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Understanding the relationship of soil carbon storage and species diversity in grasslands can provide insights into managing these ecosystems. We studied relationships among soil C and plant species richness within ~ 9700 ha of grasslands in Colorado, US. Using 141 grassland transects, we tested how soil C was related to plant species richness, grassland type, soil texture, and prairie dog presence. Soil C was significantly, positively related to plant species richness, while native perennial graminoid species richness exhibited an even stronger positive relationship. However, the relationship of soil C and plant richness was not found in all three grassland types studied, but instead was unique to the most common grassland type, mixed grass prairie, and absent from both xeric tallgrass and mesic tallgrass prairie. The presence of a single indicator species, Andropogon gerardii, showed a significant, positive relationship with soil carbon. Our best possible model explained 45% of the variance in soil C using species richness, grassland type, and their interaction. Surprisingly, soil C was negatively related to soil clay, suggesting that surface clays amplify evaporation and water runoff rather than protecting soil organic matter from decomposition. Soil C was negatively related to prairie dog presence, suggesting that prairie dogs do not enhance soil carbon sequestration; in fact, prairie dog occupied sites had significantly lower soil C, likely related to loss of topsoil from prairie dog colonies. Our results suggest that management for species richness provides the co-benefit of soil C storage, and high clay and prairie dog disturbance compromises both. Supplementary Information The online version contains supplementary material available at 10.1007/s00442-021-04992-x.

For the first time, intraspecific relationships between the macroecological metrics patchiness (P) and both abundance (A) and occupancy (O) were investigated in a faunal assemblage. As a companion study to recent work on interspecific P, A and O patterns at the same localities, intraspecific patterns were documented within each of the more dominant invertebrates forming the seagrass macrobenthos of warm–temperate Knysna estuarine bay (South Africa) and of sub-tropical Moreton Bay (Australia). As displayed interspecifically, individual species showed strong A–O patterns (mean scaling coefficient − 0.76 and mean R2 > 0.8). All P–O relations were negative and most (67%) were statistically significant, although weaker (mean R2 0.5) than A–O ones; most P–A ones were also negative but fewer (43%) achieved significance, and were even weaker (mean R2 0.4); 33% of species showed no significant interrelations of either O or A with P. No species showed only a significant P–A relationship. Compared with interspecific P–A–O data from the same assemblages, power–law scaling exponents were equivalent, but R2 values were larger. Larviparous species comprised 70% of the total studied, but 94% of those displaying significant patchiness interrelationships; 5 of the 9 showing no P–A or P–O relationships, however, were also larviparous. At Knysna, though not in Moreton Bay, larviparous species also showed higher levels of occupancy than non-larviparous ones, whilst non-larviparous species showed higher levels of patchiness. Dominant Moreton Bay species, but not those at Knysna, exhibited homogeneously sloped P–O relationships.

Environmental and dispersal filters are key determinants of species distributions of Amazonian tree communities. However, a comprehensive analysis of the role of environmental and dispersal filters is needed to understand the ecological and evolutionary processes that drive phylogenetic and taxonomic turnover of Amazonian tree communities. We compare measures of taxonomic and phylogenetic beta diversity in 41 one-hectare plots to test the relative importance of climate, soils, geology, geomorphology, pure spatial variables and the spatial variation of environmental drivers of phylogenetic and taxonomic turnover in Ecuadorian Amazon tree communities. We found low phylogenetic and high taxonomic turnover with respect to environmental and dispersal filters. In addition, our results suggest that climate is a significantly better predictor of phylogenetic turnover and taxonomic turnover than geomorphology and soils at all spatial scales. The influence of climate as a predictor of phylogenetic turnover was stronger at broader spatial scales (50 km2) whereas geomorphology and soils appear to be better predictors of taxonomic turnover at mid (5 km2) and fine spatial scales (0.5 km2) but a weak predictor of phylogenetic turnover at broad spatial scales. We also found that the combined effect of geomorphology and soils was significantly higher for taxonomic turnover at all spatial scales but not for phylogenetic turnover at large spatial scales. Geographic distances as proxy of dispersal limitation was a better predictor of phylogenetic turnover at distances of 50

Forest soil and litter is inhabited by a diverse community of animals, which directly and indirectly rely on dead organic matter as habitat and food resource. However, community composition may be driven by biotic or abiotic forces, and these vary with changes in habitat structure and resource supply associated with forest land use. To evaluate these changes, we compiled comprehensive data on the species composition of soil animal communities and environmental factors in forest types varying in land-use intensity in each of three regions in Germany, i.e., coniferous, young managed, old managed, and unmanaged beech forests. Coniferous forests featured high amounts of leaf litter and low microbial biomass concentrations contrasting in particular unmanaged beech forests. However, soil animal diversity and functional community composition differed little between forest types, indicating resilience against disturbance and forest land use. Structural equation modelling suggested that despite a significant influence of forest management on resource abundance and quality, the biomass of most soil fauna functional groups was not directly affected by forest management or resource abundance/quality, potentially because microorganisms hamper the propagation of nutrients to higher trophic levels. Instead, detritivore biomass depended heavily on soil pH. Macrofauna decomposers thrived at high pH, whereas mesofauna decomposers benefitted from low soil pH, but also from low biomass of macrofauna decomposers, potentially due to habitat modification by macrofauna decomposers. The strong influence of soil pH shows that decomposer communities are structured predominantly by regional abiotic factors exceeding the role of local biotic factors such as forest type. Supplementary Information The online version contains supplementary material available at 10.1007/s00442-021-04910-1.

