Mediterranean cork oak savannas, which are found only in southwestern Europe and northwestern Africa, are ecosystems of high socioeconomic and conservation value. Characterized by sparse tree cover ...and a diversity of understory vegetation -– ranging from shrub formations to grasslands -– that support high levels of biodiversity, these ecosystems require active management and use by humans to ensure their continued existence. The most important product of these savannas is cork, a non-timber forest product that is periodically harvested without requiring tree felling. Market devaluation of, and lower demand for, cork are causing a decline in management, or even abandonment, of southwestern Europe's cork oak savannas. Subsequent shrub encroachment into the savanna's grassland components reduces biodiversity and degrades the services provided by these ecosystems. In contrast, poverty-driven overuse is degrading cork oak savannas in northwestern Africa. "“Payment for ecosystem services"” schemes, such as Forest Stewardship Council (FSC) certification or Reducing Emissions from Deforestation and Degradation and enhancement of carbon stocks (REDD++) programs, could produce novel economic incentives to promote sustainable use and conservation of Mediterranean cork oak savanna ecosystems in both Europe and Africa.
Plant damage by invertebrate herbivores and pathogens influences the dynamics of grassland ecosystems, but anthropogenic changes in nitrogen and phosphorus availability can modify these ...relationships.
Using a globally distributed experiment, we describe leaf damage on 153 plant taxa from 27 grasslands worldwide, under ambient conditions and with experimentally elevated nitrogen and phosphorus.
Invertebrate damage significantly increased with nitrogen addition, especially in grasses and non‐leguminous forbs. Pathogen damage increased with nitrogen in grasses and legumes but not forbs. Effects of phosphorus were generally weaker. Damage was higher in grasslands with more precipitation, but climatic conditions did not change effects of nutrients on leaf damage. On average, invertebrate damage was relatively higher on legumes and pathogen damage was relatively higher on grasses. Community‐weighted mean damage reflected these functional group patterns, with no effects of N on community‐weighted pathogen damage (due to opposing responses of grasses and forbs) but stronger effects of N on community‐weighted invertebrate damage (due to consistent responses of grasses and forbs).
Synthesis. As human‐induced inputs of nitrogen and phosphorus continue to increase, understanding their impacts on invertebrate and pathogen damage becomes increasingly important. Our results demonstrate that eutrophication frequently increases plant damage and that damage increases with precipitation across a wide array of grasslands. Invertebrate and pathogen damage in grasslands is likely to increase in the future, with potential consequences for plant, invertebrate and pathogen communities, as well as the transfer of energy and nutrients across trophic levels.
As human‐induced inputs of nitrogen and phosphorus continue to increase, understanding their impacts on invertebrate and pathogen damage becomes increasingly important. Our results demonstrate that eutrophication frequently increases plant damage and that damage increases with precipitation across a wide array of grasslands. Invertebrate and pathogen damage in grasslands is likely to increase in the future, with potential consequences for plant, invertebrate and pathogen communities, as well as the transfer of energy and nutrients across trophic levels.
The ‘two-water-worlds’ hypothesis is based on stable isotope differences in stream, soil and xylem waters in dual isotope space. It postulates no connectivity between bound and mobile soil waters, ...and preferential plant water uptake of bound soil water sources. We tested the pool-weighted impact of isotopically distinct water pools for hydrological cycling, the influence of species-specific water use and the degree of ecohydrological separation.
We combined stable isotope analysis (δ18O and δ²H) of ecosystem water pools of precipitation, groundwater, soil and xylem water of two distinct species (Quercus suber, Cistus ladanifer) with observations of soil water contents and sap flow.
Shallow soil water was evaporatively enriched during dry-down periods, but enrichment faded strongly with depth and upon precipitation events. Despite clearly distinct water sources and water-use strategies, both species displayed a highly opportunistic pattern of root water uptake.
Here we offer an alternative explanation, showing that the isotopic differences between soil and plant water vs groundwater can be fully explained by spatio-temporal dynamics. Pool weighting the contribution of evaporatively enriched soil water reveals only minor annual impacts of these sources to ecosystem water cycling (c. 11% of bulk soil water and c. 14% of transpired water).
Understanding of plant hydraulic strategies (i.e. the degree of iso‐/anisohydricity) is crucial to predict the response of plants to changing environmental conditions such as climate‐change induced ...extreme drought. Several abiotic factors, including evaporative demand, have been shown to seasonally modify the isohydricity of plants. However, the impact of biotic factors such as plant–plant interactions on hydraulic strategies has seldom been explored.
