Context dependency of relationships between biodiversity and ecosystem functioning is different for multiple ecosystem functions

Increasing concern over the loss of biodiversity has led to attempts to quantify relationships between biodiversity and ecosystem functioning. While manipulative investigations have accumulated substantial evidence to support the notion that decreasing biodiversity can be detrimental to the functioning of ecosystems, recent investigations have identified the potential importance of physical processes in moderating biodiversity – ecosystem function relationships at larger geographical scales. In this study, the relationship between the genus richness of benthic macro‐invertebrates and five measures of ecosystem functioning (macrofaunal biomass, depth of the apparent redox discontinuity, fluxes of ammonium and NO_x and the abundance of nematodes) was determined over a large scale wave‐induced bed shear stress gradient on the seabed of the northern Irish Sea. Ecosystem functioning was significantly correlated to genus richness for four out of five ecosystem functions. However, wave stress moderated the genus richness – ecosystem functioning relationship for only one of the ecosystem functions; genus richness had a positive effect on the depth of the apparent redox discontinuity in the sediment at high wave stress but not at low wave stress. These results indicate that the effects of biodiversity on some ecosystem functions may be sufficiently strong to generate patterns in ecosystems where other factors are also affecting ecosystem processes, but that the biodiversity–ecosystem function relationship for can be dependent on environmental conditions for specific ecosystem functions.     

Flow regime alteration effects on the organic C dynamics in semiarid stream ecosystems

There is evidence of an ongoing alteration of the flow regime owing to climate change forcing, which has resulted in substantial increases in the frequency and magnitude of extreme events such as floods and droughts. Such changes in the flow regime may have major implications in freshwater ecosystems and, in particular, in the organic carbon dynamics in semiarid stream ecosystems. Much is known about the role of extreme flow events on structuring stream ecosystems, but few studies explored the effects of extreme flow events magnitude, timing, and sequence on stream ecosystems. To assess the effect of extreme events on stream organic C dynamics, a simple and flexible modeling approach was applied to simulate the organic carbon dynamics in a simplified river reach. The river reach model was initially calibrated and tested using long-term data for stream water velocity and amount of organic carbon in sediment. After that, multiple scenarios differing in the extreme flow events (floods and droughts) sequence and magnitude were used to simulate the effects of possible flow regime changes on the stream organic carbon dynamics. Initial expectations were that: (i) an increase in the magnitude or frequency of extreme flow events would reduce the amount of organic carbon respired within the simulated river reach, and (ii) relationship between the timings of the extreme flow events and of the litterfall input would influence considerably the effects of the extreme flow events. Results pointed out that: (i) the amount of processed carbon respect the amount entering the ecosystem was affected by extreme events such floods and droughts, but the relevance of those events differed along the year, with a maximal effect during the litterfall period; (ii) extreme event timing rather than the magnitude was more relevant to the stream organic carbon dynamics; and (iii) the amount of respired carbon in the ecosystem could be amplified or reduced depending on event sequence. Increasing awareness of the role of inland waters in the global carbon cycle and the shaping role of hydrology on the stream organic carbon dynamics stress the need to better quantify carbon fluxes and the hydrological controls on these fluxes.     

Extreme storms cause rapid but short-lived shifts in nearshore subtropical bacterial communities

Climate change scenarios predict tropical cyclones will increase in both frequency and intensity, which will escalate the amount of terrestrial run-off and mechanical disruption affecting coastal ecosystems. Bacteria are key contributors to ecosystem functioning, but relatively little is known about how they respond to extreme storm events, particularly in nearshore subtropical regions. In this study, we combine field observations and mesocosm experiments to assess bacterial community dynamics and changes in physicochemical properties during early- and late-season tropical cyclones affecting Okinawa, Japan. Storms caused large and fast influxes of freshwater and terrestrial sediment – locally known as red soil pollution – and caused moderate increases of macronutrients, especially SiO₂ and PO₄³⁻, with up to 25 and 0.5 μM respectively. We detected shifts in relative abundances of marine and terrestrially derived bacteria, including putative coral and human pathogens, during storm events. Soil input alone did not substantially affect marine bacterial communities in mesocosms, indicating that other components of run-off or other storm effects likely exert a larger influence on bacterial communities. The storm effects were short-lived and bacterial communities quickly recovered following both storm events. The early- and late-season storms caused different physicochemical and bacterial community changes, demonstrating the context-dependency of extreme storm responses in a subtropical coastal ecosystem.     

