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Body condition is an important concept in behaviour, evolution and conservation, commonly used as a proxy of an individual's performance, for example in the assessment of environmental impacts. Although body condition potentially encompasses a wide range of health state dimensions (nutritional, immune or hormonal status), in practice most studies operationalize body condition using a single (univariate) measure, such as fat storage. One reason for excluding additional axes of variation may be that multivariate descriptors of body condition impose statistical and analytical challenges. Structural equation modelling (SEM) is used in many fields to study questions relating multidimensional concepts, and we here explain how SEM is a useful analytical tool to describe the multivariate nature of body condition. In this 'Research Methods Guide' paper, we show how SEM can be used to resolve different challenges in analysing the multivariate nature of body condition, such as (a) variable reduction and conceptualization, (b) specifying the relationship of condition to performance metrics, (c) comparing competing causal hypothesis and (d) including many pathways in a single model to avoid stepwise modelling approaches. We illustrated the use of SEM on a real-world case study and provided R-code of worked examples as a learning tool. We compared the predictive power of SEM with conventional statistical approaches that integrate multiple variables into one condition variable: multiple regression and principal component analyses. We show that model performance on our dataset is higher when using SEM and led to more accurate and precise estimates compared to conventional approaches. We encourage researchers to consider SEM as a flexible framework to describe the multivariate nature of body condition and thus understand how it affects biological processes, thereby improving the value of body condition proxies for predicting organismal performance. Finally, we highlight that it can be useful for other multidimensional ecological concepts as well, such as immunocompetence, oxidative stress and environmental conditions.

We describe six datasets that contain GPS and accelerometer data of 202 Eurasian oystercatchers (Haematopusostralegus) spanning the period 2008-2021. Birds were equipped with GPS trackers in breeding and wintering areas in the Netherlands and Belgium. We used GPS trackers from the University of Amsterdam Bird Tracking System (UvA-BiTS) for several study purposes, including the study of space use during the breeding season, habitat use and foraging behaviour in the winter season, and impacts of human disturbance. To enable broader usage, all data have now been made open access. Combined, the datasets contain 6.0 million GPS positions, 164 million acceleration measurements and 7.0 million classified behaviour events (i.e., flying, walking, foraging, preening, and inactive). The datasets are deposited on the research repository Zenodo, but are also accessible on Movebank and as down-sampled occurrence datasets on the Global Biodiversity Information Facility (GBIF) and Ocean Biodiversity Information System (OBIS). ispartof: ZOOKEYS vol:1123 issue:1123 pages:31-45 ispartof: location:Bulgaria status: published

Contains fulltext : 231319.pdf (Publisher’s version ) (Open Access) 21 december 2020

AIM: In many species, density‐dependent effects on reproduction are an important driver of population dynamics. However, it is rarely considered that the direction of density dependence is expected to vary over space and time depending on anti‐predator behaviour and predator community. Aggregation may allow for effective group mobbing against avian nest predators while aggregation may also attract mammalian predators, causing negative density dependence. We aim to quantify spatial variation in the effect of conspecific breeding density on nest survival in a mobbing bird species (Eurasian oystercatcher; Haematopus ostralegus) and identify whether this variation in density dependence can be explained by the predator community. LOCATION: Country‐wide (The Netherlands). METHODS: We integrated reproductive data with breeding territory maps of Eurasian oystercatchers and occupancy maps of avian and mammalian predator species across the Netherlands for a 10‐year period. RESULTS: Spatial variation in the composition of the predator community explained the effects of neighbour density, showing decreasing nest survival when both conspecific density and mammalian dominance increased. Also, heterospecific density (from breeding godwits and lapwing) has an additional effect on the oystercatcher nest survival. Strikingly, this pattern did not extend to mammal‐free island populations. MAIN CONCLUSIONS: Our study provides evidence that both the strength and sign of density dependence can vary spatially within species, implying that it is dangerous to generalize results from a single local population to large‐scale management implications and modelling exercises. The study also suggests that conservation actions that aim to attract breeding birds should be prioritized in areas with fewer mammalian predators, but this idea requires further testing on island populations.

To understand the consequences of anthropogenic and environmental changes for wildlife populations, it is important to study how individuals differ in their sensitivity to environmental change and whether this depends on individual characteristics. An individual's reproductive performance may provide an integrative, unidimensional proxy of an individual's characteristics. In this study, we define an individual's characteristics by three such reproductive states, namely successful, failed and non-breeders in the previous year. We used a 16-year dataset of individually marked breeding Eurasian Oystercatchers Haematopus ostralegus to examine the interannual fluctuations in reproductive success and survival among breeding states, and their state-dependent sensitivity to environmental conditions. Environmental conditions included available biomass of the main prey species of breeding Oystercatchers (Ragworm, Baltic Tellin and Lugworm), tidal height, which reflects one of the main causes of nest loss (flooding), and conditions that may impact the energetic requirements during incubation, such as temperature. We also included environmental variables measured in winter, including available biomass of the main winter prey species (Blue Mussel and Common Cockle) along with factors that may affect food availability and energetic requirements for homeostasis, namely bivalve weight loss, windchill, winter severity and precipitation. Breeding birds that were successful the previous year had higher survival and were more likely to remain successful, in comparison with failed or non-breeders. The effects of environmental conditions acted in the same direction on reproductive success but had opposite effects on survival among the three breeding states, especially for windchill and Blue Mussel biomass. The contrasting state-dependent effects of the environment on survival thus averaged out when examining consequences for lifetime reproductive nest success (LRnS); instead, LRnS was largely influenced by environmental conditions acting upon reproduction. Our study indicates that an individual's previous breeding state provides an integrative measure of heterogeneity in individuals' sensitivity of reproduction and survival to environmental change. Incorporating previous breeding state as a source of individual heterogeneity in population modelling may improve predictions of future population dynamics in a rapidly changing world.

Contains fulltext : 249328.pdf (Publisher’s version ) (Open Access) 25 februari 2022

Many empirical studies motivated by an interest in stable coexistence have quantified negative density dependence, negative frequency dependence, or negative plant-soil feedback, but the links between these empirical results and ecological theory are not straightforward. Here, we relate these analyses to theoretical conditions for stabilisation and stable coexistence in classical competition models. By stabilisation, we mean an excess of intraspecific competition relative to interspecific competition that inherently slows or even prevents competitive exclusion. We show that most, though not all, tests demonstrating negative density dependence, negative frequency dependence, and negative plant-soil feedback constitute sufficient conditions for stabilisation of two-species interactions if applied to data for per capita population growth rates of pairs of species, but none are necessary or sufficient conditions for stable coexistence of two species. Potential inferences are even more limited when communities involve more than two species, and when performance is measured at a single life stage or vital rate. We then discuss two approaches that enable stronger tests for stable coexistence-invasibility experiments and model parameterisation. The model parameterisation approach can be applied to typical density-dependence, frequency-dependence, and plant-soil feedback data sets, and generally enables better links with mechanisms and greater insights, as demonstrated by recent studies.

