Disturbance theory for ecosystem ecologists: A primer.

Understanding what regulates ecosystem functional responses to disturbance is essential in this era of global change. However, many pioneering and still influential disturbance-related theorie proposed by ecosystem ecologists were developed prior to rapid global change, and before tools and metrics were available to test them. In light of new knowledge and conceptual advances across biological disciplines, we present four disturbance ecology concepts that are particularly relevant to ecosystem ecologists new to the field: (a) the directionality of ecosystem functional response to disturbance; (b) functional thresholds; (c) disturbance-succession interactions; and (d) diversity-functional stability relationships. We discuss how knowledge, theory, and terminology developed by several biological disciplines, when integrated, can enhance how ecosystem ecologists analyze and interpret functional responses to disturbance. For example, when interpreting thresholds and disturbance-succession interactions, ecosystem ecologists should consider concurrent biotic regime change, non-linearity, and multiple response pathways, typically the theoretical and analytical domain of population and community ecologists. Similarly, the interpretation of ecosystem functional responses to disturbance requires analytical approaches that recognize disturbance can promote, inhibit, or fundamentally change ecosystem functions. We suggest that truly integrative approaches and knowledge are essential to advancing ecosystem functional responses to disturbance.

Flower reflectance and floral traits data from Ökologisch-Botanischer Garten (OBG), Germany.

Not all colours are perceived and interpreted equally. The electromagnetic spectrum is perceived differently by the distinct visual systems of animal species, resulting in differences in each species' colour perception. Given the diverse colours found in flowering plants, it is interesting to consider the colour perception of insects and the co-evolution of flowering plants to attract pollinators. Here, we considered the differences between human visual systems and that of bees and flies-the two largest insect pollinator groups. We collected flower reflectance spectral data of 73 species across seven human-perceived colours using a spectrophotometer. Minimum of 3 different flowers were used to measure the reflectance properties of flower colours. The raw data can be used to visualize the different animals' visual systems i.e. it can be processed and translated into known photoreceptors of human, bee, and fly visual systems. Overall, our data will help to compare how different animals see flower colours in the natural world and will also highlight the importance of understanding the interspecific communication in plant-pollinator communities. Thus, our data will assist scientists in the future to recognize the floral colour evolution in angiosperms.

Flower colour and size-signals vary with altitude and resulting climate on the tropical-subtropical islands of Taiwan.

The diversity of flower colours in nature provides quantifiable evidence for how visitations by colour sensing insect pollinators can drive the evolution of angiosperm visual signalling. Recent research shows that both biotic and abiotic factors may influence flower signalling, and that harsher climate conditions may also promote salient signalling to entice scarcer pollinators to visit. In parallel, a more sophisticated appreciation of the visual task foragers face reveals that bees have a complex visual system that uses achromatic vision when moving fast, whilst colour vision requires slower, more careful inspection of targets. Spectra of 714 native flowering species across Taiwan from sea level to mountainous regions 3,300 m above sea level (a.s.l.) were measured. We modelled how the visual system of key bee pollinators process signals, including flower size. By using phylogenetically informed analyses, we observed that at lower altitudes including foothills and submontane landscapes, there is a significant relationship between colour contrast and achromatic signals. Overall, the frequency of flowers with high colour contrast increases with altitude, whilst flower size decreases. The evidence that flower colour signaling becomes increasingly salient in higher altitude conditions supports that abiotic factors influence pollinator foraging in a way that directly influences how flowering plants need to advertise.

Volcanic tephra deposition dataset based on interpolated field measurements following the 2021 Tajogaite Eruption on La Palma, Canary Islands, Spain.

In 2021, the Tajogaite Volcano erupted along the western slope of the Cumbre Vieja on the island of La Palma, Canary Islands, Spain. Volcanic tephra blanketed a substantial proportion of the island. By our estimations, approximately 23,000,000 m3 of pyroclastic ashes and more coarse-grained particles were deposited unto La Palma's land surface in addition to the lava flow. Five months following the initial eruption, we measured the depth of the new ash layer across the island. We combined this data with drone-based observations to compile a dataset comprising the point distribution of ash depth. A spatial interpolation was then performed using Inverse Distance Weighting (IDW) to estimate the ash depth across the island at a 2 m spatial resolution. The interpolation performed well, yielding a root mean squared error (RMSE) value of 0.34 and thus, the dataset offers immense reuse potential for spatial inquiries related to evolutionary traits, vegetation patterns, and vegetation response to disturbance on oceanic islands. In addition, the data can be used to test different spatial interpolation techniques in an effort to improve the accuracy achieved using IDW.

Extreme drought impacts have been underestimated in grasslands and shrublands globally.