Soil biotic processes remain remarkably stable after 100-year extreme weather events in experimental grassland and heath

Climate change will increase the recurrence of extreme weather events such as drought and heavy rainfall. Evidence suggests that extreme weather events pose threats to ecosystem functioning, particularly to nutrient cycling and biomass production. These ecosystem functions depend strongly on below-ground biotic processes, including the activity and interactions among plants, soil fauna, and micro-organisms. Here, experimental grassland and heath communities of three phytodiversity levels were exposed either to a simulated single drought or to a heavy rainfall event. Both weather manipulations were repeated for two consecutive years. The magnitude of manipulations imitated the local 100-year extreme weather event. Heavy rainfall events increased below-ground plant biomass and stimulated soil enzyme activities as well as decomposition rates for both plant communities. In contrast, extreme drought did not reduce below-ground plant biomass and root length, soil enzyme activities, and cellulose decomposition rate. The low responsiveness of the measured ecosystem properties in face of the applied weather manipulations rendered the detection of significant interactions between weather events and phytodiversity impossible. Our data indicate on the one hand the close interaction between below ground plant parameters and microbial turnover processes in soil; on the other hand it shows that the plant–soil system can buffer against extreme drought events, at last for the period of investigation.     

Effects of Extreme Weather Events on Plant Productivity and Tissue Die-Back are Modified by Community Composition

Extreme weather events are expected to increase in frequency and magnitude due to climate change. Their effects on vegetation are widely unknown. Here, experimental grassland and heath communities in Central Europe were exposed either to a simulated single drought or to a prolonged heavy rainfall event. The magnitude of manipulations imitated the local 100-year weather extreme according to extreme value statistics. Overall productivity of both plant communities remained stable in the face of drought and heavy rainfall, despite significant effects on tissue die-back. Grassland communities were more resistant against the extreme weather events than heath communities. Furthermore, effects of extreme weather events on community tissue die-back were modified by functional diversity, even though conclusiveness in this part is limited by the fact that only one species composition was available per diversity level within this case study. More diverse grassland communities exhibited less tissue die-back than less complex grassland communities. On the other side, more diverse heath communities were more vulnerable to extreme weather events compared to less complex heath communities. Furthermore, legumes did not effectively contribute to the buffering against extreme weather events in both vegetation types. Tissue die-back proved a strong stress response in plant communities exposed to 100-year extreme weather events, even though one important ecosystem function, namely productivity, remained surprisingly stable in this experiment. Theories and concepts on biodiversity and ecosystem functioning (insurance hypothesis, redundancy hypothesis) may have to be revisited when extreme weather conditions are considered.     

Eukaryotes contribute more than bacteria to the recovery of freshwater ecosystem functions under different drought durations

Global climate change mostly impacts river ecosystems by affecting microbial biodiversity and ecological functions. Considering the high functional redundancy of microorganisms, the unknown relationship between biodiversity and ecosystem functions obstructs river ecological research, especially under the influence of increasing weather extremes, such as in intermittent rivers and ephemeral streams (IRES). Herein, dry–wet alternation experiments were conducted in artificial stream channels for 25 and 90 days of drought, both followed by 20 days of rewetting. The dynamic recovery of microbial biodiversity and ecosystem functions (represented by ecosystem metabolism and denitrification rate) were determined to analyse biodiversity–ecosystem–function (BEF) relationships after different drought durations. There was a significant difference between bacterial and eukaryotic biodiversity recovery after drought. Eukaryotic biodiversity was more sensitive to drought duration than bacterial, and the eukaryotic network was more stable under dry–wet alternations. Based on the establishment of partial least squares path models, we found that eukaryotic biodiversity has a stronger effect on ecosystem functions than bacteria after long-term drought. Indeed, this work represents a significant step forward for further research on the ecosystem functions of IRES, especially emphasizing the importance of eukaryotic biodiversity in the BEF relationship.     

