Summary 1Relationships between above-ground net primary productivity (ANPP) of grasslands and annual precipitation are often weak at the site level, with much of the inter-annual variation in ANPP left unexplained. A potential reason for this is that the distribution of precipitation within a growing season affects productivity in addition to the total amount. 2 We analysed long-term ANPP data for three southern African temperate grasslands (mean annual precipitation ranging from 538 mm to 798 mm) to determine the effects of precipitation event size, number and spacing relative to seasonal totals. 3 Ungrazed, non-manipulated treatments at each site showed contrasting results despite sharing a common, dominant species. At the driest site, a model combining average event size and number of events per growing season provided a substantially better fit to the ANPP data than precipitation amount (seasonal total). At the wettest site, the interval between events was the most important precipitation variable. Precipitation distribution was not important at the intermediate site where amount was the best predictor of ANPP. A limit to the size of precipitation events efficiently utilized for ANPP was evident for the driest site only. 4 At each site, experimental treatments that altered species composition and soil fertility had little effect on precipitation-ANPP relationships. The lack of consistency in the relative importance of the precipitation variables among sites suggests that local, edaphic factors modify precipitation-ANPP relationships. 5 This analysis demonstrates that the distribution and size of precipitation events can affect ANPP independent of precipitation amount. As altered precipitation regimes are forecast by global climate models, the sensitivity of ecosystems to precipitation distribution should be considered when predicting responses to climate change. 6 While mean values of precipitation, and other ecosystem drivers, are typically used to predict function at the level of whole ecosystems, our results show that more complex measures of environmental variability may be required to understand ecosystem function, and to increase the accuracy of predictions of ecosystem responses to global change.

Tree biomass is an integrated measure of net growth and is critical for understanding, monitoring, and modeling ecosystem functions. Despite the importance of accurately measuring tree biomass, several fundamental barriers preclude direct measurement at large spatial scales, including the facts that trees must be felled to be weighed and that even modestly sized trees are challenging to maneuver once felled. Allometric methods allow for estimation of tree mass using structural characteristics, such as trunk diameter. Savanna trees present additional challenges, including limited available allometry and a prevalence of multiple stems per individual. Here we collected airborne lidar data over a semiarid savanna adjacent to the Kruger National Park, South Africa, and then harvested and weighed woody plant biomass at the plot scale to provide a standard against which field and airborne estimation methods could be compared. For an existing airborne lidar method, we found that half of the total error was due to averaging canopy height at the plot scale. This error was eliminated by instead measuring maximum height and crown area of individual trees from lidar data using an object-based method to identify individual tree crowns and estimate their biomass. The best object-based model approached the accuracy of field allometry at both the tree and plot levels, and it more than doubled the accuracy compared to existing airborne methods (17% vs. 44% deviation from harvested biomass). Allometric error accounted for less than one-third of the total residual error in airborne biomass estimates at the plot scale when using allometry with low bias. Airborne methods also gave more accurate predictions at the plot level than did field methods based on diameter-only allometry. These results provide a novel comparison of field and airborne biomass estimates using harvested plots and advance the role of lidar remote sensing in savanna ecosystems.

Grazing represents the most extensive use of land worldwide. Yet its impacts on ecosystem services remain uncertain because pervasive interactions between grazing pressure, climate, soil properties, and biodiversity may occur but have never been addressed simultaneously. Using a standardized survey at 98 sites across six continents, we show that interactions between grazing pressure, climate, soil, and biodiversity are critical to explain the delivery of fundamental ecosystem services across drylands worldwide. Increasing grazing pressure reduced ecosystem service delivery in warmer and species-poor drylands, whereas positive effects of grazing were observed in colder and species-rich areas. Considering interactions between grazing and local abiotic and biotic factors is key for understanding the fate of dryland ecosystems under climate change and increasing human pressure.