Tropical rainforest disturbance and conversion are critical drivers of biodiversity loss. A key knowledge gap is understanding the impacts of habitat modification on mechanisms of community assembly, which are predicted to respond differently between taxa and across spatial scales. We use a null model approach to detect trait assembly of species at local- and landscape-scales, and then subdivide communities with different habitat associations and foraging guilds to investigate whether the detection of assembly mechanisms varies between groups. We focus on two indicator taxa, dung beetles and birds, across a disturbance gradient of primary rainforest, selectively logged rainforest, and oil palm plantations in Borneo, Southeast Asia. Random community assembly was predominant for dung beetles across habitats, whereas trait convergence, indicative of environmental filtering, occurred across the disturbance gradient for birds. Assembly patterns at the two spatial scales were similar. Subdividing for habitat association and foraging guild revealed patterns hidden when focusing on the overall community. Dung beetle forest specialists and habitat generalists showed opposing assembly mechanisms in primary forest, community assembly of habitat generalists for both taxa differed with disturbance intensity, and insectivorous birds strongly influenced overall community assembly relative to other guilds. Our study reveals the sensitivity of community assembly mechanisms to anthropogenic disturbance via a shift in the relative contribution of stochastic and deterministic processes. This highlights the need for greater understanding of how habitat modification alters species interactions and the importance of incorporating species’ traits within assessments. Supplementary Information The online version contains supplementary material available at 10.1007/s00442-020-04829-z.

Marine habitat formers such as seaweeds and corals are lynchpins of coastal ecosystems, but their functional diversity and how it varies with scale and context remains poorly studied. Here, we investigate the functional diversity of seaweed assemblages across the rocky intertidal stress gradient at large (zones) and small (quadrat) scales. We quantified complementary metrics of emergent group richness, functional richness (functional space occupied) and functional dispersion (trait complementarity of dominant species). With increasing shore height, under species loss and turnover, responses of functional diversity were scale- and metric-dependent. At the large scale, functional richness contracted while—notwithstanding a decline in redundancy—emergent group richness and functional dispersion were both invariant. At the small scale, all measures declined, with the strongest responses evident for functional and emergent group richness. Comparisons of observed versus expected values based on null models revealed that functional richness and dispersion were greater than expected in the low shore but converged with expected values higher on the shore. These results show that functional diversity of assemblages of marine habitat formers can be especially responsive to environmental stress gradients at small scales and for richness measures. Furthermore, niche-based processes at the small—neighbourhood—scale can favour co-occurrence of functionally distinctive species under low, but not high, stress, magnifying differences in functional diversity across environmental gradients. As assemblages of marine habitat formers face accelerating environmental change, further studies examining multiple aspects of functional diversity are needed to elucidate patterns, processes, and ecosystem consequences of community (dis-)assembly across diverse groups. Electronic supplementary material The online version of this article (10.1007/s00442-020-04746-1) contains supplementary material, which is available to authorized users.

To improve projections of consequences of increasing intensity and frequency of drought events for grasslands, we need a thorough understanding of species performance responses to drought, of performance trade-offs and how drought resistance is related to species distributions. However, comparative and quantitative assessments of whole-plant drought resistance that allow to rigorously address these aspects are lacking for temperate grassland species. We conducted a common garden experiment with 40 common temperate grassland species to compare species survival and growth under intense drought and well-irrigated conditions. Overall, survival and growth were significantly reduced under drought, with the effect varying across species. Species ranking of drought damage and survival remained consistent with progressing drought. No performance trade-offs emerged between optimal growth and drought resistance of survival (‘growth–stress tolerance’ trade-off hypothesis), or between growth under well-watered and dry conditions (‘growth rates’ trade-off hypothesis). Species local- and large-scale association with moisture (Ellenberg F value and rainfall niche) was not related to their drought resistance. Overall, our results imply that trade-offs and differences of species fundamental drought resistance are not the main drivers of hydrological niche differentiation, species coexistence and their distribution across moisture gradients. The comparative experimental assessment of species whole-plant drought responses we present provides a basis to increase our understanding of current grassland responses to variation of moisture regimes and for projecting consequences of future changes. Electronic supplementary material The online version of this article (10.1007/s00442-020-04625-9) contains supplementary material, which is available to authorized users.

Droughts can affect invertebrate communities in wetlands, which can have bottom-up effects on the condition and survival of top predators. Shorebirds, key predators at coastal wetlands, have experienced widespread population declines and could be negatively affected by droughts. We explored, in detail, the effects of drought on multiple aspects of shorebird stopover and migration ecology by contrasting a year with average wet/dry conditions (2016) with a year with moderate drought (2017) at a major subarctic stopover site on southbound migration. We also examined the effects of drought on shorebird body mass during stopover across 14 years (historical: 1974–1982 and present-day: 2014–2018). For the detailed comparison of two years, in the year with moderate drought we documented lower invertebrate abundance at some sites, higher prey family richness in shorebird faecal samples, lower shorebird refuelling rates, shorter stopover durations for juveniles, and, for most species, a higher probability of making a subsequent stopover in North America after departing the subarctic, compared to the year with average wet/dry conditions. In the 14-year dataset, shorebird body mass tended to be lower in drier years. We show that even short-term, moderate drought conditions can negatively affect shorebird refuelling performance at coastal wetlands, which may carry-over to affect subsequent stopover decisions. Given shorebird population declines and predicted changes in the severity and duration of droughts with climate change, researchers should prioritize a better understanding of how droughts affect shorebird refuelling performance and survival. Supplementary Information The online version contains supplementary material available at 10.1007/s00442-021-05047-x.