Climate change is increasing the frequency and severity of short-term (~1 y) drought events-the most common duration of drought-globally. Yet the impact of this intensification of drought on ecosystem functioning remains poorly resolved. This is due in part to the widely disparate approaches ecologists have employed to study drought, variation in the severity and duration of drought studied, and differences among ecosystems in vegetation, edaphic and climatic attributes that can mediate drought impacts. To overcome these problems and better identify the factors that modulate drought responses, we used a coordinated distributed experiment to quantify the impact of short-term drought on grassland and shrubland ecosystems. With a standardized approach, we imposed ~a single year of drought at 100 sites on six continents. Here we show that loss of a foundational ecosystem function-aboveground net primary production (ANPP)-was 60% greater at sites that experienced statistically extreme drought (1-in-100-y event) vs. those sites where drought was nominal (historically more common) in magnitude (35% vs. 21%, respectively). This reduction in a key carbon cycle process with a single year of extreme drought greatly exceeds previously reported losses for grasslands and shrublands. Our global experiment also revealed high variability in drought response but that relative reductions in ANPP were greater in drier ecosystems and those with fewer plant species. Overall, our results demonstrate with unprecedented rigor that the global impacts of projected increases in drought severity have been significantly underestimated and that drier and less diverse sites are likely to be most vulnerable to extreme drought.

Consistent predictors of microbial community composition across spatial scales in grasslands reveal low context-dependency.

Environmental circumstances shaping soil microbial communities have been studied extensively. However, due to disparate study designs, it has been difficult to resolve whether a globally consistent set of predictors exists, or context-dependency prevails. Here, we used a network of 18 grassland sites (11 of those containing regional plant productivity gradients) to examine (i) if similar abiotic or biotic factors predict both large-scale (across sites) and regional-scale (within sites) patterns in bacterial and fungal community composition, and (ii) if microbial community composition differs consistently at two levels of regional plant productivity (low vs. high). Our results revealed that bacteria were associated with particular soil properties (such as base saturation) and both bacteria and fungi were associated with plant community composition across sites and within the majority of sites. Moreover, a discernible microbial community signal emerged, clearly distinguishing high and low-productivity soils across different grasslands independent of their location in the world. Hence, regional productivity differences may be typified by characteristic soil microbial communities across the grassland biome. These results could encourage future research aiming to predict the general effects of global changes on soil microbial community composition in grasslands and to discriminate fertile from infertile systems using generally applicable microbial indicators.

Multidimensional responses of grassland stability to eutrophication.

Eutrophication usually impacts grassland biodiversity, community composition, and biomass production, but its impact on the stability of these community aspects is unclear. One challenge is that stability has many facets that can be tightly correlated (low dimensionality) or highly disparate (high dimensionality). Using standardized experiments in 55 grassland sites from a globally distributed experiment (NutNet), we quantify the effects of nutrient addition on five facets of stability (temporal invariability, resistance during dry and wet growing seasons, recovery after dry and wet growing seasons), measured on three community aspects (aboveground biomass, community composition, and species richness). Nutrient addition reduces the temporal invariability and resistance of species richness and community composition during dry and wet growing seasons, but does not affect those of biomass. Different stability measures are largely uncorrelated under both ambient and eutrophic conditions, indicating consistently high dimensionality. Harnessing the dimensionality of ecological stability provides insights for predicting grassland responses to global environmental change.

Functional trait trade-offs define plant population stability across different biomes.

Ecological theory posits that temporal stability patterns in plant populations are associated with differences in species' ecological strategies. However, empirical evidence is lacking about which traits, or trade-offs, underlie species stability, especially across different biomes. We compiled a worldwide collection of long-term permanent vegetation records (greater than 7000 plots from 78 datasets) from a large range of habitats which we combined with existing trait databases. We tested whether the observed inter-annual variability in species abundance (coefficient of variation) was related to multiple individual traits. We found that populations with greater leaf dry matter content and seed mass were more stable over time. Despite the variability explained by these traits being low, their effect was consistent across different datasets. Other traits played a significant, albeit weaker, role in species stability, and the inclusion of multi-variate axes or phylogeny did not substantially modify nor improve predictions. These results provide empirical evidence and highlight the relevance of specific ecological trade-offs, i.e. in different resource-use and dispersal strategies, for plant populations stability across multiple biomes. Further research is, however, necessary to integrate and evaluate the role of other specific traits, often not available in databases, and intraspecific trait variability in modulating species stability.

Environmental heterogeneity modulates the effect of plant diversity on the spatial variability of grassland biomass.

