Legacy effects of premature defoliation in response to an extreme drought event modulate phytochemical profiles with subtle consequences for leaf herbivory in European beech.

Extreme droughts can have long-lasting effects on forest community dynamics and species interactions. Yet, our understanding of how drought legacy modulates ecological relationships is just unfolding. We tested the hypothesis that leaf chemistry and herbivory show long-term responses to premature defoliation caused by an extreme drought event in European beech (Fagus sylvatica L.). For two consecutive years after the extreme European summer drought in 2018, we collected leaves from the upper and lower canopy of adjacently growing drought-stressed and unstressed trees. Leaf chemistry was analyzed and leaf damage by different herbivore-feeding guilds was quantified. We found that drought had lasting impacts on leaf nutrients and on specialized metabolomic profiles. However, drought did not affect the primary metabolome. Drought-related phytochemical changes affected damage of leaf-chewing herbivores whereas damage caused by other herbivore-feeding guilds was largely unaffected. Drought legacy effects on phytochemistry and herbivory were often weaker than between-year or between-canopy strata variability. Our findings suggest that a single extreme drought event bears the potential to long-lastingly affect tree-herbivore interactions. Drought legacy effects likely become more important in modulating tree-herbivore interactions since drought frequency and severity are projected to globally increase in the coming decades.

Slower growth prior to the 2018 drought and a high growth sensitivity to previous year summer conditions predisposed European beech to crown dieback.

The record-breaking drought in 2018 caused premature leaf discoloration and shedding (early browning) in many beech (Fagus sylvatica L.) dominated forests in Central Europe. However, a high degree of variability in drought response among individual beech trees was observed. While some trees were severely impacted by the prolonged water deficits and high temperatures, others remained vital with no or only minor signs of crown vitality loss. Why some beech trees were more susceptible to drought-induced crown damage than others and whether growth recovery is possible are poorly understood. Here, we aimed to identify growth characteristics associated with the variability in drought response between individual beech trees based on a sample of 470 trees in northern Switzerland. By combining tree growth measurements and crown condition assessments, we also investigated the possible link between crown dieback and growth recovery after drought. Beech trees with early browning exhibited an overall lower growth vigor before the 2018 drought than co-occurring vital beech trees. This lower vigor is mainly indicated by lower overall growth rates, stronger growth declines in the past decades, and higher growth-climate sensitivity. Particularly, warm previous year summer conditions negatively affected current growth of the early-browning trees. These findings suggest that the affected trees had less access to critical resources and were physiologically limited in their growth predisposing them to early browning. Following the 2018 drought, observed growth recovery potential corresponded to the amount of crown dieback and the local climatic water balance. Overall, our findings emphasize that beech-dominated forests in Central Europe are under increasing pressure from severe droughts, ultimately reducing the competitive ability of this species, especially on lowland sites with shallow soils and low water holding capacity.

Plant traits poorly predict winner and loser shrub species in a warming tundra biome.

Climate change is leading to species redistributions. In the tundra biome, shrubs are generally expanding, but not all tundra shrub species will benefit from warming. Winner and loser species, and the characteristics that may determine success or failure, have not yet been fully identified. Here, we investigate whether past abundance changes, current range sizes and projected range shifts derived from species distribution models are related to plant trait values and intraspecific trait variation. We combined 17,921 trait records with observed past and modelled future distributions from 62 tundra shrub species across three continents. We found that species with greater variation in seed mass and specific leaf area had larger projected range shifts, and projected winner species had greater seed mass values. However, trait values and variation were not consistently related to current and projected ranges, nor to past abundance change. Overall, our findings indicate that abundance change and range shifts will not lead to directional modifications in shrub trait composition, since winner and loser species share relatively similar trait spaces.

Assessing the Effectiveness of in-situ Active Warming Combined With Open Top Chambers to Study Plant Responses to Climate Change.

