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  1. Abstract

    Herbivores often have highly variable impacts on plant fecundity. The relative contribution of different environmental factors operating at varying spatial scales in affecting this variability is often unclear. We examined how density‐dependent seed predation at local scales and regional differences in primary productivity are associated with variation in the magnitude of pre‐dispersal seed predation onMonarda fistulosa(Lamiaceae). WithinM. fistulosapopulations growing in a low‐productivity region (LPR), Montana, USA, and a high‐productivity region (HPR), Wisconsin, USA, we quantified the magnitude of pre‐dispersal seed predation among individual plants differing in seed head densities. Out of a total of 303M. fistulosaplants that were surveyed, we found half as many herbivores in seed heads in the LPR (n = 133 herbivores) compared to the HPR (n = 316). In the LPR, 30% of the seed heads were damaged in plants with low seed head density, while 61% of seed heads were damaged in plants with high seed head density. Seed head damage was consistently high in the HPR (about 49% across the range of seed head density) compared to the LPR (45% across a range of seed head density). However, the proportion of seeds per seed head that were destroyed by herbivores was nearly two times higher (~38% loss) in the LPR compared to HPR (22% loss). Considering the combined effects of probability of damage and seed loss per seed head, the proportion seed loss per plant was consistently higher in the HPR regardless of seed head density. Nevertheless, because of greater seed head production, the total number of viable seeds produced per plant was higher in HPR and high‐density plants, despite being exposed to greater herbivore pressure. These findings show how large‐scale factors can interact with local‐scale factors to influence how strongly herbivores suppress plant fecundity.

     
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  2. Dormancy has repeatedly evolved in plants, animals, and microbes and is hypothesized to facilitate persistence in the face of environmental change. Yet previous experiments have not tracked demography and trait evolution spanning a full successional cycle to ask whether early bouts of natural selection are later reinforced or erased during periods of population dormancy. In addition, it is unclear how well short-term measures of fitness predict long-term genotypic success for species with dormancy. Here, we address these issues using experimental field populations of the plantOenothera biennis, which evolved over five generations in plots exposed to or protected from insect herbivory. While populations existed above ground, there was rapid evolution of defensive and life-history traits, but populations lost genetic diversity and crashed as succession proceeded. After >5 y of seed dormancy, we triggered germination from the seedbank and genotyped >3,000 colonizers. Resurrected populations showed restored genetic diversity that reduced earlier responses to selection and pushed population phenotypes toward the starting conditions of a decade earlier. Nonetheless, four defense and life-history traits remained differentiated in populations with insect suppression compared with controls. These findings capture key missing elements of evolution during ecological cycles and demonstrate the impact of dormancy on future evolutionary responses to environmental change.

     
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  3. Abstract

    In many invaded grasslands, dominant exotic species can produce large amounts of litter that modify local abiotic conditions and species' interactions. These novel conditions can reduce native species abundance and promote the persistence of exotic species, yet the strength of this disparity may be influenced by how consumer pressure interacts with litter accumulation. Consumers may exacerbate this disparity by preferentially targeting native species or by promoting heterogeneity in microhabitats due to their movement and small‐scale ground disturbances that favours fast growing exotic species. How species respond to litter accumulation and consumer pressure may depend on either evolutionary differences, whereby exotics species may benefit from a lack natural predators, or by functional differences, in which species' physiological traits may confer fitness advantages to low‐light conditions or herbivory or granivory pressure.

    We examined the impact of litter presence and small mammal herbivory on the establishment and reproduction of functionally diverse exotic versus native species seeded across sites that naturally vary in resource availability in an annual invaded California grassland. We assessed whether seed mass and leaf nitrogen content (LNC) were predictive of successful establishment and reproduction.

    Litter accumulation affected exotic and native species differently, with litter significantly decreasing native recruitment and reproduction, while exotics were largely unaffected. Small mammals had a slight positive effect on the establishment of native species when litter was present but did not influence exotic species. Regardless of species provenance, larger seeded species established at a higher density while species with lower leaf nitrogen content had a higher density of reproductive individuals. Native species that successfully established and reproduced were functionally different in LNC than the resident community, while successful exotic species were functionally more similar to the resident community in LNC.

    Our study demonstrates that exotic species outperformed native species regardless of the presence of litter or herbivory pressure. Without the removal or thinning of litter, it is likely that exotic species will continue to dominate, resulting in positive feedback that further favours the persistence of exotic species within this invaded grassland system.

    Read the freePlain Language Summaryfor this article on the Journal blog.

     
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  4. Abstract

    A paradigm in the plant defence literature is that defending against herbivores comes at a cost to growth, resulting in a growth–defence trade‐off. However, while there is strong evidence for growth–defence trade‐offs across species, evidence is mixed within species.

    Several mechanisms can account for this equivocal support within species, but teasing them apart requires examining growth–defence relationships both within and among populations, an approach seldom employed.

    We examined correlations between plant biomass (growth) and terpene production (defence) within and among populations ofMonarda fistulosa, a perennial herb. We sampled populations from Montana and Wisconsin, regions that differ in resource availability characterized by different summer precipitation and associated abiotic conditions that influence plant productivity.

    We found negative, neutral and positive growth–defence correlations, depending on the scale examined. Negative correlations occurred across populations originating from divergent regions, positive correlations occurred across populations originating from within the high‐resource region and neutral correlations were found within single populations.

    Collectively, these results challenge the general expectation of ubiquitous trade‐offs and support emerging views that resource availability (as it affects productivity) shapes the evolution of defence at different scales.

    A freePlain Language Summarycan be found within the Supporting Information of this article.

     
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  5. Abstract

    Despite theoretical advances, the ecological factors and functional traits that enable species varying in seed size and fecundity to coexist remain unclear. Given inherent fecundity advantages, why don't small‐seeded species dominate communities?

    In perennial grasslands, we evaluated whether small‐seeded species are less tolerant of competition from the community dominant bunchgrass than large‐seeded species, but also less vulnerable to seed predation by mice. We also explored whether trade‐offs involving competitive tolerance include two other functional traits, height and leaf mass per area (LMA). We added seeds of 17 forb species to plots where bunchgrass competition and rodent seed predation were manipulated across sites varying in bunchgrass productivity and thus competitive intensity. Seeds were added at densities mimicking interspecific variation in fecundity among target species.

    Standardizing for differences in fecundity (i.e. seed input, which enabled us to evaluate inherent interspecific differences in susceptibility to biotic interactions), bunchgrass competition more greatly reduced recruitment and establishment of small‐ versus large‐seeded species, whereas rodent seed predation more greatly reduced the recruitment of large‐ versus small‐seeded species. Plant height and LMA were unrelated to the competition effect size.

    Small‐seeded species abundance decreased across sites increasing in bunchgrass productivity, whereas this was not the case for large‐seeded species. For adult plants but not seedlings, community‐weighted functional trait means (CWM) for seed size, height and LMA increased in plots with versus without bunchgrass competition and the CWM for seed size and height also increased at sites with greater bunchgrass productivity (for adults only). In contrast, rodent seed predation had no significant effects on CWM seed size.

    At the end of the experiment, adult abundance positively correlated with plant fecundity in plots lacking bunchgrass, indicating the inherent advantages accrued to high fecundity small‐seeded species. However, with bunchgrass competition, abundances were equalized across species due to reduced competitive tolerance of high fecundity small‐seeded species.

    Synthesis. Our results suggest that coexistence among subordinate forb species varying in seed size and fecundity is in‐part due to a trade‐off involving competitive tolerance and fecundity, mediated by seed size and associated functional traits.

     
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