Stronger Short-Term Effects of Mowing Than Extreme Summer Weather on a Subalpine Grassland

Mowing is known to favor plant diversity and influence ecosystem functioning in semi-natural grasslands. This effect could be influenced by climate variability, especially in regions with harsh climate, such as subalpine zones. In particular, short-term extreme weather fluctuations may induce rapid plant responses, affecting in turn the response to mowing. We tested the effects of concomitant summer weather manipulation and mowing on a subalpine grassland in the Central French Alps for two consecutive years. We addressed two questions: (1) How is a subalpine grassland affected by extreme summer weather? (2) Does extreme summer weather alter mowing effects on the grassland plant diversity and functioning? We used a multi-level, integrative approach assessing the responses of six abundant plant species, as well as effects on plant community structure, biomass production, and litter decomposition rates. Extreme summer weather was simulated by increasing summer temperature by 1.1°C, and decreasing summer rainfall by 80%—resulting in a 30% decrease in total annual precipitation. In addition, a heat-wave event was simulated during the first year of the experiment. This weather manipulation was combined with a late-summer mowing treatment (mown vs. unmown). Extreme summer weather mainly increased leaf senescence and decreased plant vegetative growth. Leaf litter decomposition was slowed, but only for species characterized by the fastest rates of litter decomposition. Mowing increased plant diversity by restricting the dominant grass species, thereby favoring subordinates. In the short term, this subalpine grassland was rather resistant to extreme summer weather, whereas mowing cessation remained the main factor affecting its biodiversity.     

Invader presence disrupts the stabilizing effect of species richness in plant community recovery after drought

Higher biodiversity can stabilize the productivity and functioning of grassland communities when subjected to extreme climatic events. The positive biodiversity–stability relationship emerges via increased resistance and/or recovery to these events. However, invader presence might disrupt this diversity–stability relationship by altering biotic interactions. Investigating such disruptions is important given that invasion by non‐native species and extreme climatic events are expected to increase in the future due to anthropogenic pressure. Here we present one of the first multisite invader × biodiversity × drought manipulation experiment to examine combined effects of biodiversity and invasion on drought resistance and recovery at three semi‐natural grassland sites across Europe. The stability of biomass production to an extreme drought manipulation (100% rainfall reduction; BE: 88 days, BG: 85 days, DE: 76 days) was quantified in field mesocosms with a richness gradient of 1, 3, and 6 species and three invasion treatments (no invader, Lupinus polyphyllus, Senecio inaequidens). Our results suggest that biodiversity stabilized community productivity by increasing the ability of native species to recover from extreme drought events. However, invader presence turned the positive and stabilizing effects of diversity on native species recovery into a neutral relationship. This effect was independent of the two invader's own capacity to recover from an extreme drought event. In summary, we found that invader presence may disrupt how native community interactions lead to stability of ecosystems in response to extreme climatic events. Consequently, the interaction of three global change drivers, climate extremes, diversity decline, and invasive species, may exacerbate their effects on ecosystem functioning.     

The relationship between biodiversity and ecosystem functioning in food webs

Recent theoretical and experimental work provides clear evidence that biodiversity loss can have profound impacts on functioning of natural and managed ecosystems and the ability of ecosystems to deliver ecological services to human societies. Work on simplified ecosystems in which the diversity of a single trophic level is manipulated shows that diversity can enhance ecosystem processes such as primary productivity and nutrient retention. Theory also strongly suggests that biodiversity can act as biological insurance against potential disruptions caused by environmental changes. However, these studies generally concern a single trophic level, primary producers for the most part. Changes in biodiversity also affect ecosystem functioning through trophic interactions. Here we review, through the analysis of a simple ecosystem model, several key aspects inherent in multitrophic systems that may strongly affect the relationship between diversity and ecosystem processes. Our analysis shows that trophic interactions have a strong impact on the relationships between diversity and ecosystem functioning, whether the ecosystem property considered is total biomass or temporal variability of biomass at the various trophic levels. In both cases, food-web structure and trade-offs that affect interaction strength have major effects on these relationships. Multitrophic interactions are expected to make biodiversity–ecosystem functioning relationships more complex and non-linear, in contrast to the monotonic changes predicted for simplified systems with a single trophic level.     

