Testing population differentiation in plant species – how important are environmental maternal effects

The maternal environment may contribute to population differentiation in offspring traits if growing conditions of mother plants are different. However, the magnitude of such environmental maternal effects compared with genetic differentiation is often not clear. We tested the importance of environmental maternal effects by comparing population differentiation in parental seed directly collected in the field and in F₁ seed grown under homogeneous conditions. The F₁ seeds were obtained by random crosses within populations. We used five populations in each of four plant species to analyse seed mass and growth chamber germination of both generations at the same time. In two species, we additionally tested offspring performance in the field. We found a significant population differentiation in all species and for nearly all measured traits. Population‐by‐generation interactions indicating environmental maternal effects were significant for germination (three species) and for seed mass (two species) but not for growth and reproduction. The significant interaction was partly due to a reduction of among‐population differentiation from the parental to the F₁ generation that can be explained by a decrease of maternal provisioning effects. However, in some species by trait combinations a change in population ranking and not a decrease of variation was responsible for significant population‐by‐generation interactions indicating environmental maternal effects beyond maternal provisioning. Fitting of seed mass as covariate was not successful in reducing environmental maternal effects on population differentiation in germination. We discuss alternative methods to account for environmental maternal effects in studies on genetic differentiation among populations.     

Functional trade‐offs and the phylogenetic dispersion of seed traits in a biodiversity hotspot of the Mountains of Southwest China

The diversity of traits associated with plant regeneration is often shaped by functional trade‐offs where plants typically do not excel at every function because resources allocated to one function cannot be allocated to another. By analyzing correlations among seed traits, empirical studies have shown that there is a trade‐off between seedling development and the occupation of new habitats, although only a small range of taxa have been tested; whether such trade‐off exists in a biodiverse and complex landscape remains unclear. Here, we amassed seed trait data of 1,119 species from a biodiversity hotspot of the Mountains of Southwest China and analyzed the relationship between seed mass and the number of seeds and between seed mass and time to germination. Our results showed that seed mass was negatively correlated with seed number but positively correlated with time to germination. The same trend was found regardless of variation in life‐form and phylogenetic conservatism. Furthermore, the relation between seed mass and other seed traits was randomly dispersed across the phylogeny at both the order and family levels. Collectively, results suggest that there is a functional trade‐off between seedling development and new habitat occupation for seed plants in this region. Larger seeds tend to produce fewer seedlings but with greater fitness compared to those produced by smaller seeds, whereas smaller seeds tend to have a larger number of seeds that germinate faster compared to large‐seeded species. Apart from genetic constraints, species that produce large seeds will succeed in sites where resource availability is low, whereas species with high colonization ability (those that produce a high number of seeds per fruit) will succeed in new niches. This study provides a mechanistic explanation for the relatively high levels of plant diversity currently found in a heterogeneous region of the Mountains of Southwest China.     

Maternal provisioning is structured by species’ competitive neighborhoods

Differential maternal provisioning of offspring in response to environmental conditions has been argued as ‘the missing link’ in plant life histories. Although empirical evidence suggests that maternal provisioning responses to abiotic conditions are common, there is little understanding of how differences in maternal provisioning manifest in response to competition. Frequency manipulations are commonly employed in ecological studies to assess the strength of interspecific competition, relative to intraspecific competition, and we used frequency manipulations to test how competition in two soil moisture environments affects maternal provisioning of seed mass. Specifically, for 15 pairs comprised from 25 annual plant species that occur in California, we varied the relative frequencies of conspecific to heterospecific competitors from 90% (intraspecific competition) to 10% (interspecific competition). We found that conspecific frequency affected maternal provisioning (seed mass) in 12 of the 25 species (eight significantly (p < 0.05), four marginally significantly (p < 0.07)), and that these responses included both increased (five species) and decreased (six species) seed mass, as well as one species with opposing directions of response to conspecific frequency that depended on the soil moisture environment. Conspecific frequency also affected per capita fecundity (seed number) for 17 of the 25 species (15 significantly (p < 0.05), two marginally significantly (p < 0.09)), which generally decreased seed number as conspecific frequency increased. The direction and magnitude of frequency‐dependent seed mass depended on the identity of the competitor, even among species whose fecundity was not affected by competitor identity; the latter finding reveals competitive differences among species that would otherwise appear to be competitively equivalent. Our research demonstrates how species responses to different competitive environments manifest through maternal provisioning, and that these responses alter previous estimates of environmentally‐determined maternal provisioning and reproductive output; future study is needed to understand their combined effects on population and community dynamics.     