Evidence suggests that biodiversity supports ecosystem functioning. Yet, the mechanisms driving this relationship remain unclear. Complementarity is one common explanation for these positive biodiversity-ecosystem functioning relationships. Yet, complementarity is often indirectly quantified as overperformance in mixture relative to monoculture (e.g., 'complementarity effect'). This overperformance is then attributed to the intuitive idea of complementarity or, more specifically, to species resource partitioning. Locally, however, several unassociated causes may drive this overperformance. Here, we differentiate complementarity into three types of species differences that may cause enhanced ecosystem functioning in more diverse ecosystems: (i) resource partitioning, (ii) abiotic facilitation, and (iii) biotic feedbacks. We argue that disentangling these three causes is crucial for predicting the response of ecosystems to future biodiversity loss.

The concept of plant-soil feedback is increasingly used to explain plant community assembly processes. Soil nutrient availability can be expected to play a critical role on these processes. However, little is known about the effects of nutrient availability on feedback direction and strength. A plant-soil feedback experiment was performed with the grasses Anthoxanthum odoratum and Festuca rubra, and the forbs Leontodon hispidus and Plantago lanceolata, on soil with either low or high nutrient availability. Additionally, we tested if plant-soil feedback of the two forbs under these conditions changed by inoculation of the soil with spores of an arbuscular mycorrhizal fungus. Increased nutrient availability neutralised plant-soil feedback based on shoot biomass independent of its negative or positive direction, whereas the effects on root biomass were either not altered or turned negative. Mycorrhizal fungi spore addition decreased negative feedback and increased positive feedback. Our results suggest that negative plant-soil feedback on low nutrient soil can be overcome with nutrient addition, and that positive soil biota associations on low nutrient soil may become superfluous with nutrient increase. We hypothesize that species-specific, microbial mediated plant community assembly processes occur in low rather than high nutrient environments.

Reports of declines in biomass of flying insects have alarmed the world in recent years. However, how biomass declines reflect biodiversity loss is still an open question. Here, we analyze the abundance (19,604 individuals) of 162 hoverfly species (Diptera: Syrphidae), at six locations in German nature reserves in 1989 and 2014, and generalize the results with a model varying decline rates of common vs. rare species. We show isometric decline rates between total insect biomass and total hoverfly abundance and a scale-dependent decline in hoverfly species richness, ranging between -23% over the season to -82% at the daily level. We constructed a theoretical null model to explore how strong declines in total abundance translate to changing rank-abundance curves, species persistence, and diversity measures. Observed persistence rates were disproportionately lower than expected for species of intermediate abundance, while the rarest species showed decline and appearance rates consistent with random expectation. Our results suggest that large insect biomass declines are predictive of insect diversity declines. Under current threats, even the more common species are in peril, calling for a reevaluation of hazards and conservation strategies that traditionally target already rare and endangered species only.

Contains fulltext : 245377.pdf (Publisher’s version ) (Closed access) 1 p.

Plant interactions are as important belowground as aboveground. Belowground plant interactions are however inherently difficult to quantify, as roots of different species are difficult to disentangle. Although for a couple of decades molecular techniques have been successfully applied to quantify root abundance, root identification and quantification in multispecies plant communities remains particularly challenging. Here we present a novel methodology, multispecies genotyping by sequencing (msGBS), as a next step to tackle this challenge. First, a multispecies meta‐reference database containing thousands of gDNA clusters per species is created from GBS derived High Throughput Sequencing (HTS) reads. Second, GBS derived HTS reads from multispecies root samples are mapped to this meta‐reference which, after a filter procedure to increase the taxonomic resolution, allows the parallel quantification of multiple species. The msGBS signal of 111 mock‐mixture root samples, with up to 8 plant species per sample, was used to calculate the within‐species abundance. Optional subsequent calibration yielded the across‐species abundance. The within‐ and across‐species abundances highly correlated (R 2 range 0.72–0.94 and 0.85–0.98, respectively) to the biomass‐based species abundance. Compared to a qPCR based method which was previously used to analyse the same set of samples, msGBS provided similar results. Additional data on 11 congener species groups within 105 natural field root samples showed high taxonomic resolution of the method. msGBS is highly scalable in terms of sensitivity and species numbers within samples, which is a major advantage compared to the qPCR method and advances our tools to reveal hidden belowground interactions., see also the Perspective by Josep Piñol

A growing number of studies are providing evidence that a suite of anthropogenic stressors — habitat loss and fragmentation, pollution, invasive species, climate change and overharvesting — are seriously reducing insect and other invertebrate abundance, diversity and biomass across the biosphere. These declines affect all functional groups: herbivores, detritivores, parasitoids, predators and pollinators. Insects are vitally important in a wide range of ecosystem services of which some are vitally important for food production and security (for example, pollination and pest control). […]. info:eu-repo/semantics/publishedVersion

Contains fulltext : 225690.pdf (Publisher’s version ) (Open Access) Abstract Plant-soil legacies consisting of species-specific microbial communities are hypothesized to play a critical, structuring role in plant species coexistence processes. Plant species are thought to perform worse on soil conditioned by the same species compared to soil of other species, which serves as a self-limitation mechanism and averts monodominance of strong competitors. Here, we test in a multispecies community setting, whether root colonization and resource utilization of soil patches with distinct soil legacies are consistent with this hypothesis. We grew eight grassland species together in an outdoor mesocosm set-up in unconditioned soil and created soil patches in these communities conditioned by one of four plant species, or a soil mixture of all four. During two subsequent growing seasons, we tested the effect of these conditioned soil patches on below-ground root colonization into the patches of each surrounding plant species using a novel sequencing-based approach. In addition, we tested the effect of soil conditioning on local root functioning by injecting tracers into the soil patches and measuring the recovery in above-ground biomass. Against expectations, plant species did not place less roots in own soil patches compared to foreign soil patches, nor did species take up less tracer from own compared to foreign soil patches. Using structural equation modelling, we found that tracer uptake of the plant species was to a varying degree explained by root densities in the various soil patches and by differing soil nutrient availability of the soil patches. We conclude that soil legacy effects are inextricably connected to soil nutrient availability, which needs to be taken into account in plant-soil feedback research to understand the processes that shape plant communities. Synthesis. We found that soil legacy effects in complex, multispecies semi-field conditions did not match expectations based on theory and experiments in controlled conditions. Among the many complicating factors that may modify or even overrule soil legacy effects in semi-field settings, we identified soil nutrient availability as a critical force that may, together with soil biota, shape plant species coexistence processes. 13 p.

1. Models of natural processes necessarily sacrifice some realism for the sake of tractability. Detailed, parameter‐rich models often provide accurate estimates of system behaviour but can be data‐hungry and difficult to operationalize. Moreover, complexity increases the danger of ‘over‐fitting’, which leads to poor performance when models are applied to novel conditions. This challenge is typically described in terms of a trade‐off between bias and variance (i.e. low accuracy vs. low precision). 2. In studies of ecological communities, this trade‐off often leads to an argument about the level of detail needed to describe interactions among species. Here, we used data from a grassland biodiversity experiment containing nine locally abundant plant species (the Jena ‘dominance experiment’) to parameterize models representing six increasingly complex hypotheses about interactions. For each model, we calculated goodness‐of‐fit across different subsets of the data based on sown species richness levels, and tested how performance changed depending on whether or not the same data were used to parameterize and test the model (i.e. within vs. out‐of‐sample), and whether the range of diversity treatments being predicted fell inside or outside of the range used for parameterization. 3. As expected, goodness‐of‐fit improved as a function of model complexity for all within‐sample tests. In contrast, the best out‐of‐sample performance generally resulted from models of intermediate complexity (i.e. with only two interaction coefficients per species—an intraspecific effect and a single pooled interspecific effect), especially for predictions that fell outside the range of diversity treatments used for parameterization. In accordance with other studies, our results also demonstrate that commonly used selection methods based on AIC of models fitted to the full dataset correspond more closely to within‐sample than out‐of‐sample performance. 4. Synthesis. Our results demonstrate that models which include only general intra and interspecific interaction coefficients can be sufficient for estimating species‐level abundances across a wide range of contexts and may provide better out‐of‐sample performance than do more complex models. These findings serve as a reminder that simpler models may often provide a better trade‐off between bias and variance in ecological systems, particularly when applying models beyond the conditions used to parameterize them. This paper arose as part of the ‘BEF-Coexist’ workshop, organized by Y.F., C.R., and W.S.H., with funding from the German Research Foundation (RO2397/8). A.T.C. was supported by a ‘catalyst’ postdoctoral fellowship from sDiv. S.S. was supported by the Spanish Government under a Ramón y Cajal contract (RYC-2016- 20604). Funding and support for the Jena Experiment were provided by the German Research Foundation (FOR456/1451), Friedrich Schiller University of Jena, and Max Planck Society.

