Arthropod decline in grasslands and forests is associated with landscape-level drivers.

Recent reports of local extinctions of arthropod species1, and of massive declines in arthropod biomass2, point to land-use intensification as a major driver of decreasing biodiversity. However, to our knowledge, there are no multisite time series of arthropod occurrences across gradients of land-use intensity with which to confirm causal relationships. Moreover, it remains unclear which land-use types and arthropod groups are affected, and whether the observed declines in biomass and diversity are linked to one another. Here we analyse data from more than 1 million individual arthropods (about 2,700 species), from standardized inventories taken between 2008 and 2017 at 150 grassland and 140 forest sites in 3 regions of Germany. Overall gamma diversity in grasslands and forests decreased over time, indicating loss of species across sites and regions. In annually sampled grasslands, biomass, abundance and number of species declined by 67%, 78% and 34%, respectively. The decline was consistent across trophic levels and mainly affected rare species; its magnitude was independent of local land-use intensity. However, sites embedded in landscapes with a higher cover of agricultural land showed a stronger temporal decline. In 30 forest sites with annual inventories, biomass and species number-but not abundance-decreased by 41% and 36%, respectively. This was supported by analyses of all forest sites sampled in three-year intervals. The decline affected rare and abundant species, and trends differed across trophic levels. Our results show that there are widespread declines in arthropod biomass, abundance and the number of species across trophic levels. Arthropod declines in forests demonstrate that loss is not restricted to open habitats. Our results suggest that major drivers of arthropod decline act at larger spatial scales, and are (at least for grasslands) associated with agriculture at the landscape level. This implies that policies need to address the landscape scale to mitigate the negative effects of land-use practices.

Climate-land-use interactions shape tropical mountain biodiversity and ecosystem functions.

Agriculture and the exploitation of natural resources have transformed tropical mountain ecosystems across the world, and the consequences of these transformations for biodiversity and ecosystem functioning are largely unknown1-3. Conclusions that are derived from studies in non-mountainous areas are not suitable for predicting the effects of land-use changes on tropical mountains because the climatic environment rapidly changes with elevation, which may mitigate or amplify the effects of land use4,5. It is of key importance to understand how the interplay of climate and land use constrains biodiversity and ecosystem functions to determine the consequences of global change for mountain ecosystems. Here we show that the interacting effects of climate and land use reshape elevational trends in biodiversity and ecosystem functions on Africa's largest mountain, Mount Kilimanjaro (Tanzania). We find that increasing land-use intensity causes larger losses of plant and animal species richness in the arid lowlands than in humid submontane and montane zones. Increases in land-use intensity are associated with significant changes in the composition of plant, animal and microorganism communities; stronger modifications of plant and animal communities occur in arid and humid ecosystems, respectively. Temperature, precipitation and land use jointly modulate soil properties, nutrient turnover, greenhouse gas emissions, plant biomass and productivity, as well as animal interactions. Our data suggest that the response of ecosystem functions to land-use intensity depends strongly on climate; more-severe changes in ecosystem functioning occur in the arid lowlands and the cold montane zone. Interactions between climate and land use explained-on average-54% of the variation in species richness, species composition and ecosystem functions, whereas only 30% of variation was related to single drivers. Our study reveals that climate can modulate the effects of land use on biodiversity and ecosystem functioning, and points to a lowered resistance of ecosystems in climatically challenging environments to ongoing land-use changes in tropical mountainous regions.

Nature 4.0: A networked sensor system for integrated biodiversity monitoring.