Plant productivity varies due to environmental heterogeneity, and theory suggests that plant diversity can reduce this variation. While there is strong evidence of diversity effects on temporal variability of productivity, whether this mechanism extends to variability across space remains elusive. Here we determine the relationship between plant diversity and spatial variability of productivity in 83 grasslands, and quantify the effect of experimentally increased spatial heterogeneity in environmental conditions on this relationship. We found that communities with higher plant species richness (alpha and gamma diversity) have lower spatial variability of productivity as reduced abundance of some species can be compensated for by increased abundance of other species. In contrast, high species dissimilarity among local communities (beta diversity) is positively associated with spatial variability of productivity, suggesting that changes in species composition can scale up to affect productivity. Experimentally increased spatial environmental heterogeneity weakens the effect of plant alpha and gamma diversity, and reveals that beta diversity can simultaneously decrease and increase spatial variability of productivity. Our findings unveil the generality of the diversity-stability theory across space, and suggest that reduced local diversity and biotic homogenization can affect the spatial reliability of key ecosystem functions.

Stable plant community biomass production despite species richness collapse under simulated extreme climate in the European Alps.

Direct observation of biodiversity loss in response to abrupt climate change can resolve fundamental questions about temporal community dynamics and clarify the controversial debate of biodiversity loss impacts on ecosystem functioning. We tracked local plant species loss and the corresponding change of aboveground biomass of native and non-native species by actively pushing mountain grassland ecosystems beyond their ecological thresholds in a five-year, multisite translocation experiment across the European Alps. Our results show that species loss (ranging from a 73 % to 94 % reduction in species richness) caused by simulated climate extremes (strong warming interacting with drought) did not decrease community biomass. Even without non-native species colonization, the community biomass of native species remained stable during native species richness collapse. Switching our research focus from local extinction in the face of climate change towards the beneficial impacts of persisting native species (in addition to novel plant-plant interactions) might yield insights on transformative opportunities for boosting climate resilience.

The potential bias of nitrogen deposition effects on primary productivity and biodiversity.

Atmospheric nitrogen (N) deposition is composed of both inorganic nitrogen (IN) and organic nitrogen (ON), and these sources of N may exhibit different impacts on ecosystems. However, our understanding of the impacts of N deposition is largely based on experimental gradients of INs or more rarely ONs. Thus, the effects of N deposition on ecosystem productivity and biodiversity may be biased. We explored the differential impacts of N addition with different IN:ON ratios (0:10, 3:7, 5:5, 7:3, and 10:0) on aboveground net primary productivity (ANPP) of plant community and plant diversity in a typical temperate grassland with a long-term N addition experiment. Soil pH, litter biomass, soil IN concentration, and light penetration were measured to examine the potential mechanisms underlying species loss with N addition. Our results showed that N addition significantly increased plant community ANPP by 68.33%-105.50% and reduced species richness by 16.20%-37.99%. The IN:ON ratios showed no significant effects on plant community ANPP. However, IN-induced species richness loss was about 2.34 times of ON-induced richness loss. Soil pH was positively related to species richness, and they exhibited very similar response patterns to IN:ON ratios. It implies that soil acidification accounts for the different magnitudes of species loss with IN and ON additions. Overall, our study suggests that it might be reasonable to evaluate the effects of N deposition on plant community ANPP with either IN or ON addition. However, the evaluation of N deposition on biodiversity might be overestimated if only IN is added or underestimated if only ON is added.

Linking changes in species composition and biomass in a globally distributed grassland experiment.

Global change drivers, such as anthropogenic nutrient inputs, are increasing globally. Nutrient deposition simultaneously alters plant biodiversity, species composition and ecosystem processes like aboveground biomass production. These changes are underpinned by species extinction, colonisation and shifting relative abundance. Here, we use the Price equation to quantify and link the contributions of species that are lost, gained or that persist to change in aboveground biomass in 59 experimental grassland sites. Under ambient (control) conditions, compositional and biomass turnover was high, and losses (i.e. local extinctions) were balanced by gains (i.e. colonisation). Under fertilisation, the decline in species richness resulted from increased species loss and decreases in species gained. Biomass increase under fertilisation resulted mostly from species that persist and to a lesser extent from species gained. Drivers of ecological change can interact relatively independently with diversity, composition and ecosystem processes and functions such as aboveground biomass due to the individual contributions of species lost, gained or persisting.

Field experiments underestimate aboveground biomass response to drought.

Researchers use both experiments and observations to study the impacts of climate change on ecosystems, but results from these contrasting approaches have not been systematically compared for droughts. Using a meta-analysis and accounting for potential confounding factors, we demonstrate that aboveground biomass responded only about half as much to experimentally imposed drought events as to natural droughts. Our findings indicate that experimental results may underestimate climate change impacts and highlight the need to integrate results across approaches.

Interspecific trait variability and local soil conditions modulate grassland model community responses to climate.