Contains fulltext : 217065.pdf (Publisher’s version ) (Open Access) Abstract Locally, plant species richness supports many ecosystem functions. Yet, the mechanisms driving these often-positive biodiversity?ecosystem functioning relationships are not well understood. Spatial resource partitioning across vertical resource gradients is one of the main hypothesized causes for enhanced ecosystem functioning in more biodiverse grasslands. Spatial resource partitioning occurs if species differ in where they acquire resources and can happen both above- and belowground. However, studies investigating spatial resource partitioning in grasslands provide inconsistent evidence. We present the results of a meta-analysis of 21 data sets from experimental species-richness gradients in grasslands. We test the hypothesis that increasing spatial resource partitioning along vertical resource gradients enhances ecosystem functioning in diverse grassland plant communities above- and belowground. To test this hypothesis, we asked three questions. (1) Does species richness enhance biomass production or community resource uptake across sites? (2) Is there evidence of spatial resource partitioning as indicated by resource tracer uptake and biomass allocation above- and belowground? (3) Is evidence of spatial resource partitioning correlated with increased biomass production or community resource uptake? Although plant species richness enhanced community nitrogen and potassium uptake and biomass production above- and belowground, we found that plant communities did not meet our criteria for spatial resource partitioning, though they did invest in significantly more aboveground biomass in higher canopy layers in mixture relative to monoculture. Furthermore, the extent of spatial resource partitioning across studies was not positively correlated with either biomass production or community resource uptake. Our results suggest that spatial resource partitioning across vertical resource gradients alone does not offer a general explanation for enhanced ecosystem functioning in more diverse temperate grasslands.

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Questions: The rapid climate warming in tundra ecosystems can increase nutrient availability in the soil, which may initiate shifts in vegetation composition. The direction in which the vegetation shifts will co-determine whether Arctic warming is mitigated or accelerated, making the understanding of successional trajectories urgent. One of the key factors influencing the competitive relationships between plant species is their access to nutrients, depending on the depth where they take up most nutrients. However, nutrient uptake at different soil depths by tundra plant species that differ in rooting depth is unclear. Location: Kytalyk Nature Reserve, northeast Siberia, Russia. Methods: We injected 15N to 5 cm, 15 cm and the thaw front of the soil in a moist tussock tundra. The absorption of 15N by grasses, sedges, deciduous shrubs and evergreen shrubs from the three depths was compared. Results: The results clearly show a vertical differentiation of N uptake by these plant functional types, corresponding to their rooting strategy. Shallow-rooting dwarf shrubs were more capable of absorbing nutrients from the upper soil than from deeper soil. Deep-rooting grasses and sedges were more capable of absorbing nutrients from deeper soil than the dwarf shrubs. The natural 15N abundances in control plants also indicate that graminoids can absorb more nutrients from the deeper soil than dwarf shrubs. Conclusions: Our results show that graminoids and shrubs in the Arctic differ in their N uptake strategies, with graminoids profiting from nutrients released at the thaw front, while shrubs mainly forage in upper soil layers. Our results suggest that tundra vegetation will become graminoid-dominated as permafrost thaw progresses and nutrient availability increases in the deep soil.

Summary There is consensus that plant species richness enhances plant productivity within natural grasslands, but the underlying drivers remain debated. Recently, differential accumulation of soil‐borne fungal pathogens across the plant diversity gradient has been proposed as a cause of this pattern. However, the below‐ground environment has generally been treated as a ‘black box’ in biodiversity experiments, leaving these fungi unidentified.Using next generation sequencing and pathogenicity assays, we analysed the community composition of root‐associated fungi from a biodiversity experiment to examine if evidence exists for host specificity and negative density dependence in the interplay between soil‐borne fungi, plant diversity and productivity.Plant species were colonised by distinct (pathogenic) fungal communities and isolated fungal species showed negative, species‐specific effects on plant growth. Moreover, 57% of the pathogenic fungal operational taxonomic units (OTUs) recorded in plant monocultures were not detected in eight plant species plots, suggesting a loss of pathogenic OTUs with plant diversity.Our work provides strong evidence for host specificity and negative density‐dependent effects of root‐associated fungi on plant species in grasslands. Our work substantiates the hypothesis that fungal root pathogens are an important driver of biodiversity‐ecosystem functioning relationships.

Below-ground resource partitioning is among the most prominent hypotheses for driving the positive biodiversity-ecosystem function relationship. However, experimental tests of this hypothesis in biodiversity experiments are scarce, and the available evidence is not consistent. We tested the hypothesis that resource partitioning in space, in time or in both space and time combined drives the positive effect of diversity on both plant productivity and total community resource uptake. At the community level, we predicted that total community resource uptake and biomass production above- and below-ground will increase with increased species richness or functional group richness. We predicted that, at the species level, resource partition breadth will become narrower, and that overlap between the resource partitions of different species will become smaller with increasing species richness or functional group richness. We applied multiple resource tracers (Li and Rb as potassium analogues, the water isotopologues-H2 18O and 2H2O, and 15N) in three seasons at two depths across a species and functional group richness gradient at a grassland biodiversity experiment. We used this multidimensional resource tracer study to test if plant species partition resources with increasing plant diversity across space, time or both simultaneously. At the community level, total community resource uptake of nitrogen and potassium and above- and below-ground biomass increased significantly with increasing species richness but not with increasing functional group richness. However, we found no evidence that resource partition breadth or resource partition overlap decreased with increasing species richness for any resource in space, time or both space and time combined. Synthesis. These findings indicate that below-ground resource partitioning may not drive the enhanced resource uptake or biomass production found here. Instead, other mechanisms such as facilitation, species-specific biotic feedback or above-ground resource partitioning are likely necessary for enhanced overall ecosystem function.