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The idea that tropical forest and savanna are alternative states is crucial to how we manage these biomes and predict their future under global change. Large-scale empirical evidence for alternative stable states is limited, however, and comes mostly from the multimodal distribution of structural aspects of vegetation. These approaches have been criticized, as structure alone cannot separate out wetter savannas from drier forests for example, and there are also technical challenges to mapping vegetation structure in unbiased ways. Here, we develop an alternative approach to delimit the climatic envelope of the two biomes in Africa using tree species lists gathered for a large number of forest and savanna sites distributed across the continent. Our analyses confirm extensive climatic overlap of forest and savanna, supporting the alternative stable states hypothesis for Africa, and this result is corroborated by paleoecological evidence. Further, we find the two biomes to have highly divergent tree species compositions and to represent alternative compositional states. This allowed us to classify tree species as forest vs. savanna specialists, with some generalist species that span both biomes. In conjunction with georeferenced herbarium records, we mapped the forest and savanna distributions across Africa and quantified their environmental limits, which are primarily related to precipitation and seasonality, with a secondary contribution of fire. These results are important for the ongoing efforts to restore African ecosystems, which depend on accurate biome maps to set appropriate targets for the restored states but also provide empirical evidence for broad-scale bistability.

ABSTRACT. The International Long-Term Ecological Research (ILTER) network comprises > 600 scientific groups conducting sitebased research within 40 countries. Its mission includes improving the understanding of global ecosystems and informs solutions to current and future environmental problems at the global scales. The ILTER network covers a wide range of social-ecological conditions and is aligned with the Programme on Ecosystem Change and Society (PECS) goals and approach. Our aim is to examine and develop the conceptual basis for proposed collaboration between ILTER and PECS. We describe how a coordinated effort of several contrasting LTER site-based research groups contributes to the understanding of how policies and technologies drive either toward or away from the sustainable delivery of ecosystem services. This effort is based on three tenets: transdisciplinary research; cross-scale interactions and subsequent dynamics; and an ecological stewardship orientation. The overarching goal is to design management practices taking into account trade-offs between using and conserving ecosystems toward more sustainable solutions. To that end, we propose a conceptual approach linking ecosystem integrity, ecosystem services, and stakeholder well-being, and as a way to analyze trade-offs among ecosystem services inherent in diverse management options. We also outline our methodological approach that includes: (i) monitoring and synthesis activities following spatial and temporal trends and changes on each site and by documenting cross-scale interactions; (ii) developing analytical tools for integration; (iii) promoting trans-site comparison; and (iv) developing conceptual tools to design adequate policies and management interventions to deal with trade-offs. Finally, we highlight the heterogeneity in the socialecological setting encountered in a subset of 15 ILTER sites. These study cases are diverse enough to provide a broad cross-section of contrasting ecosystems with different policy and management drivers of ecosystem conversion; distinct trends of biodiversity change; different stakeholders' preferences for ecosystem services; and diverse components of well-being issues.

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Summary1. Small fenced reserves are perceived to require management interventions to maintain ecosystems in a natural state. Such interventions are typically initiated and assessed on the basis of short-term observations, while slow changes are often misunderstood or missed entirely. Long-term monitoring is therefore crucial to understand the effect of reserve management on ecosystem structure and functioning. 2. We analysed a 15-year data set of herbaceous vegetation data for 59 monitoring sites in two nature reserves of different size and with different management regimes in a semi-arid, South African savanna. Community composition and vegetation structure (basal cover) were assessed in response to rainfall and management variables. 3. No directional changes over time were found. Basal cover as measured by two proxies (distance to the nearest tuft and tuft diameter) increased with current and previous year rainfall. The relative abundance of functional groups also responded to rainfall, with perennial grasses increasing in relative abundance following a wet year. Compositional responses, as measured by a dissimilarity index of species relative abundance, showed a 2-year lag and responses were larger following dry years than wet years. 4. The response to rainfall of distance to the nearest tuft was significantly weaker in the smaller reserve, while compositional change during dry years was larger than in the larger reserve. 5. Synthesis and applications. Most of the variation in herbaceous basal cover and community composition was associated with differences between years (time) at each site, rather than with differences between sites (space), indicating that inter-annual variation in rainfall is the most important driver of herbaceous layer dynamics in these systems. However, management did modify the effects of rainfall on herbaceous structure and community composition. The smaller reserve, which had higher grazer and waterhole density, showed greater fluctuations in key herbaceous variables. Such reserves are common in southern Africa and probably require more careful management than larger reserves in the face of global climate change.