Masting behaviour in a Mediterranean pine tree alters seed predator selection on reproductive output

• Context‐dependency in species interactions is widespread and can produce concomitant patterns of context‐dependent selection. Masting (synchronous production of large seed crops at irregular intervals by a plant population) has been shown to reduce seed predation through satiation (reduction in rates of seed predation with increasing seed cone output) and thus represents an important source of context‐dependency in plant‐animal interactions. However, the evolutionary consequences of such dynamics are not well understood.
• Here we describe masting behaviour in a Mediterranean model pine species (Pinus pinaster) and present a test of the effects of masting on selection by seed predators on reproductive output. We predicted that masting, by enhancing seed predator satiation, could in turn strengthen positive selection by seed predators for larger cone output. For this we collected six‐year data (spanning one mast year and five non‐mast years) on seed cone production and seed cone predation rates in a forest genetic trial composed by 116 P. pinaster genotypes.
• Following our prediction, we found stronger seed predator satiation during the masting year, which in turn led to stronger seed predator selection for increased cone production relative to non‐masting years.
• These findings provide evidence that masting can alter the evolutionary outcome of plant‐seed predator interactions. More broadly, our findings highlight that changes in consumer responses to resource abundance represent a widespread mechanism for predicting and understanding context dependency in plant‐consumer evolutionary dynamics.

     

Seedling establishment in a masting desert shrub parallels the pattern for forest trees

The masting phenomenon along with its accompanying suite of seedling adaptive traits has been well studied in forest trees but has rarely been examined in desert shrubs. Blackbrush (Coleogyne ramosissima) is a regionally dominant North American desert shrub whose seeds are produced in mast events and scatter-hoarded by rodents. We followed the fate of seedlings in intact stands vs. small-scale disturbances at four contrasting sites for nine growing seasons following emergence after a mast year. The primary cause of first-year mortality was post-emergence cache excavation and seedling predation, with contrasting impacts at sites with different heteromyid rodent seed predators. Long-term establishment patterns were strongly affected by rodent activity in the weeks following emergence. Survivorship curves generally showed decreased mortality risk with age but differed among sites even after the first year. There were no detectable effects of inter-annual precipitation variability or site climatic differences on survival. Intraspecific competition from conspecific adults had strong impacts on survival and growth, both of which were higher on small-scale disturbances, but similar in openings and under shrub crowns in intact stands. This suggests that adult plants preempted soil resources in the interspaces. Aside from effects on seedling predation, there was little evidence for facilitation or interference beneath adult plant crowns. Plants in intact stands were still small and clearly juvenile after nine years, showing that blackbrush forms cohorts of suppressed plants similar to the seedling banks of closed forests. Seedling banks function in the absence of a persistent seed bank in replacement after adult plant death (gap formation), which is temporally uncoupled from masting and associated recruitment events. This study demonstrates that the seedling establishment syndrome associated with masting has evolved in desert shrublands as well as in forests.     

Within-season synchrony of a masting conifer enhances seed escape

Predator satiation resulting from interannual reproductive synchrony has been widely documented in masting plants, but how reproductive synchrony within a year influences seed escape is poorly understood. We evaluated whether the intra-annual reproductive synchrony of individual white spruce trees (Picea glauca) increased seed escape from their primary predispersal seed predator, North American red squirrels (Tamiasciurus hudsonicus). Trees with cones that matured synchronously relative to those of other trees within red squirrel territories were significantly more likely to escape squirrel predation in years with both low and superabundant levels of cone production, generating a significantly positive linear selection differential for increasing intra-annual reproductive synchrony. Thus, this masting plant escapes seed predation in numbers through interannual synchrony in seed production and in time through intra-annual synchrony of seed availability.     

The evolutionary ecology of mast seeding

The past seven years have seen a revolution in understanding the causes of mast seeding In perennial plants. Before 1987, the two main theories were resource matching (i.e. plants vary their reproductive output to match variable resources) and predator satiation (i.e. losses to predators are reduced by varying the seed crop). Today, resource matching is restricted to a proximate role, and predator satiation is only one of many theories for the ultimate advantage of masting. Wind pollination, prediction of favourable years for seedling establishment, animal pollination, animal dispersal of fruits, high accessory costs of reproduction and large seed size have all been advanced as possible causes of masting. Of these, wind pollination, predator satiation and environmental prediction are important in a number of species, but the other theories have less support. In future, Important advances seem likely from quantifying synchrony within a population, and examining species with very constant reproduction between years.     