Ecosystem functions and services are severely threatened by unprecedented global loss in biodiversity. To counteract these trends, it is essential to develop systems to monitor changes in biodiversity for planning, evaluating, and implementing conservation and mitigation actions. However, the implementation of monitoring systems suffers from a trade-off between grain (i.e., the level of detail), extent (i.e., the number of study sites), and temporal repetition. Here, we present an applied and realized networked sensor system for integrated biodiversity monitoring in the Nature 4.0 project as a solution to these challenges, which considers plants and animals not only as targets of investigation, but also as parts of the modular sensor network by carrying sensors. Our networked sensor system consists of three main closely interlinked components with a modular structure: sensors, data transmission, and data storage, which are integrated into pipelines for automated biodiversity monitoring. We present our own real-world examples of applications, share our experiences in operating them, and provide our collected open data. Our flexible, low-cost, and open-source solutions can be applied for monitoring individual and multiple terrestrial plants and animals as well as their interactions. Ultimately, our system can also be applied to area-wide ecosystem mapping tasks, thereby providing an exemplary cost-efficient and powerful solution for biodiversity monitoring. Building upon our experiences in the Nature 4.0 project, we identified ten key challenges that need to be addressed to better understand and counteract the ongoing loss of biodiversity using networked sensor systems. To tackle these challenges, interdisciplinary collaboration, additional research, and practical solutions are necessary to enhance the capability and applicability of networked sensor systems for researchers and practitioners, ultimately further helping to ensure the sustainable management of ecosystems and the provision of ecosystem services.

Beyond body size: consistent decrease of traits within orthopteran assemblages with elevation.

Morphological traits provide the interface between species and their environment. For example, body size affects the fitness of individuals in various ways. Yet especially for ectotherms, the applicability of general rules of interspecific clines of body size and even more so of other morphological traits is still under debate. Here we tested relationships between elevation (as a proxy for temperature) and productivity with four ecologically relevant morphological traits of orthopteran assemblages that are related to fecundity (body size), dispersal (wing length), jumping ability (hind femur length), and predator detection (eye size). We measured traits of 160 orthopteran species that were sampled along an extensive environmental gradient at Mt. Kilimanjaro (Tanzania), spanning elevations from 790 to 4,410 m above sea level (a.s.l.) with different levels of plant productivity. For traits other than body size, we calculated the residuals from a regression on body length to estimate the variation of traits irrespective of body size. Bayesian analyses revealed that mean body size of assemblages, as well as the means of relative wing length, hind femur length, and eye size, decreased with increasing elevation. Body size and relative eye size also decreased with increasing productivity. Both phylogenetic relationships, as well as species-specific adaptations, contributed to these patterns. Our results suggest that orthopteran assemblages had higher fecundity and better dispersal and escape abilities, as well as better predator detection at higher temperatures (low elevations) than at low temperatures (high elevations). Large body sizes might be advantageous in habitats with low productivity because of a reduced risk of starvation. Likewise, large eye size might be advantageous because of the ability to detect predators in habitats with low vegetation cover, where hiding possibilities are scarce. Our study highlights that changes in temperature and productivity not only lead to interspecific changes in body size but are also related to independent changes of other morphological traits that influence the ecological fit of organisms in their environment.

A glimpse at short-term controls of evapotranspiration along the southern slopes of Kilimanjaro.

Future climate characteristics of the southern Kilimanjaro region, Tanzania, are mainly determined by local land-use and global climate change. Reinforcing increasing dryness throughout the twentieth century, ongoing land transformation processes emphasize the need for a proper understanding of the regional-scale water budget and possible implications on related ecosystem functioning and services. Here, we present an analysis of scintillometer-based evapotranspiration (ET) covering seven distinct habitat types across a massive climate gradient from the colline savanna woodlands to the upper-mountain Helichrysum zone (940 to 3960 m.a.s.l.). Random forest-based mean variable importance indicates an outstanding significance of net radiation (R net) on the observed ET across all elevation levels. Accordingly, topography and frequent cloud/fog events have a dampening effect at high elevations, whereas no such constraints affect the energy and moisture-rich submontane coffee/grassland level. By contrast, long-term moisture availability is likely to impose restrictions upon evapotranspirative net water loss in savanna, which particularly applies to the pronounced dry season. At plot scale, ET can thereby be approximated reasonably using R net, soil heat flux, and to a lesser degree, vapor pressure deficit and rainfall as predictor variables (R 2 0.59 to 1.00). While multivariate regression based on pooled meteorological data from all plots proves itself useful for predicting hourly ET rates across a broader range of ecosystems (R 2 = 0.71), additional gains in explained variance can be achieved when vegetation characteristics as seen from the NDVI are considered (R 2 = 0.87). To sum up, our results indicate that valuable insights into land cover-specific ET dynamics, including underlying drivers, may be derived even from explicitly short-term measurements in an ecologically highly diverse landscape.