Fertiliser application positively affects plants performance but reduces seed viability in seashore mallow (Kosteletzkya pentacarpos): implication for biomass production and species conservation

Kosteletzkya pentacarpos seashore mallow, is recognised as an important plant species for several industrial applications, especially in North America and Asia. In Europe, seashore mallow is highly threatened and subject of conservation actions (e.g. reintroduction). In order to define appropriate multipurpose cultivation protocols, suitable for biomass production and for conservation purposes, we investigated the effect of varying levels of soil salt content and nutrient availability on seashore mallow. As expected, seashore mallow had the best performance in terms of growth rate, flowering and fruit production when fertiliser was applied, while salt had only limited effects. Interestingly, seeds produced by fertilised plants showed a lower germination performance and higher mortality than seeds developed from non‐fertilised plants. Our results highlight a trade‐off between parental plant growth and seed performance, the former being enhanced and the latter being reduced by fertiliser application. The causes of this trade‐off are related to a transgenerational maternal effect of fertiliser application that has important implications for seashore mallow cultivation. Biomass production benefits from fertiliser application but if the quality of seeds and the fitness of the next generation are major objectives (e.g. seed production and species conservation), fertiliser application is not recommended.     

Functional trade‐offs increase species diversity in experimental plant communities

Functional trade‐offs have long been recognised as important mechanisms of species coexistence, but direct experimental evidence for such mechanisms is extremely rare. Here, we test the effect of one classical trade‐off – a negative correlation between seed size and seed number – by establishing microcosm plant communities with positive, negative and no correlation between seed size and seed number and analysing the effect of the seed size/number correlation on species richness. Consistent with theory, a negative correlation between seed size and seed number led to a higher number of species in the communities and a corresponding wider range of seed size (a measure of functional richness) by promoting coexistence of large‐ and small‐seeded species. Our study provides the first direct evidence that a seed size/number trade‐off may contribute to species coexistence, and at a wider context, demonstrates the potential role of functional trade‐offs in maintaining species diversity.     

Seed size predicts global effects of small mammal seed predation on plant recruitment

Recent studies demonstrate that by focusing on traits linked to fundamental plant life‐history trade‐offs, ecologists can begin to predict plant community structure at global scales. Yet, consumers can strongly affect plant communities, and means for linking consumer effects to key plant traits and community assembly processes are lacking. We conducted a global literature review and meta‐analysis to evaluate whether seed size, a trait representing fundamental life‐history trade‐offs in plant offspring investment, could predict post‐dispersal seed predator effects on seed removal and plant recruitment. Seed size predicted small mammal seed removal rates and their impacts on plant recruitment consistent with optimal foraging theory, with intermediate seed sizes most strongly impacted globally – for both native and exotic plants. However, differences in seed size distributions among ecosystems conditioned seed predation patterns, with relatively large‐seeded species most strongly affected in grasslands (smallest seeds), and relatively small‐seeded species most strongly affected in tropical forests (largest seeds). Such size‐dependent seed predation has profound implications for coexistence among plants because it may enhance or weaken opposing life‐history trade‐offs in an ecosystem‐specific manner. Our results suggest that seed size may serve as a key life‐history trait that can integrate consumer effects to improve understandings of plant coexistence.     

