There are two main life cycles in plants—annual and perennial1,2. These life cycles are associated with different traits that determine ecosystem function3,4. Although life cycles are textbook examples of plant adaptation to different environments, we lack comprehensive knowledge regarding their global distributional patterns. Here we assembled an extensive database of plant life cycle assignments of 235,000 plant species coupled with millions of georeferenced datapoints to map the worldwide biogeography of these plant species. We found that annual plants are half as common as initially thought5–8, accounting for only 6% of plant species. Our analyses indicate that annuals are favoured in hot and dry regions. However, a more accurate model shows that the prevalence of annual species is driven by temperature and precipitation in the driest quarter (rather than yearly means), explaining, for example, why some Mediterranean systems have more annuals than desert systems. Furthermore, this pattern remains consistent among different families, indicating convergent evolution. Finally, we demonstrate that increasing climate variability and anthropogenic disturbance increase annual favourability. Considering future climate change, we predict an increase in annual prevalence for 69% of the world’s ecoregions by 2060. Overall, our analyses raise concerns for ecosystem services provided by perennial plants, as ongoing changes are leading to a higher proportion of annual plants globally.

The soil seed bank is a major component of plant communities. However, long-term analyses of the dynamics of the seed bank and the ensuing vegetation are rare. Here, we studied the dynamics in plant communities with high dominance of annuals in Mediterranean, semiarid, and arid ecosystems for nine consecutive years. For annuals, we hypothesized that the density of the seed bank would be more stable than the density of the standing herbaceous vegetation. Moreover, we predicted that differences in temporal variability between the seed bank and the vegetation would increase with aridity, where year-to-year rainfall variability is higher. We found that the temporal variability at the population level (assessed as the standard deviation of the loge-transformed density) of the nine dominant annuals in each site did not differ between the seed bank and the ensuing vegetation in any of the sites. For the total density of annuals, patterns depended on aridity. In the Mediterranean site, the temporal variability was similar in the seed bank and the vegetation (0.40 vs. 0.40). Still, in the semiarid and arid sites, variability in the seed bank was lower than in the vegetation (0.49 vs. 1.01 and 0.63 vs. 1.38, respectively). This difference between the population-level patterns and the total density of annuals can be related to the lower population synchrony in their seed bank. In contrast, for the herbaceous perennials (all species combined), the seed bank variability was higher than in the vegetation. Overall, our results highlight the role of the seed bank in buffering the annual vegetation density with increasing climatic uncertainty typical in aridity gradients. This role is crucial under the increasing uncertainty imposed by climatic change in the region.

AbstractResource competition theory predicts coexistence and exclusion patterns based on species? R*s, the minimum resource values required for a species to persist. A central assumption of the theory is that all species have equal access to resources. However, many systems are characterized by preemption exploitation, where some species deplete resources before their competitors can access them (e.g., asymmetric light competition, contest competition among animals). We hypothesized that coexistence under preemption requires an R*-preemption trade-off?that is, the species with the priority access should have a higher R* (lower ?efficiency?). Thus, we developed an extension of resource competition theory to investigate partial and total preemption (in the latter, the preemptor is unaffected by species with lower preemption rank). We found that an R*-preemption trade-off is a necessary condition for coexistence in all models. Moreover, under total preemption, the trade-off alone is sufficient for coexistence. In contrast, under partial preemption, more conditions are needed, which restricts the parameter space of coexistence. Finally, we discuss the implications of our finding for seemingly distinct trade-offs, which we view as special cases of the R*-preemption trade-off. These trade-offs include the digger-grazer trade-off, the competition-colonization trade-off, and trade-offs related to light competition between trees and understories.AbstractResource competition theory predicts coexistence and exclusion patterns based on species? R*s, the minimum resource values required for a species to persist. A central assumption of the theory is that all species have equal access to resources. However, many systems are characterized by preemption exploitation, where some species deplete resources before their competitors can access them (e.g., asymmetric light competition, contest competition among animals). We hypothesized that coexistence under preemption requires an R*-preemption trade-off?that is, the species with the priority access should have a higher R* (lower ?efficiency?). Thus, we developed an extension of resource competition theory to investigate partial and total preemption (in the latter, the preemptor is unaffected by species with lower preemption rank). We found that an R*-preemption trade-off is a necessary condition for coexistence in all models. Moreover, under total preemption, the trade-off alone is sufficient for coexistence. In contrast, under partial preemption, more conditions are needed, which restricts the parameter space of coexistence. Finally, we discuss the implications of our finding for seemingly distinct trade-offs, which we view as special cases of the R*-preemption trade-off. These trade-offs include the digger-grazer trade-off, the competition-colonization trade-off, and trade-offs related to light competition between trees and understories.