Linking biodiversity indicators,ecosystem functioning,provision of services and human well-being in estuarine systems: Application of a conceptual framework

Assuming that human well-being strongly relies on the services provided by well-functioning ecosystems, changes in the ecological functioning of any system can have direct and indirect effects on human welfare. Intensive land use and tourism have expanded in recent decades along coastal ecosystems, together with increasing demands for water, food and energy; all of these factors intensify the exploitation of natural resources. Many of the interrelations between ecosystem functioning and the provision of ecosystem services (ES) still require quantification in estuarine systems. A conceptual framework to assess such links in a spatially and temporally explicit manner is proposed and applied to the Mondego estuary (Portugal). This framework relies on three consecutive steps and discriminates among biodiversity structural components, ecosystem functioning and stability and the services provided by the ecosystem.Disturbances in abiotic factors were found to have a direct effect on biodiversity, ecosystem functioning and the provision of ES. The observed changes in the species composition of communities had a positive effect on the ecosystem's productivity and stability. Moreover, the observed changes in the estuarine ES provision are likely to arise from changing structural and abiotic factors and in the present case from the loss or decline of locally abundant species. This study also indicates that linear relationships between biodiversity, ecosystem functioning and services provision are unlikely to occur in estuarine systems. Instead, cumulative and complex relations are observed between factors on both temporal and spatial scales. In this context, the results suggest several additional conclusions: (1) biodiversity and ecosystem functioning interaction with human well-being need to be incorporated into decision-making processes aimed at the conservative management of systems; (2) the institutional use of research results must be part of the design and implementation of sustainable management activities; and (3) more integrative tools/studies are required to account for the interactions of estuarine ecosystems with surrounding socio-economic activities. Therefore, when performing integrated assessments of ecosystem dynamics, it becomes essential to consider not only the effects of biodiversity and ecosystem functioning on services provision but also the effects that human well-being and ES provision may have on estuarine biodiversity and ecosystem functioning.The proposed framework implies taking into account both the functional and the commodities points of view upon natural ecosystems and by this representing a line of thought which will deserve further research to explore more in detail the conceptual links between biodiversity–ecosystem functioning–services provided.     

Extreme winter warming events more negatively impact small rather than large soil fauna: shift in community composition explained by traits not taxa

Extreme weather events can have negative impacts on species survival and community structure when surpassing lethal thresholds. Extreme winter warming events in the Arctic rapidly melt snow and expose ecosystems to unseasonably warm air (2–10 °C for 2–14 days), but returning to cold winter climate exposes the ecosystem to lower temperatures by the loss of insulating snow. Soil animals, which play an integral part in soil processes, may be very susceptible to such events depending on the intensity of soil warming and low temperatures following these events. We simulated week‐long extreme winter warming events – using infrared heating lamps, alone or with soil warming cables – for two consecutive years in a sub‐Arctic dwarf shrub heathland. Minimum temperatures were lower and freeze‐thaw cycles were 2–11 times more frequent in treatment plots compared with control plots. Following the second event, Acari populations decreased by 39%; primarily driven by declines of Prostigmata (69%) and the Mesostigmatic nymphs (74%). A community‐weighted vertical stratification shift occurred from smaller soil dwelling (eu‐edaphic) Collembola species dominance to larger litter dwelling (hemi‐edaphic) species dominance in the canopy‐with‐soil warming plots compared with controls. The most susceptible groups to these winter warming events were the smallest individuals (Prostigmata and eu‐edaphic Collembola). This was not apparent from abundance data at the Collembola taxon level, indicating that life forms and species traits play a major role in community assembly following extreme events. The observed shift in soil community can cascade down to the micro‐flora affecting plant productivity and mineralization rates. Short‐term extreme weather events have the potential to shift community composition through trait composition with potentially large consequences for ecosystem development.     