Recently, community ecology has emphasized the multi-facetted aspects of biological diversity by linking species traits and the environment. Here, we explored environmental correlates of taxonomically-based and traits-based compositional distances using a comprehensive data set of diatom and macroinvertebrate communities. We also explored the responses of different beta diversity components (i.e., overall beta diversity, turnover, and nestedness) of beta diversity facets (i.e., taxonomically and traits-based beta diversity) to environmental distances. Partial Mantel tests were used to test the relationships between beta diversity and environmental distance (while controlling for spatial distances). Taxonomically-based beta diversity varied much more than traits-based beta diversity, indicating strong functional convergence. We found that taxonomically-based beta diversity was largely driven by the turnover component. However, the nestedness component contributed more to overall traits-based beta diversity than the turnover component. Taxonomically-based beta diversity was significantly correlated with environmental distances for both diatoms and macroinvertebrates. Thus, we found support for the role of environmental filtering as a driver of community dissimilarities of rather different biological groups. However, the strength of these relationships between beta diversity and environmental distances varied depending on the biological group, facet, component, and the way which the environmental variables were selected to calculate the explanatory (distance) matrix. Our results indicated that both taxonomically and traits-based approaches are still needed to better understand patterns and mechanisms affecting the organization of biological communities in streams. This is because different facets of biological communities may be driven by different mechanisms. Electronic supplementary material The online version of this article (10.1007/s00442-019-04535-5) contains supplementary material, which is available to authorized users.

The exotic South African ragwort (Senecio inaequidens DC.) rapidly spread across Central Europe after its introduction, but we still do not know to what extent its timing of arrival in a plant community (i.e. before or after natives) and the composition of the native community being invaded affect (1) its capacity to invade a European grassland, (2) the performance of the native species, and (3) the direction and strength of priority effects. In a greenhouse experiment, we manipulated the timing of arrival of the exotic species (Senecio) and the composition of the native community to test the influence of these factors on the productivity and N content of exotic and native species. We also investigated if the plant species origin (native or exotic) and the native community composition affected the benefit of arriving early and the cost of arriving late in the community. The establishment success of Senecio strongly depended on its timing of arrival in a grassland community. Senecio benefited more from arriving early than did the natives. The presence of legumes in the community did not favour invasion by Senecio. When natives arrived later than Senecio, however, priority effects were weaker when legumes were part of the native community. Our results showed that inhibitory priority effects created by natives can lower the risk of invasion by Senecio. An early arrival of this species at a site with low native species abundance is a scenario that could favour invasion.

The functional composition of plant communities is a critical modulator of climate change impacts on ecosystems, but it is not a simple function of regional climate. In the Arctic tundra, where climate change is proceeding the most rapidly, communities have not shifted their trait composition as predicted by spatial temperature–trait relationships. Important causal pathways are thus missing from models of trait composition change. Here, we study causes of plant community functional variation in an oroarctic tundra landscape in Kilpisjärvi, Finland. We consider the community-weighted means of plant vegetative height, as well as two traits related to the leaf economic spectrum. Specifically, we model their responses to locally measured summer air temperature, snow conditions, and soil resource levels. For each of the traits, we also quantify the importance of intraspecific trait variation (ITV) for between-community functional differences and trait–environment matching. Our study shows that in a tundra landscape (1) snow is the most influential abiotic variable affecting functional composition, (2) vegetation height is under weak local environmental control, whereas leaf economics is under strong local environmental control, (3) the relative magnitude of ITV differs between traits, and (4) ITV is not very consequential for community-level trait–environment relationships. Our analyses highlight the importance of winter conditions for community functional composition in seasonal areas. We show that winter climate change can either amplify or counter the effects summer warming, depending on the trait. Electronic supplementary material The online version of this article (10.1007/s00442-019-04508-8) contains supplementary material, which is available to authorized users.

Plant functional traits and strategies hold the promise to explain species distribution, but few studies have linked multiple traits to multiple niche dimensions (i.e., light, water, and nutrients). Here, we analyzed for 29 liana species in a Chinese tropical seasonal rainforest how: (1) trait associations and trade-offs lead to different plant strategies; and (2) how these traits shape species’ niche dimensions. Eighteen functional traits related to light, water, and nutrient use were measured and species niche dimensions were quantified using species distribution in a 20-ha plot combined with data on canopy gaps, topographic water availability, and soil nutrients. We found a tissue toughness spectrum ranging from soft to hard tissues along which species also varied from acquisitive to conservative water use, and a resource acquisition spectrum ranging from low to high light capture and nutrient use. Intriguingly, each spectrum partly reflected the conservative–acquisitive paradigm, but at the same time, the tissue toughness and the resource acquisition spectrum were uncoupled. Resource niche dimensions were better predicted by individual traits than by multivariate plant strategies. This suggests that trait components that underlie multivariate strategy axes, rather than the plant strategies themselves determine species distributions. Different traits were important for different niche dimensions. In conclusion, plant functional traits and strategies can indeed explain species distributions, but not in a simple and straight forward way. Although the identification of global plant strategies has significantly advanced the field, this research shows that global, multivariate generalizations are difficult to translate to local conditions, as different components of these strategies are important under different local conditions. Supplementary Information The online version contains supplementary material available at 10.1007/s00442-021-04937-4.

