × giganteus) were established in Fall 2008 in research plots near Gainesville, Florida. Root decomposition rates were measured in situ from root decomposition bags over 12 months along with initial and final root tissue composition. Root potential decomposition rate constant (K) was higher in elephantgrass (3.64 g kg^?1 day^?1) and sweetcane (2.77 g kg^?1 day^?1) than in sugarcane (1.62 g kg^?1 day^?1) and energycane (1.48 g kg^?1 day^?1). Notably, K was positively related to initial root tissue total C (Total C), total fiber glucose (TFG), total fiber xylose (TFX), and total fiber carbohydrate (TFC) concentrations, but negatively related to total fiber arabinose (TFA) and lignin (TL) concentrations and specific root volume (SRV). Among the six species, elephantgrass exhibited root traits most favorable for fast decomposition: high TFG, high TFX, high TFC, high specific root length (SRL), and a low SRV, whereas giant reed, sugarcane, and energycane exhibited slow decomposition rates and the corresponding root traits. Thus, despite similar aboveground biomass yields in many cases, these species are likely to differentially affect soil C accumulation.     

Drought tolerance and growth in populations of a wide‐ranging tree species indicate climate change risks for the boreal north

Choosing drought‐tolerant planting stock in reforestation programs may help adapt forests to climate change. To inform such reforestation strategies, we test lodgepole pine (Pinus contorta Doug. ex Loud. var latifolia Englm.) population response to drought and infer potential benefits of a northward transfer of seeds from drier, southern environments. The objective is addressed by combining dendroecological growth analysis with long‐term genetic field trials. Over 500 trees originating from 23 populations across western North America were destructively sampled in three experimental sites in southern British Columbia, representing a climate warming scenario. Growth after 32 years from provenances transferred southward or northward over long distances was significantly lower than growth of local populations. All populations were affected by a severe natural drought event in 2002. The provenances from the most southern locations showed the highest drought tolerance but low productivity. Local provenances were productive and drought tolerant. Provenances from the boreal north showed lower productivity and less drought tolerance on southern test sites than all other sources, implying that maladaptation to drought may prevent boreal populations from taking full advantage of more favorable growing conditions under projected climate change.     

Climate‐associated genetic variation in Fagus sylvatica and potential responses to climate change in the French Alps

Local adaptation patterns have been found in many plants and animals, highlighting the genetic heterogeneity of species along their range of distribution. In the next decades, global warming is predicted to induce a change in the selective pressures that drive this adaptive variation, forcing a reshuffling of the underlying adaptive allele distributions. For species with low dispersion capacity and long generation time such as trees, the rapidity of the change could impede the migration of beneficial alleles and lower their capacity to track the changing environment. Identifying the main selective pressures driving the adaptive genetic variation is thus necessary when investigating species capacity to respond to global warming. In this study, we investigate the adaptive landscape of Fagus sylvatica along a gradient of populations in the French Alps. Using a double‐digest restriction‐site‐associated DNA (ddRAD) sequencing approach, we identified 7,000 SNPs from 570 individuals across 36 different sites. A redundancy analysis (RDA)‐derived method allowed us to identify several SNPs that were strongly associated with climatic gradients; moreover, we defined the primary selective gradients along the natural populations of F. sylvatica in the Alps. Strong effects of elevation and humidity, which contrast north‐western and south‐eastern site, were found and were believed to be important drivers of genetic adaptation. Finally, simulations of future genetic landscapes that used these findings allowed identifying populations at risk for F. sylvatica in the Alps, which could be helpful for future management plans.     

The implication of irrigation in climate change impact assessment: a European‐wide study

This study evaluates the impacts of projected climate change on irrigation requirements and yields of six crops (winter wheat, winter barley, rapeseed, grain maize, potato, and sugar beet) in Europe. Furthermore, the uncertainty deriving from consideration of irrigation, CO₂ effects on crop growth and transpiration, and different climate change scenarios in climate change impact assessments is quantified. Net irrigation requirement (NIR) and yields of the six crops were simulated for a baseline (1982–2006) and three SRES scenarios (B1, B2 and A1B, 2040–2064) under rainfed and irrigated conditions, using a process‐based crop model, SIMPLACE . We found that projected climate change decreased NIR of the three winter crops in northern Europe (up to 81 mm), but increased NIR of all the six crops in the Mediterranean regions (up to 182 mm yr^?1). Climate change increased yields of the three winter crops and sugar beet in middle and northern regions (up to 36%), but decreased their yields in Mediterranean countries (up to 81%). Consideration of CO₂ effects can alter the direction of change in NIR for irrigated crops in the south and of yields for C3 crops in central and northern Europe. Constraining the model to rainfed conditions for spring crops led to a negative bias in simulating climate change impacts on yields (up to 44%), which was proportional to the irrigation ratio of the simulation unit. Impacts on NIR and yields were generally consistent across the three SRES scenarios for the majority of regions in Europe. We conclude that due to the magnitude of irrigation and CO₂ effects, they should both be considered in the simulation of climate change impacts on crop production and water availability, particularly for crops and regions with a high proportion of irrigated crop area.     

Exploring sub‐individual variability: role of ontogeny,abiotic environment and seed‐dispersing birds

• Within‐individual trait variation – otherwise known as sub‐individual variation – is an important component of phenotypic variation, with both a genetic and epigenetic basis. We explore its adaptive value and the effects of ontogeny and the environment on sub‐individual variability.
• We conducted a field study to analyse the effects of tree age, soil pH, soil water content and soil nutrients on sub‐individual variability in fruit size of hawthorn (Crataegus monogyna) in three sites in northwest Spain. Additionally, we examined how bird‐mediated selection influences average and sub‐individual variation in fruit size.
• Results show that average and sub‐individual variations in fruit size were related to fitness affecting seed dispersal. Older trees produced larger fruits, but tree age did not affect sub‐individual variation in fruit size. Abiotic environmental factors differently affected sub‐individual variation and average fruit size. Seed‐dispersing birds exerted correlated selection on average and variation in fruit size, favouring trees with larger and less variable fruit size at one site.
• Our work suggests that the fruit size variation within individual trees, the sub‐individual variation, is modified by abiotic environmental factors and, additionally, is an adaptive trait that responds to natural selection.