Nutrient enrichment of natural ecosystems is a primary characteristic of the Anthropocene and a known cause of biodiversity loss, particularly in grasslands. In a global meta-analysis of 630 resource addition experiments, we conduct a simultaneous test of the three most prominent explanations of this phenomenon. Our results conclusively indicate that nitrogen is the leading cause of species loss. This result is important because of the increase in nitrogen deposition and the frequent use of nitrogen-based fertilizers worldwide. Our findings provide global-scale, experimental evidence that minimizing nitrogen inputs to ecological systems may help to conserve the diversity of grassland ecosystems. Eutrophication is a major driver of species loss in plant communities worldwide. However, the underlying mechanisms of this phenomenon are controversial. Previous studies have raised three main explanations: 1) High levels of soil resources increase standing biomass, thereby intensifying competitive interactions (the “biomass-driven competition hypothesis”). 2) High levels of soil resources reduce the potential for resource-based niche partitioning (the “niche dimension hypothesis”). 3) Increasing soil nitrogen causes stress by changing the abiotic or biotic conditions (the “nitrogen detriment hypothesis”). Despite several syntheses of resource addition experiments, so far, no study has tested all of the hypotheses together. This is a major shortcoming, since the mechanisms underlying the three hypotheses are not independent. Here, we conduct a simultaneous test of the three hypotheses by integrating data from 630 resource addition experiments located in 99 sites worldwide. Our results provide strong support for the nitrogen detriment hypothesis, weaker support for the biomass-driven competition hypothesis, and negligible support for the niche dimension hypothesis. The results further show that the indirect effect of nitrogen through its effect on biomass is minor compared to its direct effect and is much larger than that of all other resources (phosphorus, potassium, and water). Thus, we conclude that nitrogen-specific mechanisms are more important than biomass or niche dimensionality as drivers of species loss under high levels of soil resources. This conclusion is highly relevant for future attempts to reduce biodiversity loss caused by global eutrophication.

Abstract Ecological selection is a major driver of community assembly. Selection is classified as stabilizing when species with intermediate trait values gain the highest reproductive success, whereas selection is considered directional when fitness is highest for species with extreme trait values. Previous studies have investigated the effects of different selection types on trait distribution, but the effects of selection on species diversity have remained unclear. Here, we propose a framework for inferring the type and strength of selection by studying species diversity and trait distribution together against null expectations. We use a simulation model to confirm our prediction that directional selection should lead to lower species diversity than stabilizing selection despite a similar effect on trait community-weighted variance. We apply the framework to a mesocosm system of annual plants to test whether differences in species diversity between two habitats that vary in productivity are related to differences in selection on seed mass. We show that, in both habitats, species diversity was lower than the null expectation, but that species diversity was lower in the more productive habitat. We attribute this difference to strong directional selection for large-seeded species in the productive habitat as indicated by trait community-weighted mean being higher and community-weighted variance being lower than the null expectations. In the less productive habitat, we found that community-weighted variance was higher than expected by chance, suggesting that seed mass could be a driver of niche partitioning under such conditions. Altogether, our results suggest that viewing species diversity and trait distribution as interrelated patterns driven by the same process, ecological selection, is helpful in understanding community assembly.