Quantifying and mapping ecosystem services supplies and demands: a review of remote sensing applications.

Ecosystems provide services necessary for the livelihoods and well-being of people. Quantifying and mapping supplies and demands of ecosystem services is essential for continuous monitoring of such services to support decision-making. Area-wide and spatially explicit mapping of ecosystem services based on extensive ground surveys is restricted to local scales and limited due to high costs. In contrast, remote sensing provides reliable area-wide data for quantifying and mapping ecosystem services at comparatively low costs, and with the option of fast, frequent, and continuous observations for monitoring. In this paper, we review relevant remote sensing systems, sensor types, and methods applicable in quantifying selected provisioning and regulatory services. Furthermore, opportunities, challenges, and future prospects in using remote sensing for supporting ecosystem services' quantification and mapping are discussed.

Animal-Mediated Ecosystem Process Rates in Forests and Grasslands are Affected by Climatic Conditions and Land-Use Intensity.

Decomposition, vegetation regeneration, and biological control are essential ecosystem functions, and animals are involved in the underlying processes, such as dung removal, seed removal, herbivory, and predation. Despite evidence for declines of animal diversity and abundance due to climate change and land-use intensification, we poorly understand how animal-mediated processes respond to these global change drivers. We experimentally measured rates of four ecosystem processes in 134 grassland and 149 forest plots in Germany and tested their response to climatic conditions and land-use intensity, that is, grazing, mowing, and fertilization in grasslands and the proportion of harvested wood, non-natural trees, and deadwood origin in forests. For both climate and land use, we distinguished between short-term effects during the survey period and medium-term effects during the preceding years. Forests had significantly higher process rates than grasslands. In grasslands, the climatic effects on the process rates were similar or stronger than land-use effects, except for predation; land-use intensity negatively affected several process rates. In forests, the land-use effects were more pronounced than the climatic effects on all processes except for predation. The proportion of non-natural trees had the greatest impact on the process rates in forests. The proportion of harvested wood had negative effects, whereas the proportion of anthropogenic deadwood had positive effects on some processes. The effects of climatic conditions and land-use intensity on process rates mirror climatic and habitat effects on animal abundance, activity, and resource quality. Our study demonstrates that land-use changes and interventions affecting climatic conditions will have substantial impacts on animal-mediated ecosystem processes.

Species richness is more important for ecosystem functioning than species turnover along an elevational gradient.

Many experiments have shown that biodiversity enhances ecosystem functioning. However, we have little understanding of how environmental heterogeneity shapes the effect of diversity on ecosystem functioning and to what extent this diversity effect is mediated by variation in species richness or species turnover. This knowledge is crucial to scaling up the results of experiments from local to regional scales. Here we quantify the diversity effect and its components-that is, the contributions of variation in species richness and species turnover-for 22 ecosystem functions of microorganisms, plants and animals across 13 major ecosystem types on Mt Kilimanjaro, Tanzania. Environmental heterogeneity across ecosystem types on average increased the diversity effect from explaining 49% to 72% of the variation in ecosystem functions. In contrast to our expectation, the diversity effect was more strongly mediated by variation in species richness than by species turnover. Our findings reveal that environmental heterogeneity strengthens the relationship between biodiversity and ecosystem functioning and that species richness is a stronger driver of ecosystem functioning than species turnover. Based on a broad range of taxa and ecosystem functions in a non-experimental system, these results are in line with predictions from biodiversity experiments and emphasize that conserving biodiversity is essential for maintaining ecosystem functioning.

Model parameterization to simulate and compare the PAR absorption potential of two competing plant species.