Plant communities worldwide show varied responses to nutrient enrichment—including shifts in species identity, decreased diversity, and changes in functional trait composition—but the factors determining community recovery after the cessation of nutrient addition remain uncertain. We manipulated nutrient levels in a tundra community for 6 years of nutrient addition followed by 8 years of recovery. We examined how community recovery was mediated by traits related to plant resource-use strategy and plant ability to modify their environment. Overall, we observed persistent effects of fertilization on plant communities. We found that plants with fast-growing traits, including higher specific leaf area, taller stature and lower foliar C:N, were more likely to show a persistent increase in fertilized plots than control plots, maintaining significantly higher cover in fertilized plots 8 years after cessation of fertilization. Additionally, although graminoids responded most strongly to the initial fertilization treatment, forb species were more vulnerable to fertilization effects in the long-term, showing persistent decline and no recovery in 8 years. Finally, these persistent fertilization effects were accompanied by modified environmental conditions, including persistent increases in litter depth and soil phosphorous and lower soil C:N. Our results demonstrate the potential for lasting effects of nutrient enrichment in nutrient-limited systems and identify species traits related to rapid growth and nutrient-use efficiency as the main predictors of the persistence of nutrient enrichment effects. These findings highlight the usefulness of trait-based approach for understanding the persistent feedbacks of nutrient enrichment, plant dynamics, and niche construction via litter and nutrient build-up. Supplementary Information The online version contains supplementary material available at 10.1007/s00442-021-05064-w.

Ongoing globalisation and climate change are causing plant species to invade new habitats and thereby alter biodiversity and ecosystem functioning. Since numbers of plant invasions continue to increase globally, it is crucial to investigate the effects of multiple co-occurring alien species on native communities. Furthermore, priority effects due to the earlier emergence of certain species affecting fitness of later arriving species can shape community structure and affect native species performance. We investigate in a common garden pot experiment the interactions among five alien-native species pairs. First we focus on the effect of growing with either one or two alien neighbour species on a native plant, second we alter the arrival time of the alien or native neighbour by 3 weeks. Generally, native species performance decreased when surrounded by two alien species compared to only one, although the magnitude of this effect varied depending on species, with one species even performing better with alien neighbours than in monoculture. Species performance greatly decreased when arriving second in the pot, for both native and alien species. In contrast, alien species tended to benefit more from arriving early. Given that we studied annual ruderal species, their potentially lower competitive ability might explain why we detected negative effects of late arrival. We highlight the need to further elucidate underlying mechanisms of small-scale invasion dynamics to achieve generalisations concerning the response of multiple alien and native plants given their species-specific differences in response to neighbour species and arrival time. Supplementary Information The online version contains supplementary material available at 10.1007/s00442-021-05017-3.

Successional processes can be observed for many organisms and resources, but most studies of succession have focused on plants. A general framework has been proposed, advocating that successional patterns in species turnover are predominantly driven by competition, dispersal or abiotic limitation, and that the patterning of species accumulation over time gives clues to which process is most influential in a given system. We applied this framework to succession in communities of wood-living beetles, utilizing ephemeral resources in the form of 60 experimentally created dead aspen high stumps. High stumps were created at sun-exposed sites (high ambient temperature; favourable abiotic conditions) and shaded sites (low ambient temperature; abiotically limiting conditions). The sites were intermixed, ensuring similar dispersal opportunities. Beetle species richness and abundance were monitored with flight interception traps over four consecutive years. Consistent with predictions from the tested framework, several beetle functional groups accumulated species more slowly at the unfavourable shaded sites than at the favourable exposed sites. Species richness at the exposed sites increased rapidly to a plateau, consistent with a limiting effect of competition on community development. Similar results were obtained for beetle abundance and community structure. Part of the variance in beetle community structure was jointly explained by habitat and fungal community composition, suggesting that differences in the composition and developmental rate of fungal communities in the two habitats contributed to the observed patterns. Targeted experimental studies are now required to decisively establish what processes underlie the contrasting successional trajectories in the two environments. Electronic supplementary material The online version of this article (10.1007/s00442-020-04750-5) contains supplementary material, which is available to authorized users.

Despite the importance of predation in many ecosystems, gaps remain in our understanding of nocturnal marine predators. Although the kelp forests of Southern California are some of the most well-studied ecosystems, California morays, Gymnothorax mordax, are predominately nocturnal predators that have remained largely unstudied and their predatory effects on the kelp forest ecosystem are unknown. We use a multi-year data set to examine the dietary breadth of G. mordax and to determine the functional role of this predator. We also quantify bite force to examine the potential performance limitations of morays in exploiting prey. Stomach content analyses and linear selectivity index values indicate that G. mordax specializes on kelp bass, Paralabrax clathratus. Average size of kelp bass consumed varies across years, suggesting that morays respond to fluctuations in prey size availability. The scaling relationship of kelp bass standard length and moray head length reveals an ontogenetic shift, where maximum prey size increases with moray size and small prey are dropped from the diet of larger individuals. Moray bite force exhibited strong positive allometry with moray head size, suggesting that larger morays exhibit greater bite forces for their head and body size. However, we found no relationship between prey size and bite force, suggesting that a disproportional increase in bite force does not facilitate the consumption of disproportionately larger prey. Our results indicate that while G. mordax of Catalina Island is a dietary specialist, it is capable of exhibiting functional shifts in prey size and species based on their abundance. Electronic supplementary material The online version of this article (10.1007/s00442-018-4260-x) contains supplementary material, which is available to authorized users.