Temperature manipulation experiments are an effective way for testing plant responses to future climate conditions, especially for predicting shifts in plant phenological events. While passive warming techniques are widely used to elevate temperature in low stature plant communities, active warming has been applied less frequently due to the associated resource requirements. In forest ecosystems, however, active warming is crucial to simulate projected air temperature rises of 3-5 K, especially at the warm (i.e., southern and low elevation) range edges of tree species. Moreover, the warming treatment should be applied to the complete height of the experimental plants, e.g., regenerating trees in the understory. Here, we combined open top chambers (OTCs) with active heat sources, an electric heater (OTC-EH) and warming cables (OTC-WC), and tested the effectiveness of these set-ups to maintain constant temperature differences compared to ambient temperature across 18 m2 plots. This chamber size is needed to grow tree saplings in mixture in forest gaps for 3 to 10 years. With passive warming only, an average temperature increase of approx. 0.4 K as compared to ambient conditions was achieved depending on time of the day and weather conditions. In the actively warmed chambers, average warming exceeded ambient temperatures by 2.5 to 2.8 K and was less variable over time. However, active warming also reduced air humidity by about 15%. These results underline the need to complement passive warming with active warming in order to achieve constant temperature differences appropriate for climate change simulations under all weather conditions in large OTCs. Since we observed considerable horizontal and vertical temperature variation within OTCs with temperature differences of up to 16.9 K, it is essential to measure and report within-plot temperature distribution as well as temporal temperature variation. If temperature distributions within large OTCs are well characterized, they may be incorporated in the experimental design helping to identify non-linear or threshold responses to warming.

Plant functional trait change across a warming tundra biome.

The tundra is warming more rapidly than any other biome on Earth, and the potential ramifications are far-reaching because of global feedback effects between vegetation and climate. A better understanding of how environmental factors shape plant structure and function is crucial for predicting the consequences of environmental change for ecosystem functioning. Here we explore the biome-wide relationships between temperature, moisture and seven key plant functional traits both across space and over three decades of warming at 117 tundra locations. Spatial temperature-trait relationships were generally strong but soil moisture had a marked influence on the strength and direction of these relationships, highlighting the potentially important influence of changes in water availability on future trait shifts in tundra plant communities. Community height increased with warming across all sites over the past three decades, but other traits lagged far behind predicted rates of change. Our findings highlight the challenge of using space-for-time substitution to predict the functional consequences of future warming and suggest that functions that are tied closely to plant height will experience the most rapid change. They also reveal the strength with which environmental factors shape biotic communities at the coldest extremes of the planet and will help to improve projections of functional changes in tundra ecosystems with climate warming.

Biotic and abiotic drivers of tree seedling recruitment across an alpine treeline ecotone.

Treeline responses to climate change ultimately depend on successful seedling recruitment, which requires dispersal of viable seeds and establishment of individual propagules in novel environments. In this study, we evaluated the effects of several abiotic and biotic drivers of early tree seedling recruitment across an alpine treeline ecotone. In two consecutive years, we sowed seeds of low- and high-elevation provenances of Larix decidua (European larch) and Picea abies (Norway spruce) below, at, and above the current treeline into intact vegetation and into open microsites with artificially removed surface vegetation, as well as into plots protected from seed predators and herbivores. Seedling emergence and early establishment in treatment and in control plots were monitored over two years. Tree seedling emergence occurred at and several hundred metres above the current treeline when viable seeds and suitable microsites for germination were available. However, dense vegetation cover at lower elevations and winter mortality at higher elevations particularly limited early recruitment. Post-dispersal predation, species, and provenance also affected emergence and early establishment. This study demonstrates the importance of understanding multiple abiotic and biotic drivers of early seedling recruitment that should be incorporated into predictions of treeline dynamics under climate change.

Climate adaptation is not enough: warming does not facilitate success of southern tundra plant populations in the high Arctic.

Rapidly rising temperatures are expected to cause latitudinal and elevational range shifts as species track their optimal climate north and upward. However, a lack of adaptation to environmental conditions other than climate - for example photoperiod, biotic interactions, or edaphic conditions - might limit the success of immigrants in a new location despite hospitable climatic conditions. Here, we present one of the first direct experimental tests of the hypothesis that warmer temperatures at northern latitudes will confer a fitness advantage to southern immigrants relative to native populations. As rates of warming in the Arctic are more than double the global average, understanding the impacts of warming in Arctic ecosystems is especially urgent. We established experimentally warmed and nonwarmed common garden plots at Alexandra Fiord, Ellesmere Island in the Canadian High Arctic with seeds of two forb species (Oxyria digyna and Papaver radicatum) originating from three to five populations at different latitudes across the Arctic. We found that plants from the local populations generally had higher survival and obtained a greater maximum size than foreign individuals, regardless of warming treatment. Phenological traits varied with latitude of the source population, such that southern populations demonstrated substantially delayed leaf-out and senescence relative to northern populations. Our results suggest that environmental conditions other than temperature may influence the ability of foreign populations and species to establish at more northerly latitudes as the climate warms, potentially leading to lags in northward range shifts for some species.

Plastic responses to elevated temperature in low and high elevation populations of three grassland species.