Anomalous,extreme weather disrupts obligate seed dispersal mutualism: snow in a subtropical forest ecosystem

Ongoing global climate change is predicted to increase the frequency and magnitude of extreme weather events, impacting population dynamics and community structure. There is, however, a critical lack of case studies considering how climatic perturbations affect biotic interactions. Here, we document how an obligate seed dispersal mutualism was disrupted by a temporally anomalous and meteorologically extreme interlude of unseasonably frigid weather, with accompanying snowstorms, in subtropical China, during January–February 2008. Based on the analysis of 5892 fecal samples (representing six mammalian seed dispersers), this event caused a substantial disruption to the relative seed dispersal function for the raisin tree Hovenia dulcis from prestorm 6.29 (2006) and 11.47 (2007), down to 0.35 during the storm (2008). Crucially, this was due to impacts on mammalian seed dispersers and not due to a paucity of fruit, where 4.63 fruit per branch were available in January 2008, vs. 3.73 in 2006 and 3.58 in 2007. An induced dietary shift occurred among omnivorous carnivores during this event, from the consumption fruit to small mammals and birds, reducing their role in seed dispersal substantially. Induced range shift extinguished the functionality of herbivorous mammals completely, however, seed dispersal function was compensated in part by three omnivorous carnivores during poststorm years, and thus while the mutualism remained intact it was enacted by a narrower assemblage of species, rendering the system more vulnerable to extrinsic perturbations. The storm's extended effects also had anthropogenic corollaries – migrating ungulates becoming exposed to heightened levels of illegal hunting – causing long‐term modification to the seed dispersal community and mutualism dynamics. Furthermore, degraded forests proved especially vulnerable to the storm's effects. Considering increasing climate variability and anthropogenic disturbance, the impacts of such massive, aberrant events warrant conservation concern, while affording unique insights into the stability of mutualisms and the processes that structure biodiversity and mediate ecosystem dynamics.     

Effects of species evenness can be derived from species richness – ecosystem functioning relationships

Although species richness effects on ecosystem functioning have been studied thoroughly in countless experiments, the effects of the other side of diversity – species evenness – remain less identified, especially at high species richness. Due to the large number of different model ecosystems that need to be created, the explanatory power of the experimental approach for evenness is indeed limited. We show here that experimental studies on the influence of species richness on ecosystem functions contain hidden information on the influence of species evenness. Both the effects of maximum and minimum evenness, and of a key set of intermediate evenness levels, can be derived from species richness – ecosystem function curves, and that for every richness level, by using communities with low species richness as the equivalent of highly uneven communities with higher richness. We show that evenness effects on ecosystem functioning have the same direction as richness effects, however with increasing effect sizes at higher richness levels. We validated our technique for a wide range of ecosystem functions and applied it to the species richness – community biomass data from an existing biodiversity experiment. Our approach could provide a fast and easy alternative to resource‐intensive experiments in which evenness is experimentally varied, as we can build on the elaborate existing literature on species richness to assess its effects.     

Resilience and stability of a pelagic marine ecosystem

The accelerating loss of biodiversity and ecosystem services worldwide has accentuated a long-standing debate on the role of diversity in stabilizing ecological communities and has given rise to a field of research on biodiversity and ecosystem functioning (BEF). Although broad consensus has been reached regarding the positive BEF relationship, a number of important challenges remain unanswered. These primarily concern the underlying mechanisms by which diversity increases resilience and community stability, particularly the relative importance of statistical averaging and functional complementarity. Our understanding of these mechanisms relies heavily on theoretical and experimental studies, yet the degree to which theory adequately explains the dynamics and stability of natural ecosystems is largely unknown, especially in marine ecosystems. Using modelling and a unique 60-year dataset covering multiple trophic levels, we show that the pronounced multi-decadal variability of the Southern California Current System (SCCS) does not represent fundamental changes in ecosystem functioning, but a linear response to key environmental drivers channelled through bottom-up and physical control. Furthermore, we show strong temporal asynchrony between key species or functional groups within multiple trophic levels caused by opposite responses to these drivers. We argue that functional complementarity is the primary mechanism reducing community variability and promoting resilience and stability in the SCCS.     

