Climate-driven synchrony in seed production of masting deciduous and conifer tree species

Aims Understanding fluctuations in plant reproductive investment can constitute a key challenge in ecology, conservation and management. Masting events of trees (i.e. the intermittent and synchronous production of abundant seeding material) is an extreme example of such fluctuations. Our objective was to establish the degree of spatial and temporal synchrony in common four masting tree species in boreal Finland and account for potential causal drivers of these patterns.Methods We investigated the spatial intraspecific and temporal interspecific fluctuations in annual seed production of four tree species in Finland, silver birch Betula pendula Roth, downy birch Betula pubescens Ehrh., Norway spruce Picea abies (L.) H.Karst. and rowanberry Sorbus aucuparia L. We also tested to see whether variations in seed production were linked to annual weather conditions. Seeding abundance data were derived from tens of stands per species across large spatial scales within Finland during 1979 to 2014 (for rowanberries only 1986 to 2014).Important findings All species showed spatial synchrony in seed production at scales up to 1000 km. Annual estimates of seed production were strongly correlated between species. Spring and summer temperatures explained most variation in crop sizes of tree species with 0-to 2-year time lags, whereas rainfall had relatively little influence. Warm weather during flowering (May temperature) in the flowering year (Year t) and 2 years before (t ?2) were correlated with seed production. However, high May temperatures during the previous year (t^-1) adversely affected seed production. Summer temperatures in Year t^-1 was positively correlated with seed production, likely because this parameter enhances the development of flower primordials, but the effect was negative with a time lag of 2 years. The negative feedback in temperature coefficients is also likely due to patterns of resource allocation, as abundant flowering and seed production in these species is thought to reduce the subsequent initiation of potential new flower buds. Since the most important weather variables also showed spatial correlation up to 1000 km, weather parameters likely explain much of the spatial and temporal synchrony in seed production of these four studied tree species.     

Direct and indirect effects of masting on rodent populations and tree seed survival

Many plant species are thought to benefit from mast seeding as a result of increased seed survival through predator satiation. However, in communities with many different masting species, lack of synchrony in seed production among species may decrease seed survival by maintaining seed predator populations through the intermast cycle. Similarly, masting by different plant species may have different effects on the seed predator community. We conducted a three-year study in a northeastern USA temperate deciduous forest to determine if production of large seed crops by several tree species was synchronous, and if they had similar effects on all small mammal species. We found that red oak mast crops resulted in increased densities of Peromyscus leucopus and P. maniculatus , but had no effect on Clethrionomys gapperi abundance. Conversely, C. gapperi populations, but not Peromyscus populations, appeared to increase in response to a large red maple seed crop. Differences in small mammal abundance resulted in changes in species-specific seed survival: in the year of abundant C. gapperi , experimentally placed red oak acorns had significantly higher survival than in the year of high Peromyscus abundance. Red oak acorn removal was positively correlated with Peromyscus abundance, while red maple seed removal was significantly higher with increased C. gapperi abundance. Thus, species-specific seed production had differential effects on subsequent small mammal abundance, which in turn affected seed survival. We suggest that at the level of the community, even short-term lack of synchrony in production of large seed crops can cause variation in postdispersal seed survival, through differential effects on the community of small mammal seed predators.     

How does oak mast seeding affect the feeding behavior of sympatric red and roe deer?