Here, we investigated adaptations and changes in hydraulic strategies of two woody species in response to seasonal abiotic conditions, experimental drought and plant invasion in a Mediterranean cork oak (Quercus suber) ecosystem with a combined shrub invasion (Cistus ladanifer) and rain exclusion experiment.
From the dry to wet season, Q. suber shifted from a partial isohydric to an anisohydric behaviour while C. ladanifer shifted from strict anisohydric to partial isohydric. During drought, water competition by plant invasion significantly modified the hydraulic strategy of invaded Q. suber, which was accompanied by lower pre‐dawn leaf water potentials, sap flow density, leaf area index and trunk increment rates.
This altered isohydricity of invaded Q. suber trees was most likely caused by interspecific competition for water resources by water spending C. ladanifer shrubs. Both species do have the highest proportion of fine roots in the topsoil and thus, an additional water consumer, such as C. ladanifer can lead to more stressful conditions for Q. suber during times of water scarcity. Further underlying mechanisms of the altered isohydricity of Q. suber, such as potential allelopathic effects of C. ladanifer exudates on root growth of Q. suber, have to be investigated in the future.
In conclusion, we demonstrate that the degree of isohydricity of two woody Mediterranean plant species is dynamically determined by the interplay of species‐specific hydraulic traits and their abiotic and biotic environment.
Read the free Plain Language Summary for this article on the Journal blog.
Read the free Plain Language Summary for this article on the Journal blog.
Droughts can strongly affect grassland productivity and biodiversity, but responses differ widely. Nutrient availability may be a critical factor explaining this variation, but is often ignored in ...analyses of drought responses. Here, we used a standardized nutrient addition experiment covering 10 European grasslands to test if full‐factorial nitrogen, phosphorus, and potassium addition affected plant community responses to inter‐annual variation in drought stress and to the extreme summer drought of 2018 in Europe. We found that nutrient addition amplified detrimental drought effects on community aboveground biomass production. Drought effects also differed between functional groups, with a negative effect on graminoid but not forb biomass production. Our results imply that eutrophication in grasslands, which promotes dominance of drought‐sensitive graminoids over forbs, amplifies detrimental drought effects. In terms of climate change adaptation, agricultural management would benefit from taking into account differential drought impacts on fertilized versus unfertilized grasslands, which differ in ecosystem services they provide to society.
Drought impacts on grasslands vary widely. Nutrient availability may be a critical factor explaining this variation. We used a standardized nutrient addition experiment covering 10 European grasslands to test if nitrogen, phosphorus, and potassium addition affected responses to inter‐annual variation in drought stress and to the extreme summer drought of 2018. We found that nutrient addition amplified detrimental drought effects on biomass production. Drought effects differed between functional groups, with a negative effect on graminoids but not forbs. For climate change adaptation, agricultural management would benefit from taking into account differential drought impacts on fertilized versus unfertilized grasslands, which differ in ecosystem service provisioning.
Abstract
Little is currently known about how climate modulates the relationship between plant diversity and soil organic carbon and the mechanisms involved. Yet, this knowledge is of crucial ...importance in times of climate change and biodiversity loss. Here, we show that plant diversity is positively correlated with soil carbon content and soil carbon-to-nitrogen ratio across 84 grasslands on six continents that span wide climate gradients. The relationships between plant diversity and soil carbon as well as plant diversity and soil organic matter quality (carbon-to-nitrogen ratio) are particularly strong in warm and arid climates. While plant biomass is positively correlated with soil carbon, plant biomass is not significantly correlated with plant diversity. Our results indicate that plant diversity influences soil carbon storage not via the quantity of organic matter (plant biomass) inputs to soil, but through the quality of organic matter. The study implies that ecosystem management that restores plant diversity likely enhances soil carbon sequestration, particularly in warm and arid climates.