To understand the relationship between conservation measures and agricultural yields, we need to know the contributions of organisms to both ecosystem services and disservices. We studied the activity and contribution of birds and mammals to intermediate ecosystem services (predation of weed seeds or invertebrate pests) and disservices (predation of crop seeds or beneficial invertebrates) in southern Sweden between June and November 2016. We measured seed and invertebrate predation rates using trays placed in front of 32 wildlife cameras in 16 cereal fields with a local habitat contrast (8 fields adjacent to another crop field and 8 fields adjacent to a semi-natural grassland) and along a landscape heterogeneity gradient (amount of semi-natural grassland). Both activity and predation were dominated by small mammals (mainly rodents), yet only a few species contributed to predation services and disservices according to camera records. Small mammal activity and predation varied considerably over time. Small mammal activity was significantly higher at trays with crop seeds or beneficial invertebrate prey compared to trays with pest prey, and crop seed predation by small mammals was significantly higher than weed seed predation. In contrast, bird activity and predation did not differ significantly between resource types, but varied over time depending on the habitat contrast. Predation of animal prey by birds was highest after cereal harvest, independent of habitat contrast. Our study highlights that birds and in particular rodents provide important intermediate ecosystem services, but also disservices, which fluctuate strongly in intensity over time. Electronic supplementary material The online version of this article (10.1007/s00442-018-4242-z) contains supplementary material, which is available to authorized users.

In Batesian mimicry, a species lacking defences against predators benefits from mimicking the aposematic signal of a defended species, while the model may incur the costs of reduced defensive efficacy. Similar reciprocal indirect effects may emerge even when the signal is not mimicked; termed associational effects, such interactions are well known in plants sharing herbivores but have received little attention in animal studies. We investigated associational interactions in a system where unequally defended prey (chemically defended Bufo bufo and undefended Rana temporaria tadpoles), sharing general morphology but not an aposematic signal, were exposed to predation by the carp Cyprinus carpio along a gradient of relative prey abundance. In the absence of fish, the assemblage composition had no effect on the survival of Rana, while that of Bufo decreased with increasing abundance of Rana. Fish reduced the survival of tadpoles from both species. However, increased relative abundance of Bufo in the community led to enhanced survival in both Bufo and Rana. Increasing relative proportions of heterospecifics reduced metamorph mass only in Bufo, indicating greater sensitivity to interspecific competition compared to Rana; the effect was reduced in the presence of fish. Our results show that undefended non-mimetic prey enjoy reduced predation with increasing relative abundance of chemically defended prey, which in turn suffer greater mortality with an increasing proportion of the undefended species. Associational resistance/susceptibility, driven by current assemblage composition, not by selection for resemblance, can shape the dynamics of mixed communities of defended and undefended prey in the absence of mimicry.

Soil animals play important roles in ecosystem functioning and stability, but the environmental controls on their communities are not fully understood. In this study, we compiled a dataset of soil animal communities for which the abundance and body mass of multiple soil animal groups were recorded. The mass–abundance scaling relationships were then used to investigate multiple environmental controls on soil animal community composition. The data reveal latitudinal shifts from high abundances of small soil animals at high latitudes to greater relative abundances of large soil animals at low latitudes. A hierarchical linear mixed effects model was applied to reveal the environmental variables shaping these latitudinal trends. The final hierarchical model identified mean annual temperature, soil pH and soil organic carbon content as key environmental controls explaining global mass–abundance scaling relationships in soil animal communities (R2c = 0.828, Ngroup = 117). Such relationships between soil biota with climate and edaphic conditions have been previously identified for soil microbial, but not soil animal, communities at a global scale. More comprehensive global soil community datasets are needed to better understand the generality of these relationships over a broader range of global ecosystems and soil animal groups. Electronic supplementary material The online version of this article (10.1007/s00442-020-04640-w) contains supplementary material, which is available to authorized users.

Biodiversity loss and climate warming are occurring in concert, with potentially profound impacts on ecosystem functioning. We currently know very little about the combined effects of these changes on the links between the community structure, dynamics and the resulting in situ CO2 concentrations in freshwater ecosystems. Here we aimed to determine both individual and combined effects of temperature and non-resource diversity (species inedible for a given consumer) on CO2 concentration. Our analysis further aimed to establish both direct effects on CO2 concentrations and potential indirect effects that occur via changes to the phytoplankton and zooplankton biomasses. Our results showed that there were no interactive effects of changes in temperature and diversity on CO2 concentration in the water. Instead, independent increases in either temperature or non-resource diversity resulted in a substantial reduction in CO2 concentrations, particularly at the highest non-resource diversity. The effects of non-resource diversity and warming on CO2 were indirect, resulting largely from the positive impacts on total biomass of primary producers. Our study is the first to experimentally partition the impacts of temperature and diversity on the consumer–resource dynamics and associated changes to CO2 concentrations. It provides new mechanistic insights into the role of diverse plankton communities for ecosystem functioning and their importance in regulating CO2 dynamics under ongoing climate warming. Electronic supplementary material The online version of this article (10.1007/s00442-020-04593-0) contains supplementary material, which is available to authorized users.