Effects of increased nitrogen deposition and precipitation on seed and seedling production of Potentilla tanacetifolia in a temperate steppe ecosystem

Background

The responses of plant seeds and seedlings to changing atmospheric nitrogen (N) deposition and precipitation regimes determine plant population dynamics and community composition under global change.

Methodology/Principal Findings

In a temperate steppe in northern China, seeds of P. tanacetifolia were collected from a field-based experiment with N addition and increased precipitation to measure changes in their traits (production, mass, germination). Seedlings germinated from those seeds were grown in a greenhouse to examine the effects of improved N and water availability in maternal and offspring environments on seedling growth. Maternal N-addition stimulated seed production, but it suppressed seed mass, germination rate and seedling biomass of P. tanacetifolia. Maternal N-addition also enhanced responses of seedlings to N and water addition in the offspring environment. Maternal increased-precipitation stimulated seed production, but it had no effect on seed mass and germination rate. Maternal increased-precipitation enhanced seedling growth when grown under similar conditions, whereas seedling responses to offspring N- and water-addition were suppressed by maternal increased-precipitation. Both offspring N-addition and increased-precipitation stimulated growth of seedlings germinated from seeds collected from the maternal control environment without either N or water addition. Our observations indicate that both maternal and offspring environments can influence seedling growth of P. tanacetifolia with consequent impacts on the future population dynamics of this species in the study area.

Conclusion/Significance

The findings highlight the importance of the maternal effects on seed and seedling production as well as responses of offspring to changing environmental drivers in mechanistic understanding and projecting of plant population dynamics under global change.     

Quantitative and qualitative consequences of reduced pollen loads in a mixed‐mating plant

Greater pollination intensity can enhance maternal plant fitness by increasing seed set and seed quality as a result of more intense pollen competition or enhanced genetic sampling. We tested experimentally these effects by varying the pollen load from a single pollen donor on stigmas of female flowers of Dalechampia scandens (Euphorbiaceae) and measuring the effects on seed number and seed mass. Seed set increased rapidly with pollen number at low to moderate pollen loads, and a maximum set of three seeds occurred with a mean pollen load of 19 pollen grains. We did not detect a trade‐off between the number of seeds and seed mass within a fruit. Seed mass increased with increasing pollen load, supporting the hypothesis of enhanced seed quality via increased pollen‐competition intensity or genetic sampling. These results suggest that maternal fitness increases with larger pollen loads, even when the fertilization success is already high. Our results further highlight the importance of high rates of pollen arrival onto stigmas, as mediated by reliable pollinators. Comparing the pollen‐to‐seed response curve obtained in this experiment with those observed in natural populations suggests that pollen limitation may be more severe in natural populations than predicted from greenhouse studies. These results also indicate that declines in pollinator abundance may decrease plant fitness through lowered seed quality before an effect on seed set is detected.     

HETEROGENEITY IN INDIVIDUAL QUALITY AND REPRODUCTIVE TRADE‐OFFS WITHIN SPECIES

Interspecifically, a reasonable body of evidence supports a trade‐off between offspring size and number. However, at the intraspecific level, a whole manner of phenotypic correlations between offspring size and number are observed. These correlations may be predicted when heterogeneity in resource availability, or quality, is considered. Making the assumption that maternal size is a proxy for resource availability, we meta‐analytically quantified four phenotypic reproductive correlations within numerous species: (1) maternal size and offspring size, (2) maternal size and offspring number, (3) offspring number and offspring size, and (4) offspring number and offspring size after controlling for maternal size. Within species, maternal size showed a positive correlation with both offspring size and number. Despite this consistency, no correlation between offspring size and number was found. After controlling for maternal size, however, offspring size and number showed a significant negative correlation. A phylogenetic component of our analysis accounted for little heterogeneity in the data, suggesting that our findings show remarkable consistency across taxa. Overall, our results support an observable phenotypic trade‐off between offspring size and number. However, this analysis also highlights the importance of considering quality when examining trade‐offs, a task that is not always straightforward as quality is context dependant.     

