Drought-exposure history increases complementarity between plant species in response to a subsequent drought

Growing threats from extreme climatic events and biodiversity loss have raised concerns about their interactive consequences for ecosystem functioning. Evidence suggests biodiversity can buffer ecosystem functioning during such climatic events. However, whether exposure to extreme climatic events will strengthen the biodiversity-dependent buffering effects for future generations remains elusive. We assess such transgenerational effects by exposing experimental grassland communities to eight recurrent summer droughts versus ambient conditions in the field. Seed offspring of 12 species are then subjected to a subsequent drought event in the glasshouse, grown individually, in monocultures or in 2-species mixtures. Comparing productivity between mixtures and monocultures, drought-selected plants show greater between-species complementarity than ambient-selected plants when recovering from the subsequent drought, causing stronger biodiversity effects on productivity and better recovery of drought-selected mixtures after the drought. These findings suggest exposure to recurrent climatic events can improve ecosystem responses to future events through transgenerational reinforcement of species complementarity

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

Tundra species diversity and plant traits in a changing Arctic

Global air temperature is unequivocally increasing and will keep rising, more rapidly in the Arctic than in other regions. Climate warming may affect not only soil factors, e.g. temperature, moisture and nutrient availability for plants, but also vegetation. Changes in species diversity, distribution, and plant traits are expected as a consequence of direct and indirect effects of climate warming, especially in high-latitude ecosystems. Two of the main changes expected in arctic tundra are shrub expansion and loss of cryptogam diversity. Tundra vegetation shifts will result in altered feedbacks with atmosphere and permafrost through the surface energy budget and the water and carbon cycle, which might affect regional climate. Despite the high vulnerability of arctic species to climate change and the importance of tundra vegetation‒climate feedbacks, uncertainties remain in relation to species diversity and plant trait response to climate. In order to identify changes in species diversity, community composition, and plant traits that might take place under climate change in arctic tundra ecosystems, I combined observational, experimental, and dendroecological approaches. I established a set of observational plots in two contrasting habitats in northeastern Siberia, finding that species diversity and community composition were closely related to edaphic factors. These relationships were different among plant functional types, suggesting a higher vulnerability of cryptogam diversity to changes in edaphic factors and, therefore, to climate warming. Moreover, by sampling shrub individuals from experimental soil warming and fertilization plots, I found faster shrub growth with enhanced nutrient availability, a decrease in bark investment with faster growth rates, and a coordinated response of shrub traits to nutrient addition. These findings suggest a shift in growth strategy and resource acquisition towards more rapid ones with climate warming. Although shrub expansion is expected in the short term due to faster growth and denser covers, in the long term, shrubs might become more vulnerable to herbivory, pathogens, and climate extremes because of shifts in shrub resource allocation towards growth (growth‒defence trade-off). This thesis contributes, therefore, to increase our knowledge of species diversity vulnerability and plant trait shifts in a changing Arctic, which is a first step to better understand vegetation effects on the surface radiation budget in tundra ecosystems. Such an understanding is essential for reducing the uncertainties in direction and magnitude of future vegetation‒climate feedbacks

Plant trait response of tundra shrubs to permafrost thaw and nutrient addition

Plant traits reflect growth strategies and trade-offs in response to environmental conditions. Because of climate warming, plant traits might change, altering ecosystem functions and vegetation–climate interactions. Despite important feedbacks of plant trait changes in tundra ecosystems with regional climate, with a key role for shrubs, information on responses of shrub functional traits is limited. Here, we investigate the effects of experimentally increased permafrost thaw depth and (possibly thaw-associated) soil nutrient availability on plant functional traits and strategies of Arctic shrubs in northeastern Siberia. We hypothesize that shrubs will generally shift their strategy from efficient conservation to faster acquisition of resources through adaptation of leaf and stem traits in a coordinated whole-plant fashion. To test this hypothesis, we ran a 4 year permafrost thaw and nutrient fertilization experiment with a fully factorial block design and six treatment combinations – permafrost thaw (control, unheated cable, heated cable) × fertilization (no nutrient addition, nutrient addition). We measured 10 leaf and stem traits related to growth, defence and the resource economics spectrum in four shrub species (Betula nana, Salix pulchra, Ledum palustre and Vaccinium vitis-idaea), which were sampled in the experimental plots. The plant trait data were statistically analysed using linear mixed-effect models and principal component analysis (PCA). The response to increased permafrost thaw was not significant for most shrub traits. However, all shrubs responded to the fertilization treatment, despite decreased thaw depth and soil temperature in fertilized plots. Shrubs tended to grow taller but did not increase their stem density or bark thickness. We found a similar coordinated trait response for all four species at leaf and plant level; i.e. they shifted from a conservative towards a more acquisitive resource economy strategy upon fertilization. In accordance, results point towards a lower investment into defence mechanisms, and hence increased shrub vulnerability to herbivory and climate extremes. Compared to biomass and height only, detailed data involving individual plant organ traits such as leaf area and nutrient contents or stem water content can contribute to a better mechanistic understanding of feedbacks between shrub growth strategies, permafrost thaw and carbon and energy fluxes. In combination with observational data, these experimental tundra trait data allow for a more realistic representation of tundra shrubs in dynamic vegetation models and robust prediction of ecosystem functions and related climate–vegetation–permafrost feedbacks