Local persistence of plant species in the face of climate change is largely mediated by genetic adaptation and phenotypic plasticity. In species with a wide altitudinal range, population responses to global warming are likely to differ at contrasting elevations. In controlled climate chambers, we investigated the responses of low and high elevation populations (1200 and 1800 m a.s.l.) of three nutrient-poor grassland species, Trifolium montanum, Ranunculus bulbosus, and Briza media, to ambient and elevated temperature. We measured growth-related, reproductive and phenological traits, evaluated differences in trait plasticity and examined whether trait values or plasticities were positively related to approximate fitness and thus under selection. Elevated temperature induced plastic responses in several growth-related traits of all three species. Although flowering phenology was advanced in T. montanum and R. bulbosus, number of flowers and reproductive allocation were not increased under elevated temperature. Plasticity differed between low and high elevation populations only in leaf traits of T. montanum and B. media. Some growth-related and phenological traits were under selection. Moreover, plasticities were not correlated with approximate fitness indicating selectively neutral plastic responses to elevated temperature. The observed plasticity in growth-related and phenological traits, albeit variable among species, suggests that plasticity is an important mechanism in mediating plant responses to elevated temperature. However, the capacity of species to respond to climate change through phenotypic plasticity is limited suggesting that the species additionally need evolutionary adaptation to adjust to climate change. The observed selection on several growth-related and phenological traits indicates that the study species have the potential for future evolution in the context of a warming climate.

Plant population differentiation and climate change: responses of grassland species along an elevational gradient.

Mountain ecosystems are particularly susceptible to climate change. Characterizing intraspecific variation of alpine plants along elevational gradients is crucial for estimating their vulnerability to predicted changes. Environmental conditions vary with elevation, which might influence plastic responses and affect selection pressures that lead to local adaptation. Thus, local adaptation and phenotypic plasticity among low and high elevation plant populations in response to climate, soil and other factors associated with elevational gradients might underlie different responses of these populations to climate warming. Using a transplant experiment along an elevational gradient, we investigated reproductive phenology, growth and reproduction of the nutrient-poor grassland species Ranunculus bulbosus, Trifolium montanum and Briza media. Seeds were collected from low and high elevation source populations across the Swiss Alps and grown in nine common gardens at three different elevations with two different soil depths. Despite genetic differentiation in some traits, the results revealed no indication of local adaptation to the elevation of population origin. Reproductive phenology was advanced at lower elevation in low and high elevation populations of all three species. Growth and reproduction of T. montanum and B. media were hardly affected by garden elevation and soil depth. In R. bulbosus, however, growth decreased and reproductive investment increased at higher elevation. Furthermore, soil depth influenced growth and reproduction of low elevation R. bulbosus populations. We found no evidence for local adaptation to elevation of origin and hardly any differences in the responses of low and high elevation populations. However, the consistent advanced reproductive phenology observed in all three species shows that they have the potential to plastically respond to environmental variation. We conclude that populations might not be forced to migrate to higher elevations as a consequence of climate warming, as plasticity will buffer the detrimental effects of climate change in the three investigated nutrient-poor grassland species.

Patterns of genetic variation across altitude in three plant species of semi-dry grasslands.

BACKGROUND: Environmental gradients caused by altitudinal gradients may affect genetic variation within and among plant populations and inbreeding within populations. Populations in the upper range periphery of a species may be important source populations for range shifts to higher altitude in response to climate change. In this study we investigate patterns of population genetic variation at upper peripheral and lower more central altitudes in three common plant species of semi-dry grasslands in montane landscapes. METHODOLOGY/PRINCIPAL FINDINGS: In Briza media, Trifolium montanum and Ranunculus bulbosus genetic diversity, inbreeding and genetic relatedness of individuals within populations and genetic differentiation among populations was characterized using AFLP markers. Populations were sampled in the Swiss Alps at 1800 (upper periphery of the study organisms) and at 1200 m a.s.l. Genetic diversity was not affected by altitude and only in B. media inbreeding was greater at higher altitudes. Genetic differentiation was slightly greater among populations at higher altitudes in B. media and individuals within populations were more related to each other compared to individuals in lower altitude populations. A similar but less strong pattern of differentiation and relatedness was observed in T. montanum, while in R. bulbosus there was no effect of altitude. Estimations of population size and isolation of populations were similar, both at higher and lower altitudes. CONCLUSIONS/SIGNIFICANCE: Our results suggest that altitude does not affect genetic diversity in the grassland species under study. Genetic differentiation of populations increased only slightly at higher elevation, probably due to extensive (historic) gene flow among altitudes. Potentially pre-adapted genes might therefore spread easily across altitudes. Our study indicates that populations at the upper periphery are not genetically depauperate or isolated and thus may be important source populations for migration under climate change.