Summary Plant diversity enhances many ecosystem functions, including root biomass production, which drives soil carbon input. Although root decomposition accounts for a large proportion of carbon input for soil, little is known about plant diversity effect on this process. Plant diversity may affect root decomposition in two non-exclusive ways: by providing roots of different substrate quality (e.g. root chemistry) and/or by altering the soil environment (e.g. microclimate). To disentangle these two pathways, we conducted three decomposition experiments using a litter-bag approach in a grassland biodiversity experiment. We hypothesized that: (i) plant species richness negatively affects substrate quality (indicated by increased C:N ratios), which we tested by decomposing roots collected from each experimental plot in one common plot; (ii) plant species richness positively affects soil environment (indicated by increased soil water content), which we tested by decomposing standardized roots in all experimental plots; (iii) the overall effect of plant species richness on root decomposition, due to the contrast between quality and environmental effects, is neutral, which we tested by decomposing community roots in their ‘home’ plots. Plant species richness negatively affected root decomposition in all three experiments. The negative effect of plant species richness on substrate quality was largely explained by increased root C:N ratios along the diversity gradient. Functional group presence explained more variance in substrate quality than species richness. Here, the presence of grasses negatively affected substrate quality and root C:N ratios, while the presence of legumes and small herbs had positive effects. Plant species richness had a negative effect on soil environment despite its positive effect on soil water content which is known to stimulate decomposition. We argue that – instead of soil water content – a combined effect of soil temperature and seasonality might drive environmental effect of plant diversity on decomposition in our plant communities, but this remains to be tested. Synthesis. Our results demonstrate that both substrate quality and soil environment contribute to the net negative effect of plant diversity on root decomposition. This study promotes our mechanistic understanding of increased soil carbon accumulation in more diverse grassland plant communities.

Global declines in insects have sparked wide interest among scientists, politicians, and the general public. Loss of insect diversity and abundance is expected to provoke cascading effects on food webs and to jeopardize ecosystem services. Our understanding of the extent and underlying causes of this decline is based on the abundance of single species or taxonomic groups only, rather than changes in insect biomass which is more relevant for ecological functioning. Here, we used a standardized protocol to measure total insect biomass using Malaise traps, deployed over 27 years in 63 nature protection areas in Germany (96 unique location-year combinations) to infer on the status and trend of local entomofauna. Our analysis estimates a seasonal decline of 76%, and mid-summer decline of 82% in flying insect biomass over the 27 years of study. We show that this decline is apparent regardless of habitat type, while changes in weather, land use, and habitat characteristics cannot explain this overall decline. This yet unrecognized loss of insect biomass must be taken into account in evaluating declines in abundance of species depending on insects as a food source, and ecosystem functioning in the European landscape.

Grass-clover mixtures show many benefits for sustainable agriculture. In the Netherlands, organic farmers often work together in a so-called partner farm concept, with the aim to close nutrient cycles on a regional level. In this system, arable farms grow one-year grass-clover leys, as fodder for a livestock farm, in exchange for, e.g., manure. This practice could also be used in the transition of conventional farms towards a more circular regenerative and nature inclusive agriculture. In the current experiment we assessed the effect of a range of grass (Lolium perenne: Lp, Lolium multiflorum: Lm) and clover (Trifolium pratense: Tp and Trifolium repens: Tr) monocultures and mixtures on both below- and aboveground parameters in light of benefits for livestock and arable farms, and biodiversity. The grass monocultures showed good weed suppression, high root density, and especially Lp had a positive effect on soil structure. Clover, on the other hand, showed high herbage dry matter yield (particularly Tp) and Nitrogen (N) yield, and Tr showed high digestibility. Moreover, clover had a positive effect on the soil mineral N, and earthworm abundance tended to be higher in the clover monocultures. When (some of) the four species were combined in grass-clover mixtures, they combined the positive effects of the species and often even outperformed the (best) monocultures. We concluded that grass-clover mixtures increased agro-ecological functions.

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.

Population growth in passerine birds is largely driven by fecundity. If fecundity is affected, for instance by hatching failure, populations may decline. We noted high hatching failure of up to 27% per year in relict populations of the Northern wheatear (Oenanthe oenanthe) in The Netherlands, a strongly declining, migratory passerine in Europe. This hatching failure itself can cause population decline, irrespective of other adverse factors. Additionally, we investigated the cause of hatching failure. Unhatched eggs showed egg yolk infections or embryonic malformations, part of which is associated with the actions of dioxin-like compounds (DLCs). Indeed, DLCs appear to bioaccumulate in the local foodweb, where the soil contained only background concentrations, similar to those found at many other locations. DLC concentrations in Dutch eggs were six-fold higher than those in a reference population in Sweden, where egg failure was only 6%. However, Northern wheatears appear to be only moderately sensitive to the actions of DLCs, because of their specific Ah-receptor type which may moderate the receptor mediated effects of DLCs. This indicates that the concentrations of DLCs, although elevated, may not have caused the embryo malformations or the low hatching rates. We discuss whether other toxins may be important or imbalances in the nutrition and if inbreeding may play a larger role than expected. (C) 2018 Elsevier B.V. All rights reserved.

The datasets were collected with funding from the National Science Foundation (DEB 0453445; 0453665; 0613666, 0845071, 1019436 & 1558093), Netherlands Organization for Scientific Research (NWO-ALW 801-01-009), the Smithsonian Tropical Research Institute, the Centre for Tropical Forest Science, the John D. and Catherine T. MacArthur Foundation, the Mellon Foundation and the Small World Institute Fund.

This study was supported by the Netherlands Organization for Scientific Research (NWO-ALW 801-01-009; M.D.V.), the Smithsonian Tropical Research Institute (M.D.V.) and the HSBC Climate Partnership (H.C.M.). The datasets were collected with funding from the National Science Foundation (DEB 0425651 & 0948585 to S.P.H.; 0453445 to H.C.M.; 0453665 to S.J.W.; 0613666, 0845071, 1019436 & 1558093 to S.A.S., and 1242622 & 1464389 to L.S.C.), the Smithsonian Tropical Research Institute, the Centre for Tropical Forest Science, the John D. and Catherine T. MacArthur Foundation, the Mellon Foundation and the Small World Institute Fund.

Plant species richness (PSR) increases nutrient uptake which depletes bioavailable nutrient pools in soil. No such relationship between plant uptake and availability in soil was found for phosphorus (P). We explored PSR effects on P mobilisation [phosphatase activity (PA)] in soil. PA increased with PSR. The positive PSR effect was not solely due to an increase in Corg concentrations because PSR remained significant if related to PA:Corg . An increase in PA per unit Corg increases the probability of the temporal and spatial match between substrate, enzyme and microorganism potentially serving as an adaption to competition. Carbon use efficiency of microorganisms (Cmic :Corg ) increased with increasing PSR while enzyme exudation efficiency (PA:Cmic ) remained constant. These findings suggest the need for efficient C rather than P cycling underlying the relationship between PSR and PA. Our results indicate that the coupling between C and P cycling in soil becomes tighter with increasing PSR.

Flooding is a compound stress, imposing strong limitations on plant development. The expression of adaptive traits that alleviate flooding stress may be constrained if floodwater levels are too deep. For instance, adventitious root outgrowth is typically less profound in completely submerged plants than in partially submerged plants, suggesting additional constraints in full submergence. As both oxygen and carbohydrates are typically limited resources under submergence, we tested the effects of oxygen concentration in the floodwater and carbohydrate status of the plants on flooding-induced adventitious root formation in Solanum dulcamara L. Partially submerged plants continued to form adventitious roots in low-oxygen floodwater, whereas completely submerged plants developed hardly any roots, even in floodwater with twice the ambient oxygen concentration. This suggests that contact with the atmosphere, enabling internal aeration, is much more important to optimal adventitious root formation than floodwater oxygen concentrations. If plants were depleted of carbohydrates before flooding, adventitious root formation in partial submergence was poor, unless high light was provided. Thus, either stored or newly produced carbohydrates can fuel adventitious root formation. These results imply that the impact of an environmental stress factor like flooding on plant performance may strongly depend on the interplay with other environmental factors.