Climate change is increasing the frequency of droughts and the risk of severe wildfires, which can interact with shrub encroachment and browsing by wild ungulates. Wild ungulate populations are ...expanding due, among other factors, to favorable habitat changes resulting from land abandonment or land‐use changes. Understanding how ungulate browsing interacts with drought to affect woody plant mortality, plant flammability, and fire hazard is especially relevant in the context of climate change and increasing frequency of wildfires. The aim of this study is to explore the combined effects of cumulative drought, shrub encroachment, and ungulate browsing on the fire hazard of Mediterranean oak woodlands in Portugal. In a long‐term (18 years) ungulate fencing exclusion experiment that simulated land abandonment and management neglect, we investigated the population dynamics of the native shrub Cistus ladanifer, which naturally dominates the understory of woodlands and is browsed by ungulates, comparing areas with (no fencing) and without (fencing) wild ungulate browsing. We also modeled fire behavior in browsed and unbrowsed plots considering drought and nondrought scenarios. Specifically, we estimated C. ladanifer population density, biomass, and fuel load characteristics, which were used to model fire behavior in drought and nondrought scenarios. Overall, drought increased the proportion of dead C. ladanifer shrub individuals, which was higher in the browsed plots. Drought decreased the ratio of live to dead shrub plant material, increased total fuel loading, shrub stand flammability, and the modeled fire parameters, that is, rate of surface fire spread, fireline intensity, and flame length. However, total fuel load and fire hazard were lower in browsed than unbrowsed plots, both in drought and nondrought scenarios. Browsing also decreased the population density of living shrubs, halting shrub encroachment. Our study provides long‐term experimental evidence showing the role of wild ungulates in mitigating drought effects on fire hazard in shrub‐encroached Mediterranean oak woodlands. Our results also emphasize that the long‐term effects of land abandonment can interact with climate change drivers, affecting wildfire hazard. This is particularly relevant given the increasing incidence of land abandonment.
Extreme drought events and plant invasions are major drivers of global change that can critically affect ecosystem functioning and alter ecosystem-atmosphere exchange. Invaders are expanding ...worldwide and extreme drought events are projected to increase in frequency and intensity. However, very little is known on how these drivers may interact to affect the functioning and resilience of ecosystems to extreme events. Using a manipulative shrub removal experiment and the co-occurrence of an extreme drought event (2011/2012) in a Mediterranean woodland, we show that native shrub invasion and extreme drought synergistically reduced ecosystem transpiration and the resilience of key-stone oak tree species. Ecosystem transpiration was dominated by the water use of the invasive shrub Cistus ladanifer, which further increased after the extreme drought event. Meanwhile, the transpiration of key-stone tree species decreased, indicating a competitive advantage in favour of the invader. Our results suggest that in Mediterranean-type climates the invasion of water spending species and projected recurrent extreme drought events may synergistically cause critical drought tolerance thresholds of key-stone tree species to be surpassed, corroborating observed higher tree mortality in the invaded ecosystems. Ultimately, this may shift seasonally water limited ecosystems into less desirable alternative states dominated by water spending invasive shrubs.
Phloem osmolality and its components are involved in basic cell metabolism, cell growth, and in various physiological processes including the ability of living cells to withstand drought and frost. ...Osmolality and sugar composition responses to environmental stresses have been extensively studied for leaves, but less for the secondary phloem of plant stems and branches. Leaf osmotic concentration and the share of pinitol and raffinose among soluble sugars increase with increasing drought or cold stress, and osmotic concentration is adjusted with osmoregulation. We hypothesize that similar responses occur in the secondary phloem of branches. We collected living bark samples from branches of adult Pinus sylvestris, Picea abies, Betula pendula and Populus tremula trees across Europe, from boreal Northern Finland to Mediterranean Portugal. In all studied species, the observed variation in phloem osmolality was mainly driven by variation in phloem water content, while tissue solute content was rather constant across regions. Osmoregulation, in which osmolality is controlled by variable tissue solute content, was stronger for Betula and Populus in comparison to the evergreen conifers. Osmolality was lowest in mid-latitude region, and from there increased by 37% toward northern Europe and 38% toward southern Europe due to low phloem water content in these regions. The ratio of raffinose to all soluble sugars was negligible at mid-latitudes and increased toward north and south, reflecting its role in cold and drought tolerance. For pinitol, another sugar known for contributing to stress tolerance, no such latitudinal pattern was observed. The proportion of sucrose was remarkably low and that of hexoses (i.e., glucose and fructose) high at mid-latitudes. The ratio of starch to all non-structural carbohydrates increased toward the northern latitudes in agreement with the build-up of osmotically inactive C reservoir that can be converted into soluble sugars during winter acclimation in these cold regions. Present results for the secondary phloem of trees suggest that adjustment with tissue water content plays an important role in osmolality dynamics. Furthermore, trees acclimated to dry and cold climate showed high phloem osmolality and raffinose proportion.
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