Acorn crop size and pre-dispersal predation determine inter-specific differences in the recruitment of co-occurring oaks

The contribution of pre-dispersal seed predation to inter-specific differences in recruitment remains elusive. In species with no resistance mechanisms, differences in pre-dispersal predation may arise from differences in seed abundance (plant satiation) or in the ability of seeds to survive insect infestation (seed satiation). This study aimed to analyse the impact of pre-dispersal acorn predation by weevils in two co-occurring Mediterranean oaks (Quercus ilex and Quercus humilis) and to compare its relevance with other processes involved in recruitment. We monitored the patterns of acorn production and acorn infestation by weevils and we conducted experimental tests of acorn germination after weevil infestation, post-dispersal predation and seedling establishment in mixed forests. Monitoring and experimental data were integrated in a simulation model to test for the effects of pre-dispersal predation in recruitment. In both oaks pre-dispersal acorn infestation decreased with increasing acorn crop size (plant satiation). This benefited Q. ilex which exhibited stronger masting behaviour than Q. humilis, with almost a single and outstanding reproductive event in 6 years. Acorn infestation was more than twice as high in Q. humilis (47.0%) as in Q. ilex (20.0%) irrespective of the number of seeds produced by each species. Although germination of infested acorns (seed satiation) was higher in Q. humilis (60%) than in Q. ilex (21%), this could barely mitigate the higher infestation rate in the former species, to reduce seed loss. Conversely to pre-dispersal predation, no inter-specific differences were observed either in post-dispersal predation or seedling establishment. Our results indicate that pre-dispersal predation may contribute to differences in seed supply, and ultimately in recruitment, between co-existing oaks. Moreover, they suggest that seed satiation can barely offset differences in seed infestation rates. This serves as a warning against overemphasising seed satiation as a mechanism to overcome seed predation by insects. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Climate variation,reproductive frequency and acorn yield in English Oaks

Aims Annually variable but synchronous production of large seed crops (‘masting’) is a widespread phenomenon in temperate trees. Mounting concerns about the impacts of anthropogenic climate change (ACC) on plant reproduction gives urgency to our need to understand better the role of climate on tree reproduction, and in particular, mast events. Unlike our understanding of reproductive phenology however, there is little consensus regarding how climate affects plant reproductive effort or indeed the actual environmental triggers that underpin masting behaviour.     

Predator satiation and recruitment in a mast fruiting monocarpic forest herb

Background and Aims

Cross-pollination and satiation of seed predators are often invoked to explain synchronous mast reproduction in long-lived plants. However, explanations for the synchronous death of parent plants are elusive. The roles of synchronous seeding and post-reproductive mortality were investigated in a perennial monocarpic herb (Isoglossa woodii) in coastal dune forest in South Africa.

Methods

Pre-dispersal seed predation and seed production were assessed by measuring fruit and seed set of inflorescences sprayed with insecticide or water and with no spray treatments. Seed predation was measured at different densities of I. woodii plants by monitoring removal rates of seed from the forest floor. The influence of adult plants on establishment of I. woodii seedlings was assessed by monitoring growth and survivorship of seedlings in caged and uncaged 1 × 1 m plots in understorey gaps and thickets.

Key Results

Fruit and seed set were similar between spray treatments. An I. woodii stem produced 767·8 ± 160·8 seeds (mean ± s.e.) on dune crests and 1359·0 ± 234·4 seeds on the foredune. Seed rain was greater on the foredune than in other topographic locations. Seed predation rates were 32 and 54 % on dune crests and in dune slacks, respectively, and decreased with seed abundance, number of inflorescences per stem and plant height. Seedling recruitment was greater beneath synchronously dying adult plants than in natural understorey gaps (no I. woodii). However, seedling growth rate beneath I. woodii mid-way through its life-cycle was less than in gaps, although survivorship was similar.

Conclusions

The selective advantage of masting in I. woodii derives from satiation of both pre- and post-dispersal seed predators. In addition, post-seeding mortality of adult plants facilitates seedling establishment. Satiation of seed predators and the benefits of seedling establishment are strong drivers of the evolution of synchronous monocarpy in I. woodii.     