Medium-to-high elevation grasslands provide critical services in agriculture and ecosystem stabilization, through high biodiversity and providing food for wildlife. However, these ecosystems face elevated risks of disruption due to predicted soil and climate changes. Separating the effects of soil and climate, however, is difficult in situ, with previous experiments focusing largely on monocultures instead of natural grassland communities. We experimentally exposed model grassland communities, comprised of three species grown on either local or reference soil, to varied climatic environments along an elevational gradient in the European Alps, measuring the effects on species and community traits. Although species-specific biomass varied across soil and climate, species' proportional contributions to community-level biomass production remained consistent. Where species experienced low survivorship, species-level biomass production was maintained through increased productivity of surviving individuals; however, maximum species-level biomass was obtained under high survivorship. Species responded directionally to climatic variation, spatially separating differentially by plant traits (including height, reproduction, biomass, survival, leaf dry weight, and leaf area) consistently across all climates. Local soil variation drove stochastic trait responses across all species, with high levels of interactions occurring between site and species. This soil variability obscured climate-driven responses: we recorded no directional trait responses for soil-corrected traits like observed for climate-corrected traits. Our species-based approach contributes to our understanding of grassland community stabilization and suggests that these communities show some stability under climatic variation.

Determinants of community compositional change are equally affected by global change.

Global change is impacting plant community composition, but the mechanisms underlying these changes are unclear. Using a dataset of 58 global change experiments, we tested the five fundamental mechanisms of community change: changes in evenness and richness, reordering, species gains and losses. We found 71% of communities were impacted by global change treatments, and 88% of communities that were exposed to two or more global change drivers were impacted. Further, all mechanisms of change were equally likely to be affected by global change treatments-species losses and changes in richness were just as common as species gains and reordering. We also found no evidence of a progression of community changes, for example, reordering and changes in evenness did not precede species gains and losses. We demonstrate that all processes underlying plant community composition changes are equally affected by treatments and often occur simultaneously, necessitating a wholistic approach to quantifying community changes.

Biotic homogenization destabilizes ecosystem functioning by decreasing spatial asynchrony.

Our planet is facing significant changes of biodiversity across spatial scales. Although the negative effects of local biodiversity (α diversity) loss on ecosystem stability are well documented, the consequences of biodiversity changes at larger spatial scales, in particular biotic homogenization, that is, reduced species turnover across space (ß diversity), remain poorly known. Using data from 39 grassland biodiversity experiments, we examine the effects of ß diversity on the stability of simulated landscapes while controlling for potentially confounding biotic and abiotic factors. Our results show that higher ß diversity generates more asynchronous dynamics among local communities and thereby contributes to the stability of ecosystem productivity at larger spatial scales. We further quantify the relative contributions of α and ß diversity to ecosystem stability and find a relatively stronger effect of α diversity, possibly due to the limited spatial scale of our experiments. The stabilizing effects of both α and ß diversity lead to a positive diversity-stability relationship at the landscape scale. Our findings demonstrate the destabilizing effect of biotic homogenization and suggest that biodiversity should be conserved at multiple spatial scales to maintain the stability of ecosystem functions and services.

Fertilized graminoids intensify negative drought effects on grassland productivity.

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.

Root-Associated Mycobiomes of Common Temperate Plants (Calluna vulgaris and Holcus lanatus) Are Strongly Affected by Winter Climate Conditions.

Winter temperatures are projected to increase in Central Europe. Subsequently, snow cover will decrease, leading to increased soil temperature variability, with potentially different consequences for soil frost depending on e.g. altitude. Here, we experimentally evaluated the effects of increased winter soil temperature variability on the root associated mycobiome of two plant species (Calluna vulgaris and Holcus lanatus) at two sites in Germany; a colder and wetter upland site with high snow accumulation and a warmer and drier lowland site, with low snow accumulation. Mesocosm monocultures were set-up in spring 2010 at both sites (with soil and plants originating from the lowland site). In the following winter, an experimental warming pulse treatment was initiated by overhead infrared heaters and warming wires at the soil surface for half of the mesocosms at both sites. At the lowland site, the warming treatment resulted in a reduced number of days with soil frost as well as increased the average daily temperature amplitude. Contrary, the treatment caused no changes in these parameters at the upland site, which was in general a much more frost affected site. Soil and plant roots were sampled before and after the following growing season (spring and autumn 2011). High-throughput sequencing was used for profiling of the root-associated fungal (ITS marker) community (mycobiome). Site was found to have a profound effect on the composition of the mycobiome, which at the upland site was dominated by fast growing saprotrophs (Mortierellomycota), and at the lowland site by plant species-specific symbionts (e.g. Rhizoscyphus ericae and Microdochium bolleyi for C. vulgaris and H. lanatus respectively). The transplantation to the colder upland site and the temperature treatment at the warmer lowland site had comparable consequences for the mycobiome, implying that winter climate change resulting in higher temperature variability has large consequences for mycobiome structures regardless of absolute temperature of a given site.