Do large‐seeded herbs have a small range size? The seed mass–distribution range trade‐off hypothesis

We aimed to introduce and test the “seed mass–distribution range trade‐off” hypothesis, that is, that range size is negatively related to seed mass due to the generally better dispersal ability of smaller seeds. Studying the effects of environmental factors on the seed mass and range size of species, we also aimed to identify habitats where species may be at risk and need extra conservation effort to avoid local extinctions. We collected data for seed mass, global range size, and indicators for environmental factors of the habitat for 1,600 species of the Pannonian Ecoregion (Central Europe) from the literature. We tested the relationship between species’ seed mass, range size, and indicator values for soil moisture, light intensity, and nutrient supply. We found that seed mass is negatively correlated with range size; thus, a seed mass–distribution range trade‐off was validated based on the studied large species pool. We found increasing seed mass with decreasing light intensity and increasing nutrient availability, but decreasing seed mass with increasing soil moisture. Range size increased with increasing soil moisture and nutrient supply, but decreased with increasing light intensity. Our results supported the hypothesis that there is a trade‐off between seed mass and distribution range. We found that species of habitats characterized by low soil moisture and nutrient values but high light intensity values have small range size. This emphasizes that species of dry, infertile habitats, such as dry grasslands, could be more vulnerable to habitat fragmentation or degradation than species of wet and fertile habitats. The remarkably high number of species and the use of global distribution range in our study support our understanding of global biogeographic processes and patterns that are essential in defining conservation priorities.     

Variation among populations of Diodia teres (Rubiaceae) in environmental maternal effects

Previous studies have quantified variation in environmental maternal effects (EME) within populations, but these effects could differ among populations as well. In this study we grew clonal replicates of individuals from three populations of the annual plant Diodia teres in their native and non-native environments. Our goal was to estimate the effects of maternal environment and maternal population on seed and seedling traits. Seeds that were produced in this field study were then planted in two soil types to quantify effects of the offspring environment on seedling traits. There was substantial variation among populations for seed weight. We found population variation for EME, and maternal environment by offspring environment interactions. We conclude that variation among populations in EME may be an unrecognized component of local adaptation, and that attempts to control maternal effects by statistically accounting for variation in seed weight may be ineffective.     

A trait‐based trade‐off between growth and mortality: evidence from 15 tropical tree species using size‐specific relative growth rates

A life‐history trade‐off between low mortality in the dark and rapid growth in the light is one of the most widely accepted mechanisms underlying plant ecological strategies in tropical forests. Differences in plant functional traits are thought to underlie these distinct ecological strategies; however, very few studies have shown relationships between functional traits and demographic rates within a functional group. We present 8 years of growth and mortality data from saplings of 15 species of Dipterocarpaceae planted into logged‐over forest in Malaysian Borneo, and the relationships between these demographic rates and four key functional traits: wood density, specific leaf area (SLA), seed mass, and leaf C:N ratio. Species‐specific differences in growth rates were separated from seedling size effects by fitting nonlinear mixed‐effects models, to repeated measurements taken on individuals at multiple time points. Mortality data were analyzed using binary logistic regressions in a mixed‐effects models framework. Growth increased and mortality decreased with increasing light availability. Species differed in both their growth and mortality rates, yet there was little evidence for a statistical interaction between species and light for either response. There was a positive relationship between growth rate and the predicted probability of mortality regardless of light environment, suggesting that this relationship may be driven by a general trade‐off between traits that maximize growth and traits that minimize mortality, rather than through differential species responses to light. Our results indicate that wood density is an important trait that indicates both the ability of species to grow and resistance to mortality, but no other trait was correlated with either growth or mortality. Therefore, the growth mortality trade‐off among species of dipterocarp appears to be general in being independent of species crossovers in performance in different light environments.     