Disease resistance in Arabidopsisthaliana increases the competitive ability and the predicted probability of long‐term ecological success under disease pressure

Evidence is mounting that flowering by the mast-fruiting Dipterocarpaceae in Southeast Asia is triggered by ENSO events such that seeds are dispersed at the end of ENSO droughts. These droughts induce substantial defoliation and mortality of canopy trees, producing a favorable environment for seedling recruitment in the forest understory. Therefore, seedling release following droughts may have selected for synchronized, supra-annual fruiting in these rain forests. Currently, mast fruiting in Southeast Asia is generally regarded as an evolutionary response to seed predation by nomadic vertebrates. Separating the two causes for mast fruiting, seedling release and predator satiation, may be difficult if they are coupled in nature by ENSO droughts. Nevertheless, if the cue for masting is environmental, then the post-ENSO seedling environment should be considered a potential cause for masting, and if it operates in conjunction with predator satiation, then it may have provided the initial stimulus for supra-annual synchrony in fruiting.     

Climate sensitivity of reproduction in a mast-seeding boreal conifer across its distributional range from lowland to treeline forests

Mast-seeding conifers such as Picea glauca exhibit synchronous production of large seed crops over wide areas, suggesting climate factors as possible triggers for episodic high seed production. Rapidly changing climatic conditions may thus alter the tempo and spatial pattern of masting of dominant species with potentially far-reaching ecological consequences. Understanding the future reproductive dynamics of ecosystems including boreal forests, which may be dominated by mast-seeding species, requires identifying the specific cues that drive variation in reproductive output across landscape gradients and among years. Here we used annual data collected at three sites spanning an elevation gradient in interior Alaska, USA between 1986 and 2011 to produce the first quantitative models for climate controls over both seedfall and seed viability in P. glauca, a dominant boreal conifer. We identified positive associations between seedfall and increased summer precipitation and decreased summer warmth in all years except for the year prior to seedfall. Seed viability showed a contrasting response, with positive correlations to summer warmth in all years analyzed except for one, and an especially positive response to warm and wet conditions in the seedfall year. Finally, we found substantial reductions in reproductive potential of P. glauca at high elevation due to significantly reduced seed viability there. Our results indicate that major variation in the reproductive potential of this species may occur in different landscape positions in response to warming, with decreasing reproductive success in areas prone to drought stress contrasted with increasing success in higher elevation areas currently limited by cool summer temperatures.     

Aridity drives spatiotemporal patterns of masting across the latitudinal range of a dryland conifer

Masting, or the synchronous and irregular production of seed crops, is controlled by environmental conditions and resource budgets. Increasing temperatures and shifting precipitation regimes may alter the frequency and magnitude of masting, especially in species that experience chronic resource stress. Yet the effects of a changing climate on seed production are unlikely to be uniform across populations, particularly those that span broad abiotic gradients. In this study, we assessed the spatiotemporal patterns of masting across the latitudinal distribution of a widely distributed dryland conifer species, piñon pine Pinus edulis. We quantified seed cone production from 2004 to 2017 using cone abscission scars in 187 trees from 28 sites along an 1100 km latitudinal gradient to investigate the spatiotemporal drivers of seed cone production and synchrony across populations. Populations from chronically hot and dry areas (greater climatic water deficits and less monsoonal precipitation) tended to have greater interannual variability in seed cone production and smaller crop sizes. Mast years generally followed years with low vapor pressure deficits and high precipitation during key periods of the reproductive process, but the strength of these relationships varied across the region. Populations that received greater monsoonal precipitation were less sensitive to late summer vapor pressure deficits during seed cone initiation yet more sensitive to spring vapor pressure deficits during pollination. Spatially correlated patterns of vapor pressure deficit better predicted synchrony in seed cone production than geographic distance, and these patterns were conserved at distances up to 500 km. These results demonstrate that aridity drives spatiotemporal variability in seed cone production. As a result, projected increases in aridity are likely to decrease the frequency and magnitude of masting in these dry forests and woodlands. Declines in seed production may compound climatic limitations to recruitment and impede tree regeneration, with cascading effects for numerous wildlife species.     