Oak reproduction is characterized by mast seeding with high inter-annual fluctuations in fruit production. Such resource pulses can greatly affect ecosystem functioning and may cause seed consumers to alter their mobility, demography, or diet. Consequences of mast seeding for seed consumers remain poorly understood as their long timescale makes them difficult to study. We investigated impacts of oak mast seeding on the feeding behavior of two sympatric European deer species: red deer (Cervus elaphus) and roe deer (Capreolus capreolus). We analyzed their rumen content over a 31-year period in tandem with 10 years of data on oak fructification (i.e. 8 years of field monitoring and two modelled years). Acorn production is strongly correlated with consumption by both deer species. In years of high fructification, acorns represent more than 50% and 35% of red and roe deer diet, respectively, confirming assumptions that deer favor acorns when these are available. Red deer eat more acorns than roe deer both between and within years. High acorn production in mast years appears to saturate the capacity of deer to consume acorns. As the proportion of acorns increase in their diet, red deer eat more grasses and less conifer browse. No dietary shift was found for roe deer. By inducing dietary shifts in consumers, oak mast seeding can have cascading effects on ecosystem processes, notably on the damages on conifers caused by red deer and the consequences for forest dynamics.     

植物大年结实研究概述

大年结实指一个指物种群在不同年份同时产生具有很大变化的种子产量,这种生殖特征普遍存在于多年生植物中,是植物有性生殖过程中的一个重要进化对策。植物种子产量的周期性变化和种群内不同个体结实的同步程度是植物大年结实的两个重要特征。本文对大年结实的有关假说以及各自依据和缺陷进行了分析,讨论了植物种群内不同个体生殖同步性的程度和时间尺度的延续和空间尺度的异质性。     

西双版纳望天树林种子雨9年动态

种子雨是森林生态系统过程的重要组成部分。虽然国内外已有部分关于种子雨的研究, 但多数集中在温带和亚热带地区, 热带地区种子雨的长期监测研究仍显不足。为全面了解热带森林种子雨输入的数量动态和物种组成的长期规律, 本文对我国西双版纳望天树(Parashorea chinensis)林群落水平上9年(2008年5月至2017年4月)的种子雨输入进行了研究。9年共收集到木本植物的种子226种8,993,224粒, 其中大型种子(I-1型) 77种9,779粒; 中型种子(I-2型) 61种24,237粒; 小型种子(I-3型) 72种83,399粒; 粒状微小种子(II-1型) 13种1,215,235粒; 粒状极小种子(II-2型) 1种7,244,667粒; 细丝状种子(II-3型) 2种415,907粒。种子雨输入量排名前10位的优势种贡献了70%以上的种子数量。各类型种子的种子雨输入量均具有年际波动, 呈“大小年”的交替变化, 间隔期为1-2年, 物种组成也呈现一定的年际差异。持续9年的监测中, 每类型种子至少观察到了1次大年结实事件。     

Intra- and inter-specific effects of mast seeding on seed fates of two sympatric <Emphasis Type="Italic">Corylus</Emphasis> species

We released seeds of two sympatric tree species, Corylus mandshurica (seed with thinner seed hull, higher nutrition) and C. heterophylla (seeds with thicker seed hull, lower nutrition) in the masting year of C. mandshurica in 2008, and C. heterophylla in 2009, respectively, to investigate how seed masting of the two sympatric Corylus species affects seed removal and dispersal fitness of the two species differently at both intra- and inter-specific levels. At intra-specific level, the authors found mast seeding of both C. mandshurica and C. heterophylla significantly reduced seed removal, seed consumption, but increased seed dispersal distance and seed dispersal fitness of the released seeds. Mast seeding of C. mandshurica increased seed caching of C. mandshurica. At inter-specific level, the authors found mast seeding of C. mandshurica reduced seed removal of C. heterophylla, but mast seeding of C. heterophylla did not significantly reduce seed removal of C. mandshurica. Mast seeding of C. mandshurica reduced seed consumption of C. heterophylla, while mast seeding of C. heterophylla reduced seed consumption of C. mandshurica. We found mast seeding of C. mandshurica significantly reduced seed dispersal distance of C. heterophylla, while mast seeding of C. heterophylla significantly increased seed dispersal distance of C. mandshurica. We found that mast seeding of C. mandshurica significantly increased seed dispersal fitness of C. heterophylla, while mast seeding of C. heterophylla did not significantly increase seed dispersal fitness of C. mandshurica. More studies are needed to reveal the ecological consequences of mast seeding at inter-specific or community-level. Seed traits may attribute the differences of mast seeding at inter-specific level. Because seeds with thinner seed hull and higher nutrition were more harvested and eaten by rodents, mast seeding of C. mandshurica might have reduced seed removal and seed consumption, but increased dispersal fitness of C. heterophylla (seeds with thicker seed hull, lower nutrition). Therefore, synchrony among species is, or is not, selectively beneficial to the focus species depends on seed traits which determine gains from mast seeding at inter-specific level.     