Background and aims Temporal flooding is a common environmental stress for terrestrial plants. Aquatic adventitious roots (aquatic roots) are commonly formed in flooding-tolerant plant species and are generally assumed to be beneficial for plant growth by supporting water and nutrient uptake during partial flooding. However, the actual contribution of these roots to plant performance under flooding has hardly been quantified. As the investment into aquatic root development in terms of carbohydrates may be costly, these costs may - depending on the specific environmental conditions - offset the beneficial effects of aquatic roots. This study tested the hypothesis that the balance between potential costs and benefits depends on the duration of flooding, as the benefits are expected to outweigh the costs in long-term but not in short-term flooding. Methods The contribution of aquatic roots to plant performance was tested in Solanum dulcamara during 1-4 weeks of partial submergence and by experimentally manipulating root production. Nutrient uptake by aquatic roots, transpiration and photosynthesis were measured in plants differing in aquatic root development to assess the specific function of these roots. Key results As predicted, flooded plants benefited from the presence of aquatic roots. The results showed that this was probably due to the contribution of roots to resource uptake. However, these beneficial effects were only present in long-term but not in short-term flooding. This relationship could be explained by the correlation between nutrient uptake and the flooding duration-dependent size of the aquatic root system. Conclusions The results indicate that aquatic root formation is likely to be selected for in habitats characterized by long-term flooding. This study also revealed only limited costs associated with adventitious root formation, which may explain the maintenance of the ability to produce aquatic roots in habitats characterized by very rare or short flooding events.

Flooding is expected to increase in frequency and severity in the future. The ecological consequences of flooding are the combined result of species-specific plant traits and ecological context. However, the majority of past flooding research has focused on individual model species under highly controlled conditions. An early summer flooding event in a grassland biodiversity experiment in Jena, Germany, provided the opportunity to assess flooding responses of 60 grassland species in monocultures and 16-species mixtures. We examined plant biomass, species-specific traits (plant height, specific leaf area (SLA), root aerenchyma, starch content) and soil porosity. We found that, on average, plant species were less negatively affected by the flood when grown in higher-diversity plots in July 2013. By September 2013, grasses were unaffected by the flood regardless of plant diversity, and legumes were severely negatively affected regardless of plant diversity. Plants with greater SLA and more root aerenchyma performed better in September. Soil porosity was higher in higher-diversity plots and had a positive effect on plant performance. As floods become more frequent and severe in the future, growing flood-sensitive plants in higher-diversity communities and in soil with greater soil aeration may attenuate the most negative effects of flooding.

Contains fulltext : 176862.pdf (Publisher’s version ) (Closed access) Plant diversity influences many ecosystem functions including root decomposition. However, due to the presence of multiple pathways via which plant diversity may affect root decomposition, our mechanistic understanding of their relationships is limited. In a grassland biodiversity experiment, we simultaneously assessed the effects of three pathways—root litter quality, soil biota, and soil abiotic conditions—on the relationships between plant diversity (in terms of species richness and the presence/absence of grasses and legumes) and root decomposition using structural equation modeling. Our final structural equation model explained 70% of the variation in root mass loss. However, different measures of plant diversity included in our model operated via different pathways to alter root mass loss. Plant species richness had a negative effect on root mass loss. This was partially due to increased Oribatida abundance, but was weakened by enhanced root potassium (K) concentration in more diverse mixtures. Equally, grass presence negatively affected root mass loss. This effect of grasses was mostly mediated via increased root lignin concentration and supported via increased Oribatida abundance and decreased root K concentration. In contrast, legume presence showed a net positive effect on root mass loss via decreased root lignin concentration and increased root magnesium concentration, both of which led to enhanced root mass loss. Overall, the different measures of plant diversity had contrasting effects on root decomposition. Furthermore, we found that root chemistry and soil biota but not root morphology or soil abiotic conditions mediated these effects of plant diversity on root decomposition. 13 p.

In the past two decades, a large number of studies have investigated the relationship between biodiversity and ecosystem functioning, most of which focussed on a limited set of ecosystem variables. The Jena Experiment was set up in 2002 to investigate the effects of plant diversity on element cycling and trophic interactions, using a multi-disciplinary approach. Here, we review the results of 15 years of research in the Jena Experiment, focussing on the effects of manipulating plant species richness and plant functional richness. With more than 85,000 measures taken from the plant diversity plots, the Jena Experiment has allowed answering fundamental questions important for functional biodiversity research.[br/] First, the question was how general the effect of plant species richness is, regarding the many different processes that take place in an ecosystem. About 45% of different types of ecosystem processes measured in the 'main experiment', where plant species richness ranged from 1 to 60 species, were significantly affected by plant species richness, providing strong support for the view that biodiversity is a significant driver of ecosystem functioning. Many measures were not saturating at the 60-species level, but increased linearly with the logarithm of species richness. There was, however, great variability in the strength of response among different processes. One striking pattern was that many processes, in particular belowground processes, took several years to respond to the manipulation of plant species richness, showing that biodiversity experiments have to be long-term, to distinguish trends from transitory patterns. In addition, the results from the Jena Experiment provide further evidence that diversity begets stability, for example stability against invasion of plant species, but unexpectedly some results also suggested the opposite, e.g. when plant communities experience severe perturbations or elevated resource availability. This highlights the need to revisit diversity-stability theory.[br/] Second, we explored whether individual plant species or individual plant functional groups, or biodiversity itself is more important for ecosystem functioning, in particular biomass production. We found strong effects of individual species and plant functional groups on biomass production, yet these effects mostly occurred in addition to, but not instead of, effects of plant species richness.[br/] Third, the Jena Experiment assessed the effect of diversity on multitrophic interactions. The diversity of most organisms responded positively to increases in plant species richness, and the effect was stronger for above-than for belowground organisms, and stronger for herbivores than for carnivores or detritivores. Thus, diversity begets diversity. In addition, the effect on organismic diversity was stronger than the effect on species abundances.[br/] Fourth, the Jena Experiment aimed to assess the effect of diversity on N, P and C cycling and the water balance of the plots, separating between element input into the ecosystem, element turnover, element stocks, and output from the ecosystem. While inputs were generally less affected by plant species richness, measures of element stocks, turnover and output were often positively affected by plant diversity, e.g. carbon storage strongly increased with increasing plant species richness. Variables of the N cycle responded less strongly to plant species richness than variables of the C cycle.[br/] Fifth, plant traits are often used to unravel mechanisms underlying the biodiversity-ecosystem functioning relationship. In the Jena Experiment, most investigated plant traits, both above-and belowground, were plastic and trait expression depended on plant diversity in a complex way, suggesting limitation to using database traits for linking plant traits to particular functions.[br/] Sixth, plant diversity effects on ecosystem processes are often caused by plant diversity effects on species interactions. Analyses in the Jena Experiment including structural equation modelling suggest complex interactions that changed with diversity, e.g. soil carbon storage and greenhouse gas emission were affected by changes in the composition and activity of the belowground microbial community. Manipulation experiments, in which particular organisms, e.g. belowground invertebrates, were excluded from plots in split-plot experiments, supported the important role of the biotic component for element and water fluxes.[br/] Seventh, the Jena Experiment aimed to put the results into the context of agricultural practices in managed grasslands. The effect of increasing plant species richness from 1 to 16 species on plant biomass was, in absolute terms, as strong as the effect of a more intensive grassland management, using fertiliser and increasing mowing frequency. Potential bioenergy production from high-diversity plots was similar to that of conventionally used energy crops. These results suggest that diverse 'High Nature Value Grasslands' are multifunctional and can deliver a range of ecosystem services including production-related services.[br/] A final task was to assess the importance of potential artefacts in biodiversity-ecosystem functioning relationships, caused by the weeding of the plant community to maintain plant species composition. While the effort ( in hours) needed to weed a plot was often negatively related to plant species richness, species richness still affected the majority of ecosystem variables. Weeding also did not negatively affect monoculture performance; rather, monocultures deteriorated over time for a number of biological reasons, as shown in plant-soil feedback experiments.[br/] To summarize, the Jena Experiment has allowed for a comprehensive analysis of the functional role of biodiversity in an ecosystem. A main challenge for future biodiversity research is to increase our mechanistic understanding of why the magnitude of biodiversity effects differs among processes and contexts. It is likely that there will be no simple answer. For example, among the multitude of mechanisms suggested to underlie the positive plant species richness effect on biomass, some have received limited support in the Jena Experiment, such as vertical root niche partitioning. However, others could not be rejected in targeted analyses. Thus, from the current results in the Jena Experiment, it seems likely that the positive biodiversity effect results from several mechanisms acting simultaneously in more diverse communities, such as reduced pathogen attack, the presence of more plant growth promoting organisms, less seed limitation, and increased trait differences leading to complementarity in resource uptake. Distinguishing between different mechanisms requires careful testing of competing hypotheses. Biodiversity research has matured such that predictive approaches testing particular mechanisms are now possible.