Shrub growth rate and bark responses to soil warming and nutrient addition – A dendroecological approach in a field experiment

Tundra shrubs are slow-growing species limited by low air temperature and scarce nutrient availability. However, shrub expansion has been widely observed in the Arctic during the last decades and attributed to climate warming. Shift in shrub growth, wood structure and abundance affects the surface albedo and permafrost thawing and these changes may feedback to climate. Despite the importance of shrub–climate feedbacks, uncertainties about shrub growth sensitivity to climate remain. Here, we explored the indirect effects of climate warming on shrub growth (vertical and radial), bark thickness, and bark investment in four arctic shrub species. We combined a field experiment addressing two suggested growth drivers – thawing depth and nutrient availability – with dendroecology in a Siberian tundra ecosystem. We used heating cables to increase the thawing depth. To enhance the nutrient availability, we fertilized the surface soil layers. We found that shrub growth was mainly limited by nutrient availability, as indicated by the fertilization treatment effects on shrub growth ring widths. We also found a bark thickness decrease with the combined soil heating and nutrient addition treatment and a negative correlation between bark investment and growth rate for two of the species. These findings suggest that tundra shrubs, especially deciduous species, will grow faster and taller driven by an increasing nutrient availability in the surface soil layers. However, shrubs might become more vulnerable to pests, herbivory, and climate extremes, such as frost or drought events, due to thinner bark and lower bark investment. Using dendroecological approaches in field experiments simulating projected climate scenarios for the Arctic, and an increasing number of study species and locations will reduce uncertainties related to shrub growth sensitivity to climate and other processes driving shrub dynamics.</p

Drivers of shortwave radiation fluxes in Arctic tundra across scales

Vegetation composition and water surface area are changing in many tundra regions due to climate warming, which is twice as strong in the Arctic as compared to the global mean. Such land cover changes feed back to climate and permafrost thaw through altering the surface energy budget. We quantified the influence of vegetation type, canopy characteristics, and patchiness on the tundra shortwave radiation components. We used in situ measurements and vegetation mapping to parametrise a 3D radiative transfer model (DART) for summer conditions at the Kytalyk test site in northeast Siberia. We analysed model results assessing the most important drivers of canopy albedo, transmittance, and absorptance of photosynthetically active radiation (PAR). Tundra albedo was strongly influenced by the fractional cover of water surfaces. Albedo decreased with increasing shrub cover. However, plant area index effects on albedo were not statistically significant. Canopy transmittance and PAR absorptance (fAPAR) were almost entirely controlled by plant area index at the landscape scale. Only about one half of the total plant area index consisted of green leaves, while wood and standing dead leaves contributed equally to the other half.While spatial patterns and patch sizes of vegetation types and open water did not significantly influence the radiation budget at the landscape scale, it contributed to the large variability at the local scale. Such local variability of shortwave radiation may impact evapotranspiration and primary productivity at a range of scales. Therefore, the variation of radiation fluxes within single vegetation types potentially affects larger scale energy, water, and carbon fluxes