Mechanisms driving interspecific variation in regional synchrony of trees reproduction

Seed production in many plants is characterized by large interannual variation, which is synchronized at subcontinental scales in some species but local in others. The reproductive synchrony affects animal migrations, trophic responses to resource pulses and the planning of management and conservation. Spatial synchrony of reproduction is typically attributed to the Moran effect, but this alone is unable to explain interspecific differences in synchrony. We show that interspecific differences in the conservation of seed production-weather relationships combine with the Moran effect to explain variation in reproductive synchrony. Conservative timing of weather cues that trigger masting allows populations to be synchronized at distances >1000 km. Conversely, if populations respond to variable weather signals, synchrony cannot be achieved. Our study shows that species vary in the extent to which their weather cueing is spatiotemporally conserved, with important consequences, including an interspecific variation of masting vulnerability to climate change.     

Spatial dynamics of Norwegian tetraonid populations

Different species in a given site or population of a given species in different sites may fluctuate in synchrony if they are affected similarly by factors such as spatially autocorrelated climate, predation, or by dispersal between populations of one species. We used county wise time series of hunting bag records of four Norwegian tetraonid species covering 24 years to examine patterns of interspecific and intraspecific synchrony. We estimated synchrony at three spatial scales; national, regional (consisting of counties with similar climate), and county level. Ecologically related species with overlapping distributions exhibited strong synchrony across Norway, but there was much variation between the different regions and counties. Regions with a long coastline to both the North Sea and the Norwegian Ocean exhibited an overall stronger synchrony than those consisting of more continental areas. Intraspecific synchrony was generally low across all counties, but stronger synchrony between counties within regions defined by climatic conditions. Synchrony was negatively related to distance between populations in three of four species. Only the synchrony in willow ptarmigan showed a clear negative relationship with distance, while the other species had both strong positive and negative correlations at short distances. Strong interspecific synchrony between some species pairs within regions and weak intraspecific synchrony across counties within regions suggest a stronger synchronizing effect from environmental factors such as weather or predation and less effect from dispersal. Our results suggest that the complete tetraonid community is structured by environmental factors affecting the different species similarly and causes widespread interspecific synchrony. Local factors affecting the population dynamics nevertheless frequently forces neighbouring populations out of phase.     

Desynchronization and re-synchronization of reproduction by Astragalus scaphoides, a plant that flowers in alternate years

Mast seeding, the synchronous seed production by plants at irregular intervals, has been widely studied from the perspective of its fitness benefits, but much less is known about the proximate factors that cause plants to reproduce synchronously. In this article, I follow up on more than two decades of research investigating proximate mechanisms of mast seeding by Astragalus scaphoides, an iteroparous perennial forb. We use long-term monitoring in relation to two environmental manipulations to evaluate the importance of exogenous environmental factors versus endogenous feedbacks for synchrony in this species. Our past research showed that synchrony in this species is explained by the pollen-coupling hypothesis: plants that flower synchronously set seed and deplete stored resources, whereas plants that happen to flower asynchronously are pollen limited, set fewer seeds, and do not deplete resources, and flower again until they are resynchronized. Continued monitoring of two experimental manipulations, water addition, and flower removal, provides additional support for this model, and also reveals subtle effects of water availability on synchrony. Water addition decreased flowering, rather than increasing it as expected based on simple correlations with weather variables, suggesting that precipitation does not synchronize reproduction. Exogenous drivers are generally considered to be the primary synchronizing factors in plant reproduction. Our work in this system suggests that endogenous feedbacks may be more important than has been previously assumed.     