Dioecy has a demographic disadvantage compared with hermaphroditism: only about half of reproductive adults produce seeds. Dioecious species must therefore have fitness advantages to compensate for this cost through increased survival, growth, and/or reproduction. We used a full life cycle approach to quantify the demographic costs and benefits associated with dioecy while controlling for demographic differences between dioecious and hermaphroditic species related to other functional traits. The advantage of this novel approach is that we can focus on the effect of breeding system across a diverse tree community. We built a composite integral projection model for hermaphroditic and dioecious tree populations from Barro Colorado Island, Panama, using long-term demographic and newly collected reproductive data. Integration of all costs and benefits showed that compensation was realized through increased seed production, resulting in no net costs of dioecy. Compensation was also facilitated by the low contribution of reproduction to population growth. Estimated positive effects of dioecy on tree growth and survival were small and insignificant for population growth rates. Our model revealed that, for long-lived organisms, the cost of having males is smaller than generally expected. Hence, little compensation is required for dioecious species to maintain population growth rates similar to those of hermaphroditic species.

Summary In contrasting habitats, locally adapted populations are expected to evolve through directional selection. Hydrological gradients provide a scenario where strong selection forces have led to species segregation in communities along the hydrological niche axes due to specific adaptations to their ambient environment. However, much less is known about the processes enabling species with wide ecological amplitudes, such as Solanum dulcamara L., to occupy the contrasting niches at the far ends of the gradient. Here, we investigate the question whether local adaptation is indeed the driving mechanism enabling this species to inhabit contrasting wet and dry habitats. Using molecular analyses and performing a common garden experiment, we examined whether genetic and phenotypic differentiation had occurred between populations originating from flooding- and drought-prone habitat pairs from nine different locations. We also tested whether plants were better adapted to their home site stress. We found a significant genetic variation but hardly any phenotypic differentiation between populations from contrasting habitats for plants grown under optimal conditions. Surprisingly, comparison of the genetic differentiation in neutral markers with that in phenotypic traits using the QST-FST (estimated by ФPT) approach revealed that the QST values were consistently and significantly smaller than the ФPT values. This is in contrast to the hypothesis that directional selection is an important component enabling S. dulcamara to survive under contrasting habitat conditions. In addition, no home site effects were detected for responses to flooding and drought treatments. All plants were characterized by high levels of plasticity for traits associated with flooding and drought tolerance, such as rapid adventitious root formation upon flooding and increased root to shoot ratio in response to drought, irrespective of habitat of origin. Moreover, no trade-offs in such flooding and drought responses were found. Synthesis. Our results show that S. dulcamara has not evolved locally adapted populations in response to flooding and drought stress, despite genetic differentiation and despite the presumably strong selection gradient. The generally high levels of adaptive plasticity in traits increasing flooding and drought tolerance may be the main mechanism allowing S. dulcamara to occupy hydrologically contrasting habitats.

The identification of patterns in life-history strategies across the tree of life is essential to our prediction of population persistence, extinction, and diversification. Plants exhibit a wide range of patterns of longevity, growth, and reproduction, but the general determinants of this enormous variation in life history are poorly understood. We use demographic data from 418 plant species in the wild, from annual herbs to supercentennial trees, to examine how growth form, habitat, and phylogenetic relationships structure plant life histories and to develop a framework to predict population performance. We show that 55% of the variation in plant life-history strategies is adequately characterized using two independent axes: the fast-slow continuum, including fast-growing, short-lived plant species at one end and slow-growing, long-lived species at the other, and a reproductive strategy axis, with highly reproductive, iteroparous species at one extreme and poorly reproductive, semelparous plants with frequent shrinkage at the other. Our findings remain consistent across major habitats and are minimally affected by plant growth form and phylogenetic ancestry, suggesting that the relative independence of the fast-slow and reproduction strategy axes is general in the plant kingdom. Our findings have similarities with how life-history strategies are structured in mammals, birds, and reptiles. The position of plant species populations in the 2D space produced by both axes predicts their rate of recovery from disturbances and population growth rate. This life-history framework may complement trait-based frameworks on leaf and wood economics; together these frameworks may allow prediction of responses of plants to anthropogenic disturbances and changing environments.

Background and aims: Competition is an important force shaping plant communities. Here we test the hypothesis that high overall root length density and selective root placement in nutrient patches, as two alternative strategies, confer competitive advantage in species mixtures. Methods: We performed a full-factorial pairwise competition experiment with eight grassland species in soil with homogeneously distributed nutrients, or with nutrients concentrated in a single patch. We measured species-specific relative growth rate, root length density, selective root placement, and ion uptake rates of all species in monocultures and in mixtures. Results: Grasses showed higher specific root length overall and forbs a higher selective root placement in the nutrient patch. However, relative growth rate and root length density were more strongly related to competitive ability (measured as relative yield per plant), with little distinction between grasses and forbs. Conclusions: Our results suggest that short-term competitive success was related to fast growth and high root densities, irrespective of nutrient heterogeneity. Developing a large root mass quickly may overwhelm the importance of other traits in the establishment phase of plants, although these other traits may prove to be important in the long run.