Plant trait response of tundra shrubs to permafrost thaw and nutrient addition

Plant traits reflect growth strategies and trade-offs in response to environmental conditions. Because of climate warming, plant traits might change, altering ecosystem functions and vegetation-climate interactions. Despite important feedbacks of plant trait changes in tundra ecosystems with regional climate, with a key role for shrubs, information on responses of shrub functional traits is limited. Here, we investigate the effects of experimentally increased permafrost thaw depth and (possibly thawassociated) soil nutrient availability on plant functional traits and strategies of Arctic shrubs in northeastern Siberia. We hypothesize that shrubs will generally shift their strategy from efficient conservation to faster acquisition of resources through adaptation of leaf and stem traits in a coordinated whole-plant fashion. To test this hypothesis, we ran a 4 year permafrost thaw and nutrient fertilization experiment with a fully factorial block design and six treatment combinations - permafrost thaw (control, unheated cable, heated cable) fertilization (no nutrient addition, nutrient addition). We measured 10 leaf and stem traits related to growth, defence and the resource economics spectrum in four shrub species (Betula nana, Salix pulchra, Ledum palustre and Vaccinium vitis-idaea), which were sampled in the experimental plots. The plant trait data were statistically analysed using linear mixed-effect models and principal component analysis (PCA). The response to increased permafrost thaw was not significant for most shrub traits. However, all shrubs responded to the fertilization treatment, despite decreased thaw depth and soil temperature in fertilized plots. Shrubs tended to grow taller but did not increase their stem density or bark thickness. We found a similar coordinated trait response for all four species at leaf and plant level; i.e. they shifted from a conservative towards a more acquisitive resource economy strategy upon fertilization. In accordance, results point towards a lower investment into defence mechanisms, and hence increased shrub vulnerability to herbivory and climate extremes. Compared to biomass and height only, detailed data involving individual plant organ traits such as leaf area and nutrient contents or stem water content can contribute to a better mechanistic understanding of feedbacks between shrub growth strategies, permafrost thaw and carbon and energy fluxes. In combination with observational data, these experimental tundra trait data allow for a more realistic representation of tundra shrubs in dynamic vegetation models and robust prediction of ecosystem functions and related climate- vegetation-permafrost feedbacks

Drivers of shortwave radiation fluxes in Arctic tundra across scales

Vegetation composition and water surface area are changing in many tundra regions due to climate warming, which is twice as strong in the Arctic as compared to the global mean. Such land cover changes feed back to climate and permafrost thaw through altering the surface energy budget. We quantified the influence of vegetation type, canopy characteristics, and patchiness on the tundra shortwave radiation components. We used in situ measurements and vegetation mapping to parametrise a 3D radiative transfer model (DART) for summer conditions at the Kytalyk test site in northeast Siberia. We analysed model results assessing the most important drivers of canopy albedo, transmittance, and absorptance of photosynthetically active radiation (PAR). Tundra albedo was strongly influenced by the fractional cover of water surfaces. Albedo decreased with increasing shrub cover. However, plant area index effects on albedo were not statistically significant. Canopy transmittance and PAR absorptance (fAPAR) were almost entirely controlled by plant area index at the landscape scale. Only about one half of the total plant area index consisted of green leaves, while wood and standing dead leaves contributed equally to the other half.While spatial patterns and patch sizes of vegetation types and open water did not significantly influence the radiation budget at the landscape scale, it contributed to the large variability at the local scale. Such local variability of shortwave radiation may impact evapotranspiration and primary productivity at a range of scales. Therefore, the variation of radiation fluxes within single vegetation types potentially affects larger scale energy, water, and carbon fluxes

Drought-exposure history increases complementarity between plant species in response to a subsequent drought

Growing threats from extreme climatic events and biodiversity loss have raised concerns about their interactive consequences for ecosystem functioning. Evidence suggests biodiversity can buffer ecosystem functioning during such climatic events. However, whether exposure to extreme climatic events will strengthen the biodiversity-dependent buffering effects for future generations remains elusive. We assess such transgenerational effects by exposing experimental grassland communities to eight recurrent summer droughts versus ambient conditions in the field. Seed offspring of 12 species are then subjected to a subsequent drought event in the glasshouse, grown individually, in monocultures or in 2-species mixtures. Comparing productivity between mixtures and monocultures, drought-selected plants show greater between-species complementarity than ambient-selected plants when recovering from the subsequent drought, causing stronger biodiversity effects on productivity and better recovery of drought-selected mixtures after the drought. These findings suggest exposure to recurrent climatic events can improve ecosystem responses to future events through transgenerational reinforcement of species complementarity