Temporal heterogeneity in nutrient availability is known to increase phytoplankton diversity by allowing more species to coexist under different resource niches. Spatial heterogeneity has also been positively correlated with species diversity. Here we investigated how temporal and spatial differences in nutrient addition together impact biodiversity in metacommunities varying in the degree of connectivity among the patches. We used a microcosm experimental design to test two spatiotemporal ways of supplying nutrients: synchronously (nutrients were added regionally—to all four patches at the same time) and asynchronously (nutrients were added locally—to a different patch each time), combined with two different degrees of connectivity among the patches (low or high connectivity). We used three species of algae and one species of cyanobacteria as the primary producers; and five ciliate and two rotifer species as the grazers. We expected higher diversity in metacommunities receiving an asynchronous nutrient supply, assuming stronger development of heterogeneous patches with this condition rather than with synchronous nutrient supply. This result was expected, however, to be dependent on the degree of connectivity among patches. We found significant effects of nutrient addition in both groups of organisms. Phytoplankton diversity increased until the fourth week (transiently) and zooplankton richness was persistently higher under asynchronous nutrient addition. Our results were consistent with our hypothesis that asynchronicity in nutrient supply would create a more favorable condition for species to co-occur. However, this effect was, in part, transient and was not influenced by the degree of connectivity. Electronic supplementary material The online version of this article (10.1007/s00442-020-04768-9) contains supplementary material, which is available to authorized users.

Ectotherms depend on the environmental temperature for thermoregulation and exploit thermal regimes that optimise physiological functioning. They may also frequent warmer conditions to up-regulate their immune response against parasite infection and/or impede parasite development. This adaptive response, known as ‘behavioural fever’, has been documented in various taxa including insects, reptiles and fish, but only in response to endoparasite infections. Here, a choice chamber experiment was used to investigate the thermal preferences of a tropical freshwater fish, the Trinidadian guppy (Poecilia reticulata), when infected with a common helminth ectoparasite Gyrodactylus turnbulli, in female-only and mixed-sex shoals. The temperature tolerance of G. turnbulli was also investigated by monitoring parasite population trajectories on guppies maintained at a continuous 18, 24 or 32 °C. Regardless of shoal composition, infected fish frequented the 32 °C choice chamber more often than when uninfected, significantly increasing their mean temperature preference. Parasites maintained continuously at 32 °C decreased to extinction within 3 days, whereas mean parasite abundance increased on hosts incubated at 18 and 24 °C. We show for the first time that gyrodactylid-infected fish have a preference for warmer waters and speculate that sick fish exploit the upper thermal tolerances of their parasites to self medicate.

Shrub encroachment induced by global change and human disturbance strongly affects ecosystem structure and function. In this study, we explore the degree to which invading leguminous shrubs affected neighboring grasses, including via the transfer of fixed nitrogen (N). We measured N concentrations and natural abundance 15N of shoot tissues from three dominant grasses from different plant functional groups across seven distances along a local transect (up to 500 cm) to the leguminous shrub, Caragana microphylla. C. microphylla did transfer fixed N to neighboring grasses, but the amount and distance of N transferred were strongly species-specific. Shoot N concentrations decreased significantly with distance from C. microphylla, for a rhizomatous grass, Leymus chinensis, and a bunchgrass, Achnatherum sibiricum. However, N concentrations of another bunchgrass, Stipa grandis, were higher only directly underneath the shrub canopy. Shoot δ15N values of L. chinensis were enriched up to 500 cm from the shrub, but for S. grandis were enriched only below the shrub canopy. In contrast, δ15N of A. sibiricum did not change along the 500-cm transect. Our results indicated the rhizomatous grass transferred fixed N over long distances while bunchgrasses did not. The presence of C. microphylla increased the shoot biomass of L. chinensis but decreased that of S. grandis and A. sibiricum. These findings highlight the potential role of nutrient-acquisition strategies of neighboring grasses in moderating the interspecific variation of fixed N transfer from the leguminous shrub. Overall, leguminous shrubs have either positive or negative effects on the neighboring grasses and dramatically affect plant community composition and structure. Electronic supplementary material The online version of this article (doi:10.1007/s00442-015-3538-5) contains supplementary material, which is available to authorized users.

The scale of resource heterogeneity may influence how resources are locally partitioned between co-existing large and small organisms such as trees and grasses in savannas. Scale-related plant responses may, in turn, influence herbivore use of the vegetation. To examine these scale-dependent bi-trophic interactions, we varied fertilizer [(nitrogen (N)/phosphorus (P)/potassium (K)] applications to patches to create different scales of nutrient patchiness (patch size 2 × 2 m, 10 × 10 m, or whole-plot 50 × 50 m) in a large field experiment in intact African savanna. Within-patch fertilizer concentration and the total fertilizer load per plot were independently varied. We found that fertilization increased the leaf N and P concentrations of trees and grasses, resulting in elevated utilization by browsers and grazers. Herbivory off-take was particularly considerable at higher nutrient concentrations. Scale-dependent effects were weak. The net effect of fertilization and herbivory was that plants in fertilized areas tended to grow less and develop smaller rather than larger standing biomass compared to plants growing in areas that remained unfertilized. When all of these effects were considered together at the community (plot) level, herbivory completely eliminated the positive effects of fertilization on the plant community. While this was true for all scales of fertilization, grasses tended to profit more from coarse-grained fertilization and trees from fine-grained fertilization. We conclude that in herbivore-dominated communities, such as the African savanna, nutrient patchiness results in the herbivore community profiting rather more than the plant community, irrespective of the scale of patchiness. At the community level, the allometric scaling theory’s prediction of plant—and probably also animal—production does not hold or may even be reversed as a result of complex bi-trophic interactions. Electronic supplementary material The online version of this article (doi:10.1007/s00442-016-3627-0) contains supplementary material, which is available to authorized users.