Density-dependent reproductive success among sympatric dipterocarps during a major mast fruiting event

Although masting in Southeast Asia is characterized by the interspecific synchronization of reproduction, little is known about the variation in regeneration strategies among sympatric tree species during major masting events. Herein, we examined the hypothesis that non-abundant species achieve greater seed survival per seed production at the pre- and post-dispersal stages by synchronizing reproduction with abundant species. During the masting event from May to September 2010, we installed seed traps in a primary forest plot of the Deramakot Forest Reserve, Borneo. To identify the possible causes of post-dispersal seed mortality, we conducted a seed-sowing experiment from September 2010 to July 2011 in a primary forest plot with high community-level dipterocarp seed density and two surrounding secondary forests with low seed densities. An abundant species (Shorea multiflora) produced more seeds than other species and exhibited a lower survival rate during the pre-dispersal stage. The ratio of aborted seeds was greater in species with greater seed production, while the ratio of seeds predated by insects was not, suggesting that resource limitations and/or greater inbreeding frequency may explain inefficient seed production. Interspecific variation was rarely observed for post-dispersal seed survival rates. Our study highlights the density-dependent variation in reproductive success between abundant and non-abundant dipterocarp species at the pre-dispersal stage. Non-abundant species achieved greater reproductive success by synchronizing reproduction with the masting of abundant species, which might be an important mechanism for sustainability of rare species populations.     

Evolutionary ecology of mast-seeding in temperate and tropical oaks (Quercus spp.)

Mast-seeding is the synchronous production of large seed crops within a population or community of species every two or more years. This paper addresses three non-mutually exclusive hypotheses explaining the evolution of mast-seeding in temperate tree species, especially the genus Quercus: (1) mast-seeding is a consequence of mast-flowering which evolves to increased pollination efficiency in mast-flowering years; (2) mast-seeding has evolved as an anti-predator adaptation by which large seed crops during mast years satiate the seed predators and allow survival of some of the seeds; (3) selection on seed size by habitat can indirectly affect the evolution of masting if trees with large seeds require more time to accumulate reserves to mature those seeds. I find support for the pollination hypothesis in several wind-pollinated temperate tree species but not oaks. However, oaks show evidence favoring the predation and seed size hypotheses. I then develop a model to illustrate the relationships among the three hypotheses in their effects on the evolution of masting. Finally, using data from herbaria and Floras, the influence of selection via flowering, fruiting, and seed size in the evolution of masting in tropical oaks is discussed. I conclude that the need for a supra-annual cue to synchronize flowering and fruiting as well as the larger seed size found in many tropical oak species should contribute to the evolution of masting to a greater extent than seen among temperate oaks.     

Masting in Buxus balearica: assessing fruiting patterns and processes at a large spatial scale

Masting consists of the synchronous highly variable seed production among years by a plant population. We studied spatiotemporal variation in fruit production in ten populations of Buxus balearica (six in the Balearic Islands and four in the Iberian Peninsula) from 2001 to 2004 in the light of masting. In some of them we assessed, by means of both observational and experimental data, the relationship between fruit production and some abiotic variables, the role of previous reproduction, the "pollination efficiency" and the "predator satiation" hypotheses, as well as the consequences for seedling density and survival. Fruit production in B. balearica showed substantial between-year variation, especially in island compared to mainland populations. Correlative evidence indicated that this variation and its geographic pattern were related to differences between regions in rainfall variability, cost of reproduction and the degree of ambophily. We found no indication of predator satiation. However, experimental tests failed to support many of our results, namely a negative effect of previous reproduction on future flowering in island populations and lower pollen limitation with increasing flower production. We therefore warn against exclusive reliance on correlations when testing hypotheses related to masting. In addition, seedling recruitment increased after some episodes of high fruit production but probably additional factors had a role in recruitment, suggesting that mast events not always translate into increased reproductive success. Although a limited time series only allows considering B. balearica as showing 'putative' masting, weather and pollination-related processes are good candidates for further exploration of fruiting patterns and processes at a large spatial scale.     

Reproductive success of individuals with different fruit production patterns. What does it mean for the predator satiation hypothesis?