Summary •Plants may experience different environmental cues throughout their development which interact in determining their phenotype. This paper tests the hypothesis that environmental conditions experienced early during ontogeny affect the phenotypic response to subsequent environmental cues. •This hypothesis was tested by exposing different accessions of Rumex palustris to different light and nutrient conditions, followed by subsequent complete submergence. •Final leaf length and submergence-induced plasticity were affected by the environmental conditions experienced at early developmental stages. In developmentally older leaves, submergence-induced elongation was lower in plants previously subjected to high-light conditions. Submergence-induced elongation of developmentally younger leaves, however, was larger when pregrown in high light. High-light and low-nutrient conditions led to an increase of nonstructural carbohydrates in the plants. There was a positive correlation between submergence-induced leaf elongation and carbohydrate concentration and content in roots and shoots, but not with root and shoot biomass before submergence. •These results show that conditions experienced by young plants modulate the responses to subsequent environmental conditions, in both magnitude and direction. Internal resource status interacts with cues perceived at different developmental stages in determining plastic responses to the environment.

In savannas, the tree-grass balance is governed by water, nutrients, fire and herbivory, and their interactions. We studied the hypothesis that herbivores indirectly affect vegetation structure by changing the availability of soil nutrients, which, in turn, alters the competition between trees and grasses. Nine abandoned livestock holding-pen areas (kraals), enriched by dung and urine, were contrasted with nearby control sites in a semi-arid savanna. About 40 years after abandonment, kraal sites still showed high soil concentrations of inorganic N, extractable P, K, Ca and Mg compared to controls. Kraals also had a high plant production potential and offered high quality forage. The intense grazing and high herbivore dung and urine deposition rates in kraals fit the accelerated nutrient cycling model described for fertile systems elsewhere. Data of a concurrent experiment also showed that bush-cleared patches resulted in an increase in impala dung deposition, probably because impala preferred open sites to avoid predation. Kraal sites had very low tree densities compared to control sites, thus the high impala dung deposition rates here may be in part driven by the open structure of kraal sites, which may explain the persistence of nutrients in kraals. Experiments indicated that tree seedlings were increasingly constrained when competing with grasses under fertile conditions, which might explain the low tree recruitment observed in kraals. In conclusion, large herbivores may indirectly keep existing nutrient hotspots such as abandoned kraals structurally open by maintaining a high local soil fertility, which, in turn, constrains woody recruitment in a negative feedback loop. The maintenance of nutrient hotspots such as abandoned kraals by herbivores contributes to the structural heterogeneity of nutrient-poor savanna vegetation.

• Plants can respond to their environment by morphological plasticity. Generally, the potential benefits of adaptive plastic responses are beyond doubt under predictable environmental changes. However, the net benefits may be less straightforward when plants encounter temporal stresses, such as flooding in river flood plains. • Here, we tested whether the balance of costs and benefits associated with flooding-induced shoot elongation depends on the flooding regime, by subjecting Rumex palustris plants with different elongation capacity to submergence of different frequency and duration. • Our results showed that reaching the surface by shoot elongation is associated with fitness benefits, as under less frequent, but longer, flooding episodes plants emerging above the floodwater had greater biomass production than plants that were kept below the surface. As we predicted, slow-elongating plants had clear advantages over fast-elongating ones if submergence was frequent but of short duration, indicating that elongation also incurs costs. • Our data suggest that high costs select for weak plasticity under frequent environmental change. In contrast to our predictions, however, fast-elongating plants did not have an overall advantage over slow-elongating plants when floods lasted longer. This indicates that the delicate balance between benefits and costs of flooding-induced elongation depends on the specific characteristics of the flooding regime.

Summary 1. Life table response experiment (LTRE) analyses have been widely used to examine the sources of differences in the long-term deterministic growth rate (r =l ogk) of stage-structured populations that live in spatially distinct habitats or under distinct experimental conditions. However, existing methods for LTRE analysis ignore the fact that persistent temporal variation in matrix elements results in a long-term stochastic growth rate (a =l ogks) that is different from the deterministic growth rate (r) and thus do not take into account environmental stochasticity. 2. Here, we develop a stochastic extension of LTRE methods that can be used to compare stochastic growth rates among populations that differ in the observed variability of their matrix elements over time. We illustrate our method with actual data and explore a range of questions that may be addressed with these new tools. Specifically, we investigate how variability in weather conditions affected the population dynamics of the short-lived perennial plant species Anthyllis vulneraria and examine how differences in stochastic growth rates (a) are determined by contributions of mean matrix elements and variability in matrix elements. 3. We find that, consistent with the life history of the species, differences in mean fertility and growth made the largest contribution to differences in a, whereas in terms of variability fertility made the largest contribution in most populations. However, we also find that in all populations, the magnitude of the total contribution of mean matrix elements outweighed that of variability. Finally, increasing soil depth significantly lowered contributions of variability in matrix elements, but it was not related to contributions of differences in mean matrix elements. 4. Synthesis. Stochastic life table response experiment analysis described here provides the first systematic way of incorporating observed differences in temporal variability into the comparison of natural populations. A key finding from this study is that populations occurring on relatively deeper soils were better buffered against climatic variation than populations occurring on shallow soils. We expect this new approach to analyse temporal variability to prove especially useful in the analysis of natural populations experiencing environmental change.

Summary 1. When range shifts or invasions of plant species are studied, it is important to know whether large-scale spatial variation in a species’ demography can be ignored or approximated by variation observed over smaller spatial scales. 2. Here, we studied the population dynamics of three similar (as shown by elasticity analysis) shortlived perennial plant species in multiple sites in different European countries over 2 years. We constructed a total of 40 transition matrices and analysed the spatio-temporal variation in the projected population growth rate (k) with spatially nested life table response experiments (LTRE). 3. All species (Carlina vulgaris, Tragopogon pratensis and Hypochaeris radicata )s howed considerable life-history variation among regions on top of variation among sites within regions. 4. Net variance contributions (NVC), a novel LTRE statistic, revealed that in each species, variation in one group of vital rates contributed most to variation in k among regions as well as among sites. However, that most important type of vital rates differed between species: plant growth in C. vulgaris, flower head production in T. pratensis and establishment probability of seedlings in H. radicata. The rankings of the NVCs of other vital rates varied between site and region effects, suggesting that buffering through negative vital rate correlations varies over different spatial scales, while the identity of the main contributor tok variation is more constant. 5. Temporal effects were smaller than spatial effects, but the LTREs showed strong interactions between time and space (region or site), suggesting that the effect of, e.g. climate fluctuations are not synchronized throughout the distribution of a species. 6. Synthesis. This study shows that the life histories of plant species are distinguishable even when mean elasticity values show only small differences, and that life histories vary over the distribution range of a species. Demographic differences over large spatial scales can therefore only be partly substituted by small scale spatial variation in modelling studies on the population dynamics of a species across its entire distribution.

Individuals respond to different environments by developing different phenotypes, which is generally seen as a mechanism through which individuals can buffer adverse environmental conditions and increase their fitness. To understand the consequences of phenotypic plasticity it is necessary to study how changing a particular trait of an individual affects either its survival, growth, reproduction or a combination of these demographic vital rates (i.e. fitness components). Integrating vital rate changes due to phenotypic plasticity into models of population dynamics allows detailed study of how phenotypic changes scale up to higher levels of integration and forms an excellent tool to distinguish those plastic trait changes that really matter at the population level. A modeling approach also facilitates studying systems that are even more complex: traits and vital rates often co-vary or trade-off with other traits that may show plastic responses over environmental gradients. Here we review recent developments in the literature on population models that attempt to include phenotypic plasticity with a range of evolutionary assumptions and modeling techniques. We present in detail a model framework in which environmental impacts on population dynamics can be followed analytically through direct and indirect pathways that importantly incorporate phenotypic plasticity, trait-trait and trait-vital rate relationships. We illustrate this framework with two case studies: the population-level consequences of phenotypic responses to nutrient enrichment of plant species occurring in nutrient-poor habitats and of responses to changes in flooding regimes due to climate change. We conclude with exciting prospects for further development of this framework: selection analyses, modeling advances and the inclusion of spatial dynamics by considering dispersal traits as well.