Biodiversity effects on ecosystem functioning: emerging issues and their experimental test in aquatic environments

Recent experiments, mainly in terrestrial environments, have provided evidence of the functional importance of biodiversity to ecosystem processes and properties. Compared to terrestrial systems, aquatic ecosystems are characterised by greater propagule and material exchange, often steeper physical and chemical gradients, more rapid biological processes and, in marine systems, higher metazoan phylogenetic diversity. These characteristics limit the potential to transfer conclusions derived from terrestrial experiments to aquatic ecosystems whilst at the same time provide opportunities for testing the general validity of hypotheses about effects of biodiversity on ecosystem functioning. Here, we focus on a number of unique features of aquatic experimental systems, propose an expansion to the scope of diversity facets to be considered when assessing the functional consequences of changes in biodiversity and outline a hierarchical classification scheme of ecosystem functions and their corresponding response variables. We then briefly highlight some recent controversial and newly emerging issues relating to biodiversity‐ecosystem functioning relationships. Based on lessons learnt from previous experimental and theoretical work, we finally present four novel experimental designs to address largely unresolved questions about biodiversity‐ecosystem functioning relationships. These include (1) investigating the effects of non‐random species loss through the manipulation of the order and magnitude of such loss using dilution experiments; (2) combining factorial manipulation of diversity in interconnected habitat patches to test the additivity of ecosystem functioning between habitats; (3) disentangling the impact of local processes from the effect of ecosystem openness via factorial manipulation of the rate of recruitment and biodiversity within patches and within an available propagule pool; and (4) addressing how non‐random species extinction following sequential exposure to different stressors may affect ecosystem functioning. Implementing these kinds of experimental designs in a variety of systems will, we believe, shift the focus of investigations from a species richness‐centred approach to a broader consideration of the multifarious aspects of biodiversity that may well be critical to understanding effects of biodiversity changes on overall ecosystem functioning and to identifying some of the potential underlying mechanisms involved.     

Spatial co‐localisation of extreme weather events: a clear and present danger

Extreme weather events have become a dominant feature of the narrative surrounding changes in global climate with large impacts on ecosystem stability, functioning and resilience; however, understanding of their risk of co‐occurrence at the regional scale is lacking. Based on the UK Met Office’s long‐term temperature and rainfall records, we present the first evidence demonstrating significant increases in the magnitude, direction of change and spatial co‐localisation of extreme weather events since 1961. Combining this new understanding with land‐use data sets allowed us to assess the likely consequences on future agricultural production and conservation priority areas. All land‐uses are impacted by the increasing risk of at least one extreme event and conservation areas were identified as the hotspots of risk for the co‐occurrence of multiple event types. Our findings provide a basis to regionally guide land‐use optimisation, land management practices and regulatory actions preserving ecosystem services against multiple climate threats.     

Effects of hydromorphological integrity on biodiversity and functioning of river ecosystems

River channels tend to a dynamic equilibrium driven by the dynamics of water and sediment discharge. The resulting fluctuating pattern of channel form is affected by the slope, the substrate erodibility, and the vegetation in the river corridor and in the catchment. Geomorphology is basic to river biodiversity and ecosystem functioning since the channel pattern provides habitat for the biota and physical framework for ecosystem processes. Human activities increasingly change the natural drivers of channel morphology on a global scale (e.g. urbanization increases hydrological extremes, and clearing of forests for agriculture increases sediment yield). In addition, human actions common along world rivers impact channel dynamics directly, e.g. river regulation simplifies and fossilizes channel form. River conservation and restoration must incorporate mechanisms of channel formation and ecological consequences of channel form and dynamics. This article (1) summarizes the role of channel form on biodiversity and functioning of river ecosystems, (2) describes spatial complexity, connectivity and dynamism as three key hydromorphological attributes, (3) identifies prevalent human activities that impact these key components and (4) analyzes gaps in current knowledge and identifies future research topics.     

Legacy effects and memory loss: how contingencies moderate the response of rocky intertidal biofilms to present and past extreme events

Understanding how historical processes modulate the response of ecosystems to perturbations is becoming increasingly important. In contrast to the growing interest in projecting biodiversity and ecosystem functioning under future climate scenarios, how legacy effects originating from historical conditions drive change in ecosystems remains largely unexplored. Using experiments in combination with stochastic antecedent modelling, we evaluated how extreme warming, sediment deposition and grazing events modulated the ecological memory of rocky intertidal epilithic microphytobenthos (EMPB). We found memory effects in the non‐clustered scenario of disturbance (60 days apart), where EMPB biomass fluctuated in time, but not under clustered disturbances (15 days apart), where EMPB biomass was consistently low. A massive grazing event impacted on EMPB biomass in a second run of the experiment, also muting ecological memory. Our results provide empirical support to the theoretical expectation that stochastic fluctuations promote ecological memory, but also show that contingencies may lead to memory loss.     