The predator satiation hypothesis states that synchronous periodic production of seeds is an adaptive strategy evolved to reduce the pressure of seed predators. The seed production pattern is crucial to the predator satiation hypothesis, but there are few studies documenting the success of individuals that are in synchrony and out of synchrony with the whole population. This study is based on long-term data on seed production of Sorbus aucuparia and specialised pre-dispersal seed predation by Argyresthia conjugella, in a subalpine spruce forest in the Western Carpathians (Poland). At the population level, we tested whether functional and numerical responses of predators to the variation of fruit production operate. At the individual level, we tested whether individuals with higher interannual variability in their own seed crops and higher synchrony with the population have higher percentages of uninfested fruits. The intensity of pre-dispersal seed predation was high (average 70 %; range 19–100 %). There were both functional and numerical responses of predators to the variation of fruit production at the population level. We found that individuals that were expected to be preferred under seed predator pressure had higher reproductive success. With increasing synchrony of fruit production between individual trees and the population, the percentage of infested fruits decreased. There was also a negative relationship between the interannual variation in individual fruit production and the percentage of infested fruits. These results confirm selection for individuals with a masting strategy. However, the population does not seem well adapted to strong seed predation pressure and we suggest that this may be a result of prolonged diapause of A. conjugella.     

Temporal variation in the female components of reproductive success over the extended flowering season of a Mediterranean perennial herb

We examined the reproductive success of the perennial herb Lobularia maritima during its extended flowering and fruiting season. The within- and between-year variability of the female components of reproductive success (from flower, fruit and seed production to seed survival, seed germination and seedling establishment) were analysed during four flowering seasons. All the components of reproductive success studied showed a significant within- and between-year variation. September was the period of the year with the maximum values of flower and fruit production, and the highest germination and establishment rates. Nevertheless, seed losses due to both predispersal and postdispersal seed predation during this period were also the highest, seriously reducing seed output in this period. On the other hand, in those periods in which seed production was low, i.e. January and May, the percentage of seeds lost to seed predators was the lowest. Reproductive success in each period of the flowering season was estimated using a simple demographic model, in which the information concerning all the components already calculated was integrated. The two variables used to estimate reproductive success in each period, i.e. the number of new individuals produced per plant and the probability of a seed becoming an adult plant, showed relatively small differences over the year. These results suggest a counter-balance of the different components of reproductive success in this species, with favourable and unfavourable periods for the different components being compensated during its extended flowering season.     

Masting of rowan <Emphasis Type="Italic">Sorbus aucuparia</Emphasis> L. and consequences for the apple fruit moth <Emphasis Type="Italic">Argyresthia conjugella</Emphasis> Zeller

 Masting of rowan Sorbus aucuparia L. has been studied in 45 sites in southern Norway for 22 years. We present data on the year-to-year variation in fruit setting of rowan, and show that masting is spatially synchronous in Norway and probably all over Fennoscandia. The apple fruit moth Argyresthia conjugella Zeller is an important seed predator on rowan. We present data on the abundance of apple fruit moth in rowanberries during these years and discuss the consequences of masting and intermasting of rowan for apple fruit moth as a pest of apple. We conclude that growth and climate have little impact on flowering intensity and suggest that masting of rowan is an adaptive defense against seed predation and a new example of predator satiation: intermast years inhibit predators and prepare the rowan for the subsequent mast. Received: September 3, 2001 / Accepted: February 24, 2003     

Reproductive collapse in European beech results from declining pollination efficiency in large trees

Climate warming increases tree mortality which will require sufficient reproduction to ensure population viability. However, the response of tree reproduction to climate change remains poorly understood. Warming can reduce synchrony and interannual variability of seed production (“masting breakdown”) which can increase seed predation and decrease pollination efficiency in trees. Here, using 40 years of observations of individual seed production in European beech (Fagus sylvatica), we showed that masting breakdown results in declining viable seed production over time, in contrast to the positive trend apparent in raw seed count data. Furthermore, tree size modulates the consequences of masting breakdown on viable seed production. While seed predation increased over time mainly in small trees, pollination efficiency disproportionately decreased in larger individuals. Consequently, fecundity declined over time across all size classes, but the overall effect was greatest in large trees. Our study showed that a fundamental biological relationship—correlation between tree size and viable seed production—has been reversed as the climate has warmed. That reversal has diverse consequences for forest dynamics; including for stand- and biogeographical-level dynamics of forest regeneration. The tree size effects suggest management options to increase forest resilience under changing climates.     