Global climate models predict more frequent periods of drought stress alternated by heavier, but fewer rainfall events in the future. Biodiversity studies have shown that such changed drought stress may be mitigated by plant species richness. Here, we investigate if grassland communities, differing in species richness, respond differently to climatic extremes within the growing season. In a 3-year outdoor mesocosm experiment, four grassland species in both monoculture and mixture were subjected to a rainfall distribution regime with two levels: periods of severe drought in the summer intermitted by extreme rainfall events versus regular rainfall over time. Both treatments received the same amount of water over the season. Extreme rainfall combined with drought periods resulted in a 15% decrease in aboveground biomass in the second and third year, compared to the regular rainfall regime. Root biomass was also reduced in the extreme rainfall treatment, particularly in the top soil layer (− 40%). All species developed higher water use efficiencies (less negative leaf δ13C) in extreme rainfall than in regular rainfall. These responses to the rainfall/drought treatment were independent of species richness, although the mixtures were on an average more productive in terms of biomass than the monocultures. Our experimental results suggest that mixtures are similarly able to buffer these within-season rainfall extremes than monocultures, which contrasts with findings in the studies on natural droughts. Our work demonstrates the importance of investigating the interactions between rainfall distribution and drought periods for understanding effects of climate change on plant community performance. Electronic supplementary material The online version of this article (10.1007/s00442-019-04476-z) contains supplementary material, which is available to authorized users.

Aquatic habitats are often characterized by both high diversity and the threat of multiple anthropogenic stressors. Our research deals with temporal and spatial aspects of two of the main threats for biodiversity, namely eutrophication and fragmentation. It is known that pulsed nutrient addition creates temporal differences in environmental conditions, promoting higher diversity by preventing the best competitor from dominating. Furthermore, a metacommunity landscape with intermediate connectivity increases autotrophs’ diversity and stability. However, it is yet unclear if these two factors are additive in increasing diversity and if the effects extend to the consumer level. With the goal of understanding how eutrophication impacts biodiversity in a metacommunity landscape, we hypothesized that pulsed nutrient addition will increase diversity among both autotrophs and heterotrophs, and this effect will be even greater in a metacommunity landscape. We simulated eutrophication and fragmentation in a microcosm experiment using phytoplankton as primary producers and microzooplankton as grazers. Four treatment combinations were tested including two different landscapes (metacommunity and isolated community) and two forms of nutrient supply (pulsed and continuous): metacommunity/continuous nutrient addition (MC); metacommunity/pulsed nutrient addition (MP); isolated community/continuous nutrient addition (IC); isolated community/pulsed nutrient addition (IP). As expected, pulsed nutrient addition had a persistent positive effect on phytoplankton diversity, with a weaker influence of landscape type. In contrast, the grazer community strongly benefited from a metacommunity landscape, with less significance of pulsed or continuous nutrient addition. Overall, the metacommunity landscape with pulsed nutrient supply supported higher diversity of primary producers and grazers. Electronic supplementary material The online version of this article (10.1007/s00442-018-4319-8) contains supplementary material, which is available to authorized users.

Este artículo contiene 16 páginas, 3 tablas, 2 figuras., We studied community–environment relationships of lake macrophytes at two metacommunity scales using data from 16 regions across the world. More specifically, we examined (a) whether the lake macrophyte communities respond similar to key local environmental factors, major climate variables and lake spatial locations in each of the regions (i.e., within-region approach) and (b) how well can explained variability in the community–environment relationships across multiple lake macrophyte metacommunities be accounted for by elevation range, spatial extent, latitude, longitude, and age of the oldest lake within each metacommunity (i.e., across-region approach). In the within-region approach, we employed partial redundancy analyses together with variation partitioning to investigate the relative importance of local variables, climate variables, and spatial location on lake macrophytes among the study regions. In the across-region approach, we used adjusted R2 values of the variation partitioning to model the community–environment relationships across multiple metacommunities using linear regression and commonality analysis. We found that niche filtering related to local lake-level environmental conditions was the dominant force structuring macrophytes within metacommunities. However, our results also revealed that elevation range associated with climate (increasing temperature amplitude affecting macrophytes) and spatial location (likely due to dispersal limitation) was important for macrophytes based on the findings of the across-metacommunities analysis. These findings suggest that different determinants influence macrophyte metacommunities within different regions, thus showing context dependency. Moreover, our study emphasized that the use of a single metacommunity scale gives incomplete information on the environmental features explaining variation in macrophyte communities., Open access funding provided by University of Oulu including Oulu University Hospital. JA appreciates financial support from the Ella and Georg Ehrnrooth Foundation. BAL was supported by National Research, Development and Innovation Office—NKFIH, OTKA PD120775 Grant and by the Bolyai János Research Scholarship of the Hungarian Academy of Sciences. S.K. was supported by NWO Veni grant 86312012. Sampling of the coastal Brazilian lakes was financed by NWO grant W84-549; The National Geographic Society grant 7864-5; and CNPq grants 480122, 490409, 311427.