Species with greater seed mass are more tolerant of conspecific neighbours: a key driver of early survival and future abundances in a tropical forest

Multiple niche‐based processes including conspecific negative density dependence (CNDD) determine plant regeneration and community structure. We ask how interspecific and intraspecific density‐dependent interactions relate to plant life histories and associated functional traits. Using hierarchical models, we analysed how such interactions affected first‐year survival of seedling recruits of 175 species in a tropical forest, and how species abundances and functional traits are related to interspecific variation in density‐dependent effects. Conspecific seedling neighbour effects prevailed over the effects of larger conspecific and all heterospecific neighbours. Tolerance of seedling CNDD enhanced recruit survival and subsequent abundance, all of which were greater among larger seeded, slow‐growing and well‐defended species. Niche differentiation along the growth–survival trade‐off and tolerance of seedling CNDD strongly correlated with regeneration success, with manifest consequences for community structure. The ability of larger seeded species to better tolerate CNDD suggests a novel mechanism for CNDD to contribute to seed‐size variation and promote species coexistence through a tolerance–fecundity trade‐off.     

Performance and responses to competition in two congeneric annual species: does seed heteromorphism matter?

Variations in seed characteristics observed in heteromorphic species may affect various stages of their life cycles, e.g. seed dormancy, germination characteristics or even adult plant performance. Highly specialised seed morphs – described as colonisers and maintainers – exhibit a trade‐off between colonisation capacity and competitive traits. The performance of distinct seed morph progeny under competitive conditions, and especially in multi‐species arrangements, had previously not been given much attention. In this study, we compared performance and response to competition among distinct seed morph progenies in two congeneric, co‐occurring species: the invasive Bidens frondosa and the non‐invasive Bidens tripartita. We hypothesised that maintainer seed morphs of both species would perform better under increased plant densities and within inter‐morphic mixtures, while coloniser morphs would show stronger responses to increased densities and perform relatively poorly in inter‐morphic mixtures. We conducted a growth trial and a greenhouse experiment, which revealed that seed morph progeny differed significantly in plant height when grown without competition, while under competitive conditions such differences became less apparent. The observed pattern was more strongly pronounced in B. frondosa, which showed a general predominance in stature and biomass over its non‐invasive congener. Although seed morphs performed equally well under competitive conditions, increased plant height and more rapid germination can favour the maintainer seed morph on sites where vegetation is already present.     

Transgenerational consequences of plant responses to herbivory: an adaptive maternal effect?

Herbivory has many effects on plants, ranging from shifts in primary processes such as photosynthesis, growth, and phenology to effects on defense against subsequent herbivores and other species interactions. In this study, I investigated the effects of herbivory on seed and seedling characteristics of several families of wild radish (Raphanus raphanistrum) to test the hypothesis that herbivory may affect the quality of offspring and the resistance of offspring to plant parasites. Transgenerational effects of herbivory may represent adaptive maternal effects or factors that constrain or amplify natural selection on progeny. Caterpillar (Pieris rapae) herbivory to greenhouse-grown plants caused plants in some families to produce smaller seeds and those in other families to produce larger seeds compared with undamaged controls. Seed mass was positively associated with probability of emergence in the field. The number of setose trichomes, a putative plant defense, was higher in the progeny of damaged plants in some families and lower in the progeny of damaged plants in other families. In a field experiment, plant families varied in their resistance to several herbivores and pathogens as well as in growth rate and time to flowering. Seeds from damaged parent plants were more likely to become infested with a plant virus. Although herbivory on maternal plants did not directly affect interactions of offspring with other plant parasites, seed mass influenced plant resistance to several attackers. Thus, herbivory affected seed characters, which mediated interactions between plants and their parasites. Finally, irrespective of seed mass, herbivory on maternal plants influenced components of progeny fitness, which was dependent on plant family. Natural selection may act on plant responses to herbivory that affect seedling-parasite interactions and, ultimately, fitness.     