When sex is not enough: ecological correlates of resprouting capacity in congeneric tropical forest shrubs

In moist tropical forests resprouting may be an important component of life history, contributing to asexual reproduction through the clonal spread of individuals derived from shoot fragments. However, in contrast to other ecosystems where resprouting is common, the ecological correlates of resprouting capacity in tropical forests remain largely unexplored. In this study we characterized shade tolerance, resprouting capacity and sexual reproductive success of eight co-occurring Piper species from lowland forests of Panama. In field experiments we found that shade-tolerant Piper species had a higher capacity to regenerate from excised or pinned stem fragments than light-demanding species in both gap and understory light conditions. In contrast, shade-tolerant species had lower recruitment probabilities from seeds, as a consequence of lower initial seed viability, and lower seedling emergence rates. All Piper species needed gap conditions for successful seedling establishment. Of 8,000 seeds sown in the understory only 0.2% emerged. In gaps, seed germination of light-demanding species was between 10 and 50%, whereas for shade-tolerant species it was 0.5–9.8%. We propose that the capacity to reproduce asexually from resprouts could be adaptive for shade-tolerant species that are constantly exposed to damage from falling litter in the understory. Resprouting may allow Piper populations to persist and spread despite the high rate of pre-dispersal seed predation and low seed emergence rates. Across Piper species, we detected a trade-off between resprouting capacity and the annual viable seed production per plant but not with annual seed mass produced per plant. This suggests that species differences in sexual reproductive success may not necessarily result from differential resource allocation. Instead we suggest that low sexual reproductive success in the understory may in part reflect reduced genetic diversity in populations undergoing clonal growth, resulting in self-fertilization and in-breeding depression.     

Masting in a temperate tree: Evidence for environmental prediction?

Many plant species produce large fruit crops in some years and then produce few or no fruits in others. Synchronous, inter‐annual variation in plant reproduction is known as ‘masting’ and its adaptive significance has yet to be fully resolved. For 8 consecutive years, I quantified every fruit produced by 22 females of a New Zealand tree species (Dysoxylum spectabile), which has an unusual habit of taking a full calendar year to mature fruits after flowering. Fruit production varied strongly among years and was tightly synchronized among trees. Annual variability in fruit production declined with total reproductive output, indicating trees with lower fecundity exhibited a stronger tendency to mast. Although unrelated to temperature, annual fruit production was positively related to precipitation during annual periods of fruit development, and negatively related to fruit production in the previous year. Seedlings had higher rates of survivorship in a wet, high‐seed year than in a dry, low‐seed year, suggesting that seedlings might be drought sensitive. Therefore, D. spectabile produced large fruit crops during periods of high rainfall prior to fruit maturation, which may enhance survivorship of drought‐intolerant seeds. Results were inconsistent with several hypotheses that are widely believed to be the most likely explanations for masting. Instead, results were consistent with the environmental prediction hypothesis, suggesting that this hypothesis may be more important than previously appreciated.     

Masting uncoupling: mast seeding does not follow all mast flowering episodes in a dioecious juniper tree

Evolutionary selective forces, like predator satiation and pollination efficiency, are acknowledged to be major causes of masting (the variable, periodic and synchronic production of seeds in a population). However, a number of recent studies indicate that resources might also play an important role on shaping masting patterns. Dioecious masting species offer a privileged framework to study the role of resources on masting variation, since male and female plants often experience different reproductive costs and selective pressures. We followed masting and reproductive investment (RI) of the dioecious tree Juniperus thurifera in two populations along 10 years and studied the different response of males and females to experimentally increased water and nutrient availability in a third population. Juniperus thurifera females invested in reproduction three times more resources than males. Such disparity generated different resource‐use strategies in male and female trees. Tree‐ring growth and water use efficiency (WUE) confirmed that sexes differed in their resource investment temporal pattern, with males using current resources for reproduction and females using resources accumulated during longer periods. Watered and fertilized female trees presented significantly higher flowering reproductive investments than males and experienced an extraordinary mast‐flowering event. However, seeding RI and mast seeding were not affected by the treatment. This suggests that although resource availability affects the reproductive output of this species, there are other major forces regulating masting on J. thurifera. During 10 years, J. thurifera male and female trees presented high and low flowering years more or less synchronously. However, not all mast flowering episodes resulted in mast seeding, leading to masting uncoupling between flowering and seeding. Since flowering costs represent only 1% of females’ total reproductive investments, masting uncoupling could be a beneficial bet‐hedging strategy to maximize reproductive output in spite of unpredictable catastrophic events.