Abstract Ecological theory predicts that the soil seed bank stabilizes the composition of annual plant communities in the face of environmental variability. However, long-term data on the community dynamics in the seed bank and the standing vegetation are needed to test this prediction. We tested the hypothesis that the composition of the seed bank undergoes lower temporal variability than the standing vegetation in a 9-year study in Mediterranean, semi-arid and arid ecosystems. The composition of the seed bank was estimated by collecting soil cores from the studied sites on an annual basis. Seedling emergence under optimal watering conditions was measured in each soil core for three consecutive years, to account for seed dormancy. In all sites, the composition of the seed bank differed from the vegetation throughout the years. Small-seeded and dormant-seeded species had a higher frequency in the seed bank than in the standing vegetation. In contrast, functional group membership (grasses vs. forbs) did not explain differences in species frequency between the seed bank and the vegetation after controlling for differences between grasses and forbs in seed mass and seed dormancy. Contrary to predictions, the magnitude of year-to-year variability (the mean compositional dissimilarity between consecutive years) was not lower in the seed bank than in the vegetation in all sites. However, long-term compositional trends in the seed bank were weaker than in the vegetation in the Mediterranean and semi-arid sites. In the arid site where year-to-year variability was highest, no long-term trends were observed. Synthesis. The effect of the seed bank on the temporal variability of the vegetation in annual communities depends on site conditions and time-scale. While the year-to-year variability of the seed bank is similar to the vegetation, the soil seed bank can buffer long-term trends.

AbstractLife-history trade-offs among species are major drivers of community assembly. Most studies investigate how trade-offs promote deterministic coexistence of species. It remains unclear how trade-offs may instead promote historically contingent exclusion of species, where species dominance is affected by initial abundances, causing alternative community states via priority effects. Focusing on the establishment-longevity trade-off, in which high longevity is associated with low competitive ability during establishment, we study the transient dynamics and equilibrium outcomes of competitive interactions in a simulation model of plant community assembly. We show that in this model, the establishment-longevity trade-off is a necessary but not sufficient condition for alternative stable equilibria, which also require low fecundity for both species. An analytical approximation of our simulation model demonstrates that alternative stable equilibria are driven by demographic stochasticity in the number of seeds arriving at each establishment site. This site-scale stochasticity is affected only by fecundity and therefore occurs even in infinitely large communities. In many cases where the establishment-longevity trade-off does not cause alternative stable equilibria, the trade-off still decreases the rate of convergence toward the single equilibrium, resulting in decades of transient dynamics that can appear indistinguishable from alternative stable equilibria in empirical studies.

ABSTRACT Ecological selection is a major driver of community assembly. Selection is classified as stabilizing when species with intermediate trait values gain the highest reproductive success, whereas selection is considered directional when fitness is highest for species with extreme trait values. Previous studies have investigated the effects of different selection types on trait distribution, but the effects of selection on species diversity have remained unclear. Here, we propose a framework for inferring the type and strength of selection by studying species diversity and trait distribution together against null expectations. We use a simulation model to confirm our prediction that directional selection should lead to lower species diversity than stabilizing selection despite a similar effect on trait community-weighted variance. We apply the framework to a mesocosm system of annual plants to test whether differences in species diversity between two habitats that vary in productivity are related to differences in selection on seed mass. We show that, in both habitats, species diversity was lower than the null expectation, but that species diversity was lower in the more productive habitat. We attribute this difference to strong directional selection for large-seeded species in the productive habitat as indicated by trait community-weighted-mean being higher and community-weighted variance being lower than the null expectations. In the less productive habitat, we found that community-weighted variance was higher than expected by chance, suggesting that seed mass could be a driver of niche partitioning under such conditions. Altogether, our results suggest that viewing species diversity and trait distribution as interrelated patterns driven by the same process, ecological selection, is helpful in understanding community assembly.