We report on two experiments with oilseed rape (Brassica napus L.) to test if partial root-zone drying techniques improve yield in a crop in which vegetative and reproductive growth overlap (indeterminate growth habit), and to investigate what plant morphological responses contribute to the yield that is realized. Deficit irrigation resulted in smaller plants with smaller yields but larger seeds compared to treatments with shallow groundwater (first experiment) and with fully watered conditions (second experiment). Different partial root-zone drying treatments (water supply patterns) under deficit irrigation, however, had little effect on plant growth and yield components (number of branches, branch lengths, number of pods, etc.). Our results suggest that partial root-zone drying doesn’t work well with oilseed rape. Detailed measurements of soil water contents and root distribution indicate that roots were extremely plastic, effectively foraging for water, and these root responses may have overwhelmed physiological effects of partial root drying on the shoot. Furthermore, in crops with indeterminate growth with a short vegetative growth phase, partial root-zone drying may be ineffective in enhancing the major yield components. Further reasons for the lack of success are discussed.

Summary 1. An increasing body of evidence suggests that within-species diversity plays an important role for community and ecosystem functioning, alters complex trophic interactions and affects patterns of species diversity and coexistence. Nonetheless, we lack a good understanding of how genotypic trait variation translates into shifts in the relative abundance of genotypes within populations. 2. In this study, we show that genotypic selection strongly alters dominance relationships among genotypes over a period of 5 years. This resulted in remarkably consistent changes in the proportional representation of genotypes, and in a concomitant decline of diversity and evenness in our experimental populations. 3. High growth rates and the production of large offspring were positively associated with genotypic performance. Vegetative abundances of genotypes translated monotonically into flowering frequencies. 4. Synthesis. We conclude that genotypic selection markedly affects patterns of diversity and consistently alters genotypic abundance and mean trait distributions in plant populations over a relatively short period of time.

We studied the effects of genotypic and plastic variation in vertical and horizontal spacer lengths on plant performance in a stoloniferous herb subjected to opposing selection regimes. We hypothesized that longer vertical structures are beneficial if plants are subjected to competition, but they should negatively affect plant performance if plants are exposed to aboveground disturbance. To test these hypotheses we subjected 34 genotypes of Trifolium repens to competition and disturbance treatments. Competition was imposed by a grass canopy consisting of Lolium perenne, and disturbance was simulated by regularly clipping the target plants and all the surrounding vegetation at 1 cm above soil level. Conform to our hypothesis, genotypes with longer vertical structures (petioles) produced fewer ramets than genotypes with shorter petioles in the disturbance treatment. However, genotypes with longer petioles did not perform better under competition than genotypes with shorter petioles. Genotypes with highly plastic vertical structures tended to produce more shoot mass under competition, and they produced fewer ramets if subjected to disturbance. Unexpectedly, horizontal structures (stolon internodes) expanded in response to competition which, furthermore, was associated with enhanced plant performance. However, producing longer internodes is inherently associated with costs in terms of increased resource allocation to the longer structures, but not to benefits in terms of increased resource capture. Positive correlations among the length and plasticity of vertical and horizontal structures may explain the apparent positive effect of producing longer internodes on plant performance. Our data thus support the notion that trait correlations may weaken selective forces acting on a focal trait in a specific environment if opposing selection pressures act on genetically correlated traits.

1 An increasing number of studies have shown that plants produce more root mass when sharing rooting space with an intraspecific neighbour as compared with plants growing alone. This so-called self/non-self discrimination has been suggested as a mechanism by which plants may prevent wasteful competition with their own roots and enhance their competitive ability for nutrients with roots of neighbouring plants. The overproduction of root biomass is said to result in a ‘tragedy of the commons’, because it appears to occur at the expense of reproductive biomass. 2 Studies on self/non-self root discrimination have commonly used a split-root design to distinguish self from non-self competition, while keeping the total amount of nutrients available per plant the same. This design has recently been criticized because the rooting volume differs between treatments. 3 Here, we use three general hypotheses to explain the published results without invoking the mechanism of self/non-self discrimination. The hypotheses propose that differences in root mass are due to differences in rooting volume, and differences in nutrient availability determine whole plant growth. More root mass without more growth results in less reproductive biomass. 4 A reanalysis of the results of root self/non-self discrimination confirms these hypotheses. Root overproduction in the presence of another plant, as found in nearly all studies, is consistent with effects of a larger soil volume available to these plants as compared with plants growing alone. Under the same total nutrient availability, total plant weight was the same or higher when more roots were produced. Inevitably, a larger root production with the same total biomass implies that less reproductive biomass is produced. 5 Although our analysis can explain most of the results of the split-root experiments, we cannot rule out the possibility that self/non-self root discrimination did take place. We discuss a limited set of experiments for which volume effects cannot explain the results, suggesting, in fact, that direct self/non-self root interactions have operated. We suggest experimental designs that can demonstrate their ecological significance in the future. 6 We conclude that there is ample evidence that plants can sense the volume of available rooting space, and a limited number of studies on individual roots show that plant roots may sense the identity of neighbouring roots and respond accordingly. The significance of these responses for whole plant growth and reproduction in relation to well-known resource competition effects is yet largely unknown and in urgent need of further study.

Summary 1. Plants are able to detect the presence of their neighbours below-ground. The associated root responses may affect plant performance, plant–plant interactions and community dynamics, but the extent and direction of these responses is heavily debated. 2. Some studies suggest that plants will over-proliferate roots in response to neighbours at the expense of reproduction, which was framed as a ‘tragedy of the commons’. Others propose an ‘ideal free distribution’ hypothesis stating that plants produce roots simply as a function of the amount of available nutrients. However, experimental evidence for either hypothesis that is unbiased by confounding effects of rooting volume and plant size in their experimental set-ups is still lacking. 3. We grew split-root pea plants in the presence or absence of a below-ground neighbour at a range of rooting volumes, while providing equal amounts of nutrients per plant. Path analyses were used to disentangle the direct effects of neighbour presence on allocation patterns from the confounding effects of rooting volume and plant size. 4. Within the chosen range of rooting volumes, the presence of a below-ground neighbour generally reduced plant root mass by 21% and total mass by 9%. A doubling of rooting volume generally increased plant root mass by 22% and total mass by 11%. Pod mass was only directly and positively correlated with vegetative mass. 5. The presence of a below-ground neighbour induced less root allocation but more pod allocation, whereas increased rooting volume caused a reduction in reproductive allocation. A large part of these effects, however, was indirectly mediated through the influence on plant total mass. 6. Synthesis. Not considering the effects of rooting volume and plant size may lead to misinterpretations of plant growth strategies in response to neighbours. Accounting for these factors, we found pea allocating less mass to roots in the presence of a below-ground neighbour. The obtained results can help to reconcile the various responses to below-ground neighbours as they are published in the literature.