The role of the tolerance–fecundity trade‐off in maintaining intraspecific seed trait variation in a widespread dimorphic herb

Coexistence of species with different seed sizes is a long‐standing issue in community ecology, and a trade‐off between fecundity and stress tolerance has been proposed to explain co‐occurrence in heterogeneous environments. Here we tested an intraspecific extension of this model: whether such trade‐off also explains seed trait variation among populations of widespread plants under stress gradients. We collected seeds from 14 populations of Plantago coronopus along the Atlantic coast in North Africa and Europe. This herb presents seed dimorphism, producing large basal seeds with a mucilaginous coat that facilitates water absorption (more stress tolerant), and small apical seeds without coats (less stress tolerant). We analysed variation among populations in number, size and mucilage production of basal and apical seeds, and searched for relationships between local environment and plant size. Populations under higher stress (higher temperature, lower precipitation, lower soil organic matter) had fewer seeds per fruit, higher predominance of basal relative to apical seeds, and larger basal seeds with thicker mucilaginous coats. These results strongly suggest a trade‐off between tolerance and fecundity at the fruit level underpins variation in seed traits among P. coronopus populations. However, seed production per plant showed the opposite pattern to seed production per fruit, and seemed related to plant size and other life‐cycle components, as an additional strategy to cope with environmental variation across the range. The tolerance–fecundity model may constitute, under stress gradients, a broader ecological framework to explain trait variation than the classical seed size–number compromise, although several fecundity levels and traits should be considered to understand the diverse strategies of widespread plants to maximise fitness in each set of local conditions.     

Seed traits,seed‐reserve utilization and offspring performance across pre‐industrial to future CO2 concentrations in a Mediterranean community

Atmospheric CO₂ enrichment can affect plants directly via impacts on their performance, and indirectly, by environment‐specific traits passed down from the mother plant to the offspring. Such maternal effects can significantly alter plant species composition, especially in annual ecosystems where the entire community is recruited from seeds each year. This study assessed impacts of future, high CO₂ (440 and 600 ppm) and pre‐industrial, low CO₂ (280 ppm) on seed traits and offspring performance in three plant functional groups (grasses, legumes, forbs) comprising 17 annual species of a semi‐arid Mediterranean community. In grasses, seed size and seed‐reserve utilization as expressed by root elongation tended to be higher at high than at low maternal CO₂, but total seed protein concentration and protein pool decreased with increasing maternal CO₂. The response of seed size to high CO₂ increased with increasing leaf‐mass fraction in grasses, and decreased with decreasing concentration of leaf non‐structural carbohydrates in legumes. Offspring development was studied at ambient CO₂, and showed reduced emergence success of high‐CO₂ progeny compared with low‐CO₂ progeny in forbs. Total biomass was lower in high‐CO₂ than in low‐CO₂ offspring across all functional groups. The biomass response to high maternal CO₂ in legume offspring correlated inversely with seed size, resulting in up to 25% lower biomass in large‐seeded species. Under the scenario of maternal effects combined with projected changes in biomass and seed production under direct exposure to high CO₂, legumes might gain and forbs and grasses might lose from future CO₂ enrichment. Most changes in seed traits and offspring performance were greater between pre‐industrial and near‐future CO₂ than between near‐ and remote‐future CO₂ concentrations. Hence, maternal effects of increasing CO₂ may contribute to current changes in plant productivity and species composition, and they need to be considered when predicting impacts of global change on plant communities.     