Mountain pastures dominated by the pasture grass Setaria sphacelata in the Andes of southern Ecuador are heavily infested by southern bracken (Pteridium arachnoideum), a major problem for pasture management. Field observations suggest that bracken might outcompete the grass due to its competitive strength with regard to the absorption of photosynthetically active radiation (PAR). To understand the PAR absorption potential of both species, the aims of the current paper are to (1) parameterize a radiation scheme of a two-big-leaf model by deriving structural (LAI, leaf angle parameter) and optical (leaf albedo, transmittance) plant traits for average individuals from field surveys, (2) to initialize the properly parameterized radiation scheme with realistic global irradiation conditions of the Rio San Francisco Valley in the Andes of southern Ecuador, and (3) to compare the PAR absorption capabilities of both species under typical local weather conditions. Field data show that bracken reveals a slightly higher average leaf area index (LAI) and more horizontally oriented leaves in comparison to Setaria. Spectrometer measurements reveal that bracken and Setaria are characterized by a similar average leaf absorptance. Simulations with the average diurnal course of incoming solar radiation (1998-2005) and the mean leaf-sun geometry reveal that PAR absorption is fairly equal for both species. However, the comparison of typical clear and overcast days show that two parameters, (1) the relation of incoming diffuse and direct irradiance, and (2) the leaf-sun geometry play a major role for PAR absorption in the two-big-leaf approach: Under cloudy sky conditions (mainly diffuse irradiance), PAR absorption is slightly higher for Setaria while under clear sky conditions (mainly direct irradiance), the average bracken individual is characterized by a higher PAR absorption potential. (approximately 74 MJ m(-2) year(-1)). The latter situation which occurs if the maximum daily irradiance exceeds 615 W m(-2) is mainly due to the nearly orthogonal incidence of the direct solar beam onto the horizontally oriented frond area which implies a high amount of direct PAR absorption during the noon maximum of direct irradiance. Such situations of solar irradiance favoring a higher PAR absorptance of bracken occur in approximately 36% of the observation period (1998-2005). By considering the annual course of PAR irradiance in the San Francisco Valley, the clear advantage of bracken on clear days (36% of all days) is completely compensated by the slight but more frequent advantage of Setaria under overcast conditions (64% of all days). This means that neither bracken nor Setaria show a distinct advantage in PAR absorption capability under the current climatic conditions of the study area.

Author Correction: Heterogeneity-diversity relationships differ between and within trophic levels in temperate forests.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Heterogeneity-diversity relationships differ between and within trophic levels in temperate forests.

The habitat heterogeneity hypothesis predicts that biodiversity increases with increasing habitat heterogeneity due to greater niche dimensionality. However, recent studies have reported that richness can decrease with high heterogeneity due to stochastic extinctions, creating trade-offs between area and heterogeneity. This suggests that greater complexity in heterogeneity-diversity relationships (HDRs) may exist, with potential for group-specific responses to different facets of heterogeneity that may only be partitioned out by a simultaneous test of HDRs of several species groups and several facets of heterogeneity. Here, we systematically decompose habitat heterogeneity into six major facets on ~500 temperate forest plots across Germany and quantify biodiversity of 12 different species groups, including bats, birds, arthropods, fungi, lichens and plants, representing 2,600 species. Heterogeneity in horizontal and vertical forest structure underpinned most HDRs, followed by plant diversity, deadwood and topographic heterogeneity, but the relative importance varied even within the same trophic level. Among substantial HDRs, 53% increased monotonically, consistent with the classical habitat heterogeneity hypothesis but 21% were hump-shaped, 25% had a monotonically decreasing slope and 1% showed no clear pattern. Overall, we found no evidence of a single generalizable mechanism determining HDR patterns.

Middle Stone Age foragers resided in high elevations of the glaciated Bale Mountains, Ethiopia.