Previous studies of Eurasian grasslands have suggested that nutrient ratios, rather than absolute nutrient availabilities and associated productivity, may be driving plant species richness patterns. However, the underlying assumption that species occupy distinct niches along nutrient ratio gradients remains to be tested. We analysed plant community composition and nutrient status of 644 Eurasian wet grassland plots. The importance of nutrient ratios driving variation in species composition was analysed using ordination methods (DCA and CCA). Subsequently, we assessed the niche position and width along the most important nutrient ratio gradient [N:P] for each species. We found that the N:P ratio explained part of the variation in species composition independent from conventional explanatory variables. The N:P ratio explained less variation than soil moisture or pH, but more than productivity or the availability of N and P separately, highlighting its importance for grassland species composition. Species occupied distinct niches along the N:P gradient, and species’ niche widths decreased toward extreme nutrient limitation. After correcting for niche position, there was no overall difference in niche width between endangered and non-endangered species. Surprisingly, endangered species with niche optima at the extreme P-limited end of the gradient had broader niches than their non-endangered counterparts. As species occupied distinct niches along a nutrient ratio gradient, future grassland conservation efforts may benefit from targeting changes in nutrient ratios, i.e. the balance between N and P, rather than only focussing on a general reduction in nutrient availability. However, what management interventions can be used for this purpose remains unclear. Electronic supplementary material The online version of this article (10.1007/s00442-018-4086-6) contains supplementary material, which is available to authorized users.

Savanna grasslands are characterized by high spatial heterogeneity in vegetation structure, aboveground biomass and nutritional quality, with high quality short-grass grazing lawns forming mosaics with patches of tall bunch grasses of lower quality. This heterogeneity can arise because of local differences in consumption, because of differences in productivity, or because both processes enforce each other (more production and consumption). However, the relative importance of both processes in maintaining mosaics of lawn and bunch grassland types has not been measured. Also their interplay been not been assessed across landscape gradients. In a South African savanna, we, therefore, measured the seasonal changes in primary production, nutritional quality and herbivore consumption (amount and percentage) of grazing lawns and adjacent bunch grass patches across a rainfall gradient. We found both higher amounts of primary production and, to a smaller extent, consumption for bunch grass patches. In addition, for bunch grasses primary production increased towards higher rainfall while foliar nitrogen concentrations decreased. Foliar nitrogen concentrations of lawn grasses decreased much less with increasing rainfall. Consequently, large herbivores targeted the biomass produced on grazing lawns with on average 75 % of the produced biomass consumed. We conclude that heterogeneity in vegetation structure in this savanna ecosystem is better explained by small-scale differences in productivity between lawn and bunch grass vegetation types than by local differences in consumption rates. Nevertheless, the high nutritional quality of grazing lawns is highly attractive and, therefore, important for the maintenance of the heterogeneity in species composition (i.e. grazing lawn maintenance). Electronic supplementary material The online version of this article (doi:10.1007/s00442-016-3698-y) contains supplementary material, which is available to authorized users.

Many species have been introduced worldwide into areas outside their natural range. Often these non-native species are introduced without their natural enemies, which sometimes leads to uncontrolled population growth. It is rarely reported that an introduced species provides a new resource for a native species. The rose hips of the Japanese rose, Rosa rugosa, which has been introduced in large parts of Europe, are infested by the native monophagous tephritid fruit fly Rhagoletis alternata. We studied differences in fitness benefits between R. alternata larvae using R. rugosa as well as native Rosa species in the Netherlands. R. alternata pupae were larger and heavier when the larvae fed on rose hips of R. rugosa. Larvae feeding on R. rugosa were parasitized less frequently by parasitic wasps than were larvae feeding on native roses. The differences in parasitization are probably due to morphological differences between the native and non-native rose hips: the hypanthium of a R. rugosa hip is thicker and provides the larvae with the possibility to feed deeper into the hip, meaning that the parasitoids cannot reach them with their ovipositor and the larvae escape parasitization. Our study shows that native species switching to a novel non-native host can experience fitness benefits compared to the original native host.

Effects of temperature changes on phytoplankton communities seem to be highly context-specific, but few studies have analyzed whether this context specificity depends on differences in the abiotic conditions or in species composition between studies. We present an experiment that allows disentangling the contribution of abiotic and biotic differences in shaping the response to two aspects of temperature change: permanent increase of mean temperature versus pulse disturbance in form of a heat wave. We used natural communities from six different sites of a floodplain system as well as artificially mixed communities from laboratory cultures and grew both, artificial and natural communities, in water from the six different floodplain lakes (sites). All 12 contexts (2 communities × 6 sites) were first exposed to three different temperature levels (12, 18, 24 °C, respectively) and afterward to temperature pulses (4 °C increase for 7 h day−1). Temperature-dependent changes in biomass and community composition depended on the initial composition of phytoplankton communities. Abiotic conditions had a major effect on biomass of phytoplankton communities exposed to different temperature conditions, however, the effect of biotic and abiotic conditions together was even more pronounced. Additionally, phytoplankton community responses to pulse temperature effects depended on the warming history. By disentangling abiotic and biotic effects, our study shows that temperature-dependent effects on phytoplankton communities depend on both, biotic and abiotic constraints. Electronic supplementary material The online version of this article (doi:10.1007/s00442-016-3693-3) contains supplementary material, which is available to authorized users.