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Niv DeMalach

Publications and source records attributed to Niv DeMalach.

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Anthropogenic disturbance expands the climatic limits of annual plant dominance

Disturbance regimes and nutrient inputs are changing worldwide, with consequences for the structure and functioning of plant communities. Classical life-history theory predicts that disturbance should shift communities from long-lived perennials toward short-lived annuals, and that nutrient enrichment may amplify this shift. However, these predictions have not been tested experimentally across broad environmental gradients. Here, using a global coordinated grassland experiment spanning 37 sites, we tested how physical disturbance, vegetation removal and shallow soil tillage, and fertilisation reshape annual-perennial balance, and whether disturbance relaxes the climatic limits of annual dominance. Disturbance nearly doubled the proportion of annual species and more than doubled the relative cover of annuals, whereas fertilisation had little influence and did not interact with disturbance. The disturbance-driven shift arose through contrasting pathways: in graminoids and legumes, it reflected the loss of perennial cover, while in forbs, the expansion of annual cover. In the absence of disturbance, annual dominance was restricted to systems with extremely hot and dry summers, but disturbance nearly tripled the extent of climate space in which annuals dominated. By rapidly reassembling after disturbance, annuals may help maintain vegetation cover, but their expansion also signals loss of perennial cover and the long-term ecosystem functions associated with it.

q-bio.OT

Revising the global biogeography of annual and perennial plants

There are two main life cycles in plants, annual and perennial. These life cycles are associated with different traits that determine ecosystem function. 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 data points to map the worldwide biogeography thereof. We found that annuals are half as common as initially thought, accounting for only 6% of plant species. Our analyses indicate annuals are favored in hot and dry regions. However, a more accurate model shows annual species' prevalence 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 deserts. Furthermore, this pattern remains consistent among different families, indicating convergent evolution. Finally, we demonstrate that increasing climate variability and anthropogenic disturbance increase annual favorability. 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 perennials, as ongoing changes are leading to a more annuals-dominated world.

q-bio.PE

Contrasting dynamics of seed banks and standing vegetation of annuals and perennials along a rainfall gradient

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.

q-bio.PE

Density-dependent and independent mechanisms jointly reduce species performance under nitrogen enrichment

Nitrogen (N) deposition is a primary driver of species loss in plant communities globally. However, the mechanisms by which high N availability causes species loss remain unclear. Many hypotheses for species loss with increasing N availability highlight density-dependent mechanisms, i.e., changes in species interactions. However, an alternative set of hypotheses highlights density-independent detrimental effects of nitrogen (e.g., N toxicity). We tested the role of density-dependent and density-independent mechanisms in reducing species performance. For this aim, we used 120 experimental plant communities comprised of annual species growing together in containers under four fertilization treatments: (1) no nutrient addition(, (2) all nutrients except N (P, K, and micronutrients), (3) Low N, and (4) high N. Each fertilization treatment included two sowing densities to differentiate between the effects of competition (N * density interactions) and other detrimental effects of N. We focused on three performance attributes: the probability of reaching the reproduction period, biomass growth, and population growth. We found that individual biomass and population growth rates decreased with increasing sowing density in all nutrient treatments, implying that species interactions were predominantly negative. The common grass had a higher biomass and population growth under N enrichment, regardless of sowing density. In contrast, the legume showed a density-independent reduction in biomass growth with increasing N. Lastly, the small forb showed a density-dependent reduction in population growth, i.e., the decline occurred only under high density. Our results demonstrate that density-dependent and density-independent mechanisms operate simultaneously to reduce species performance under high N availability. Yet, their relative importance varies among species and life stages.

q-bio.PE

Coexistence under hierarchical resource exploitation: the role of R*-preemption tradeoff

Resource 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 hypothesize that coexistence under preemption requires an R*-preemption tradeoff, i.e., 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 tradeoff is a necessary condition for coexistence in all models. Moreover, under total preemption, the tradeoff alone is sufficient for coexistence. In contrast, under partial preemption, more conditions are needed, which restricts the parameter space of coexistence. Finally, we discussed the implications of our finding for seemingly distinct tradeoffs, which we view as special cases of R*-preemption tradeoff. These tradeoffs include the digger-grazer, the competition-colonization, and tradeoffs related to light competition between trees and understories.

q-bio.PE

Alternative states in plant communities driven by a life-history tradeoff and demographic stochasticity

Life-history tradeoffs among species are major drivers of community assembly. Most studies investigate how tradeoffs promote deterministic coexistence of species. It remains unclear how tradeoffs may instead promote historically contingent exclusion of species, where species dominance is affected by initial abundances, causing alternative community states. Focusing on the establishment-longevity tradeoff, 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 tradeoff is a necessary but not sufficient condition for alternative stable equilibria that require also 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 only affected by fecundity and therefore occurs even in infinitely large communities. In many cases where the establishment-longevity tradeoff does not cause alternative stable equilibria, it 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.

q-bio.PE

The soil seed bank can buffer long-term compositional changes in annual plant communities

Ecological theory predicts that the soil seed bank stabilises 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 nine-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. Overall, 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.

q-bio.PE

Seed mass diversity along resource gradients: the role of allometric growth rate and size-asymmetric competition

The large variation in seed mass among species inspired a vast array of theoretical and empirical research attempting to explain this variation. So far, seed mass variation was investigated by two classes of studies: one class focuses on species varying in seed mass within communities, while the second focuses on variation between communities, most often with respect to resource gradients. Here, we develop a model capable of simultaneously explaining variation in seed mass within and between communities. The model describes resource competition (for both soil and light resources) in annual communities and incorporates two fundamental aspects: light asymmetry (higher light acquisition per unit biomass for larger individuals) and growth allometry (negative dependency of relative growth rate on plant biomass). Results show that both factors are critical in determining patterns of seed mass variation. In general, growth allometry increases the reproductive success of small-seeded species while light asymmetry increases the reproductive success of large-seeded species. Increasing availability of soil resources increases light competition, thereby increasing the reproductive success of large-seeded species and ultimately the community (weighted) mean seed mass. An unexpected prediction of the model is that maximum variation in community seed mass (a measure of functional diversity) occurs under intermediate levels of soil resources. Extensions of the model incorporating size-dependent seed survival and disturbance also show patterns consistent with empirical observations. These overall results suggest that the mechanisms captured by the model are important in determining patterns of species and functional diversity.

q-bio.PE

Plant species accumulation curves are determined by evenness and spatial aggregation in drylands worldwide

Species accumulation curves (SAC), i.e. the relationship between species richness and the number of sampling units in a given community, can be used to describe diversity patterns while accounting for the well-known scale-dependence of species richness. Despite their value, the functional form and the parameters of SAC, as well as their determinants, have barely been investigated in plant communities, particularly in drylands. We characterized the SAC of perennial plant communities from 233 dryland ecosystems from six continents by comparing the fit of major functions (power-law, logarithmic and Michaelis-Menten). We tested the theoretical prediction that the effects of aridity and soil pH on SAC are mediated by vegetation attributes such as evenness, cover, and spatial aggregation. We found that the logarithmic relationship was the most common functional form, followed by Michaelis-Menten and power-law. Functional form was mainly determined by evenness while the SAC parameters (intercept and slope) were largely determined by spatial aggregation. In addition, aridity decreased small scale richness (intercept of SAC) but did not affect accumulation rate (slope of the SAC). Our results highlight the role that attributes such as spatial aggregation and evenness play as main mediators of the SAC of vegetation in drylands, the Earth's largest biome.

q-bio.PE