Studies of early human settlement in alpine environments provide insights into human physiological, genetic, and cultural adaptation potentials. Although Late and even Middle Pleistocene human presence has been recently documented on the Tibetan Plateau, little is known regarding the nature and context of early persistent human settlement in high elevations. Here, we report the earliest evidence of a prehistoric high-altitude residential site. Located in Africa's largest alpine ecosystem, the repeated occupation of Fincha Habera rock shelter is dated to 47 to 31 thousand years ago. The available resources in cold and glaciated environments included the exploitation of an endemic rodent as a key food source, and this played a pivotal role in facilitating the occupation of this site by Late Pleistocene hunter-gatherers.

Radar vision in the mapping of forest biodiversity from space.

Recent progress in remote sensing provides much-needed, large-scale spatio-temporal information on habitat structures important for biodiversity conservation. Here we examine the potential of a newly launched satellite-borne radar system (Sentinel-1) to map the biodiversity of twelve taxa across five temperate forest regions in central Europe. We show that the sensitivity of radar to habitat structure is similar to that of airborne laser scanning (ALS), the current gold standard in the measurement of forest structure. Our models of different facets of biodiversity reveal that radar performs as well as ALS; median R² over twelve taxa by ALS and radar are 0.51 and 0.57 respectively for the first non-metric multidimensional scaling axes representing assemblage composition. We further demonstrate the promising predictive ability of radar-derived data with external validation based on the species composition of birds and saproxylic beetles. Establishing new area-wide biodiversity monitoring by remote sensing will require the coupling of radar data to stratified and standardized collected local species data.

Plant and animal functional diversity drive mutualistic network assembly across an elevational gradient.

Species' functional traits set the blueprint for pair-wise interactions in ecological networks. Yet, it is unknown to what extent the functional diversity of plant and animal communities controls network assembly along environmental gradients in real-world ecosystems. Here we address this question with a unique dataset of mutualistic bird-fruit, bird-flower and insect-flower interaction networks and associated functional traits of 200 plant and 282 animal species sampled along broad climate and land-use gradients on Mt. Kilimanjaro. We show that plant functional diversity is mainly limited by precipitation, while animal functional diversity is primarily limited by temperature. Furthermore, shifts in plant and animal functional diversity along the elevational gradient control the niche breadth and partitioning of the respective other trophic level. These findings reveal that climatic constraints on the functional diversity of either plants or animals determine the relative importance of bottom-up and top-down control in plant-animal interaction networks.

Multiple forest attributes underpin the supply of multiple ecosystem services.

Trade-offs and synergies in the supply of forest ecosystem services are common but the drivers of these relationships are poorly understood. To guide management that seeks to promote multiple services, we investigated the relationships between 12 stand-level forest attributes, including structure, composition, heterogeneity and plant diversity, plus 4 environmental factors, and proxies for 14 ecosystem services in 150 temperate forest plots. Our results show that forest attributes are the best predictors of most ecosystem services and are also good predictors of several synergies and trade-offs between services. Environmental factors also play an important role, mostly in combination with forest attributes. Our study suggests that managing forests to increase structural heterogeneity, maintain large trees, and canopy gaps would promote the supply of multiple ecosystem services. These results highlight the potential for forest management to encourage multifunctional forests and suggest that a coordinated landscape-scale strategy could help to mitigate trade-offs in human-dominated landscapes.

Predictors of elevational biodiversity gradients change from single taxa to the multi-taxa community level.

The factors determining gradients of biodiversity are a fundamental yet unresolved topic in ecology. While diversity gradients have been analysed for numerous single taxa, progress towards general explanatory models has been hampered by limitations in the phylogenetic coverage of past studies. By parallel sampling of 25 major plant and animal taxa along a 3.7 km elevational gradient on Mt. Kilimanjaro, we quantify cross-taxon consensus in diversity gradients and evaluate predictors of diversity from single taxa to a multi-taxa community level. While single taxa show complex distribution patterns and respond to different environmental factors, scaling up diversity to the community level leads to an unambiguous support for temperature as the main predictor of species richness in both plants and animals. Our findings illuminate the influence of taxonomic coverage for models of diversity gradients and point to the importance of temperature for diversification and species coexistence in plant and animal communities.