Effects of variation in resource acquisition during different stages of the life cycle on life‐history traits and trade‐offs in a burying beetle

Individual variation in resource acquisition should have consequences for life‐history traits and trade‐offs between them because such variation determines how many resources can be allocated to different life‐history functions, such as growth, survival and reproduction. Since resource acquisition can vary across an individual's life cycle, the consequences for life‐history traits and trade‐offs may depend on when during the life cycle resources are limited. We tested for differential and/or interactive effects of variation in resource acquisition in the burying beetle Nicrophorus vespilloides. We designed an experiment in which individuals acquired high or low amounts of resources across three stages of the life cycle: larval development, prior to breeding and the onset of breeding in a fully crossed design. Resource acquisition during larval development and prior to breeding affected egg size and offspring survival, respectively. Meanwhile, resource acquisition at the onset of breeding affected size and number of both eggs and offspring. In addition, there were interactive effects between resource acquisition at different stages on egg size and offspring survival. However, only when females acquired few resources at the onset of breeding was there evidence for a trade‐off between offspring size and number. Our results demonstrate that individual variation in resource acquisition during different stages of the life cycle has important consequences for life‐history traits but limited effects on trade‐offs. This suggests that in species that acquire a fixed‐sized resource at the onset of breeding, the size of this resource has larger effects on life‐history trade‐offs than resources acquired at earlier stages.     

The relationship between individual seed quality and maternal plant body size in crowded herbaceous vegetation

Aims In most natural plant populations, there is a strong right-skewed distribution of body sizes for reproductive plants—i.e. the vast majority are relatively small, suppressed weaklings that manage not just to survive effects of crowding/competition and other hazards but also to produce offspring. Recent research has shown that because of their relatively large numbers, these relatively small resident plants collectively contribute most of the seed offspring production available for the population in the next generation. However, the success of these offspring will depend in part on their quality, e.g. reflected by seed size and resource content. Accordingly, in the present study, we used material from natural populations of herbaceous species to test the null hypothesis that there is no significant relationship between body size variation in resident plants—resulting from between-site variation in the intensity of crowding/competition—and variation in the mass or N content of their individual seeds.Methods Using populations of 56 herbaceous species common in eastern Ontario, total above-ground dry plant mass, mean mass per seed and mean nitrogen (N) content per seed were recorded for a sample of the largest resident plants and also for the smallest reproductive plants growing in local neighbourhoods with the most severe crowding/competition from near neighbours.Important findings Mass per seed was numerically smaller from the smallest resident plants for most study species, but with few exceptions, this was not significantly different (P> 0.05) from mass per seed from the largest resident plants. The results therefore showed no general effect of maternal plant body size on individual seed mass, or N content. This suggests that the reproductive output of the smaller half of the resident plant size distribution within these populations is likely to contribute not just most of the seed production available for the next generation but also seed offspring that are just as likely—on a per individual basis—to achieve seedling/juvenile recruitment success as the seed offspring produced by the largest resident plants. This conflicts with the traditional 'size-advantage' hypothesis for predicting plant fitness under severe competition, and instead supports the recent 'reproductive-economy-advantage' hypothesis, where competitive fitness is promoted by capacity to produce offspring that—despite severe body size suppression imposed by neighbour effects—in turn have capacity to produce grand-offspring.     

Trait plasticity alters the range of possible coexistence conditions in a competition–colonisation trade‐off

Most of the classical theory on species coexistence has been based on species‐level competitive trade‐offs. However, it is becoming apparent that plant species display high levels of trait plasticity. The implications of this plasticity are almost completely unknown for most coexistence theory. Here, we model a competition–colonisation trade‐off and incorporate trait plasticity to evaluate its effects on coexistence. Our simulations show that the classic competition–colonisation trade‐off is highly sensitive to environmental circumstances, and coexistence only occurs in narrow ranges of conditions. The inclusion of plasticity, which allows shifts in competitive hierarchies across the landscape, leads to coexistence across a much broader range of competitive and environmental conditions including disturbance levels, the magnitude of competitive differences between species, and landscape spatial patterning. Plasticity also increases the number of species that persist in simulations of multispecies assemblages. Plasticity may generally increase the robustness of coexistence mechanisms and be an important component of scaling coexistence theory to higher diversity communities.