Climatic and biotic extreme events moderate long‐term responses of above‐ and belowground sub‐Arctic heathland communities to climate change

Climate change impacts are not uniform across the Arctic region because interacting factors causes large variations in local ecosystem change. Extreme climatic events and population cycles of herbivores occur simultaneously against a background of gradual climate warming trends and can redirect ecosystem change along routes that are difficult to predict. Here, we present the results from sub‐Arctic heath vegetation and its belowground micro‐arthropod community in response to the two main drivers of vegetation damage in this region: extreme winter warming events and subsequent outbreaks of the defoliating autumnal moth caterpillar (Epirrita autumnata). Evergreen dwarf shrub biomass decreased (30%) following extreme winter warming events and again by moth caterpillar grazing. Deciduous shrubs that were previously exposed to an extreme winter warming event were not affected by the moth caterpillar grazing, while those that were not exposed to warming events (control plots) showed reduced (23%) biomass from grazing. Cryptogam cover increased irrespective of grazing or winter warming events. Micro‐arthropods declined (46%) following winter warming but did not respond to changes in plant community. Extreme winter warming and caterpillar grazing suppressed the CO₂ fluxes of the ecosystem. Evergreen dwarf shrubs are disadvantaged in a future sub‐Arctic with more stochastic climatic and biotic events. Given that summer warming may further benefit deciduous over evergreen shrubs, event and trend climate change may both act against evergreen shrubs and the ecosystem functions they provide. This is of particular concern given that Arctic heath vegetation is typically dominated by evergreen shrubs. Other components of the vegetation showed variable responses to abiotic and biotic events, and their interaction indicates that sub‐Arctic vegetation response to multiple pressures is not easy to predict from single‐factor responses. Therefore, while biotic and climatic events may have clear impacts, more work is needed to understand their net effect on Arctic ecosystems.     

Trophic and non‐trophic interactions influence the mechanisms underlying biodiversity–ecosystem functioning relationships under different abiotic conditions

Plant diversity effects on ecosystem functioning usually have been studied from a plant perspective. However, the mechanisms underlying biodiversity–ecosystem functioning relationships may also depend on positive or negative interactions between plants and other biotic and abiotic factors, which remain poorly understood. Here we assessed whether plant–herbivore and/or plant–detritivore interactions modify the biodiversity–ecosystem functioning relationship and the mechanisms underlying biodiversity effects, including complementarity and selection effects, biomass allocation, vertical distribution of roots, and plant survival using a microcosm experiment. We also evaluated to what extent trophic and non‐trophic interactions are affected by abiotic conditions by studying drought effects. Our results show that biotic and abiotic conditions influence the shape of the biodiversity–ecosystem function relationship, varying from hump‐shaped to linear. For instance, total biomass increased linearly with plant richness in the presence of detritivores, but not in the absence of detritivores. Moreover, detritivore effects on belowground plant productivity were highly context dependent, varying in the presence of herbivores. Plant interactions with soil biota, especially with herbivores, influenced the mechanisms underlying diversity effects. Herbivores increased plant complementarity and modified biomass allocation and vertical distribution of roots. Furthermore, biotic–abiotic interactions influenced plant productivity differently across plant functional groups. Our findings emphasize the importance of complex biotic interactions underlying biodiversity effects, and that these biotic interactions may change with abiotic conditions. Despite minor changes in productivity in the short‐term, soil biota‐induced changes in plant–plant interactions and plant survival are likely to have significant long‐term consequences for ecosystem functioning. Considering the context‐dependency of multichannel interactions may contribute to reconciling differences among observed patterns in biodiversity studies. Further, abiotic conditions modified the effects of biotic interactions, suggesting that changes in environmental conditions may not only affect ecosystems directly, but also change the biotic composition of and dynamics within ecosystems.