Variation in ovule and seed size and associated size-number trade-offs in angiosperms

Unlike pollen and seed size, the extent and causes of variation in ovule size remain unexplored. Based on 45 angiosperm species, we assessed whether intra- and interspecific variation in ovule size is consistent with cost minimization during ovule production or allows maternal plants to dominate conflict with their seeds concerning resource investment. Despite considerable intraspecific variation in ovule volume (mean CV = 0.356), ovule production by few species was subject to a size-number trade-off. Among the sampled species, ovule volume varied two orders of magnitude, whereas seed volume varied four orders of magnitude. Ovule volume varied positively among species with flower mass and negatively with ovule number. Tenuinucellate ovules were generally larger that crassinucellate ovules, and species with apical placentation (which mostly have uniovulate ovaries) had smaller ovules than those with other placentation types. Seed volume varied positively among species with fruit mass and seed development time, but negatively with seed number. Seeds grew a median 93-fold larger than the ovules from which they originated. Our results provide equivocal evidence that selection minimizes ovule size to allow efficient resource allocation after fertilization, but stronger evidence that ovule size affords maternal plants an advantage in parent-offspring conflict.     

Predator satiation and extreme mast seeding in 11 species of Chionochloa (Poaceae)

Variation in annual flowering effort is described for 16 long datasets from 11 species of Chionochloa (Poaceae) in New Zealand. All populations exhibited extreme mast seeding. The most variable species was C. crassiuscula (coefficient of variation, CV=3.02) over 26 years at Takahe Valley, Fiordland, which is the highest published CV we know of worldwide. The other populations also had high CVs (lowest CV=1.42, mean CV=1.84) which were higher than for other well‐studied genera such as Picea, Pinus and Quercus. There were also frequent years of zero flowering (mean across all populations was 37.2% zero years; maximum 53% for C. rubra and C. crassiuscula over 19 years) whereas zero years are rare in other published masting datasets.Flowering was highly synchronous among species within a site (mean r=0.886), and also (though significantly less so) among sites. Among sites, synchrony was not significantly higher within‐species (mean r=0.711) than between‐species (r=0.690). Warm summer temperatures led to heavy flowering the following summer. Flowering synchrony increased with increasing synchrony in local deseasonalised summer temperatures, and decreased with increasing distance between sites.Mast seeding has been shown in Chionochloa to reduce losses to specialist flower or seed predators. Among‐species synchrony may be adaptive if species share a common seed predator. Developing seeds of at least 10 Chionochloa species are attacked by larvae of an undescribed cecidomyiid. In Takahe Valley, where masting is most pronounced, cecidomyiids attacked all six Chionochloa species in all four years studied. Mean annual losses were almost constant (10.0 to 13.4%) while flowering effort varied 100‐fold. The invariant losses are consistent with other evidence that the cecidomyiid may have extended diapause, which would make it harder to satiate by mast seeding. We hypothesise that one possible factor favouring such extremely high levels of mast seeding in Chionochloa is that its seed predator is very hard to satiate.     

Sensitivity to Seed Germination Schedule by Scatter‐Hoarding Pére David's Rock Squirrels During Mast and Non‐Mast Years

Mast seeding by oaks (Quercus) and other seed species has fundamental impacts on the behavior of individual scatter‐hoarding animals as well as their population and community dynamics. Scatter‐hoarding squirrels are highly sensitive to acorn germination schedule and stop germination by removing the embryo of non‐dormant acorns to prevent losses of energy and nutrients by rapid germination. However, we know little about how this behavior is affected by mast seeding. We investigated foraging decisions made by free‐ranging Pére David's rock squirrel (Sciurotamias davidianus) with three species that produce non‐dormant seeds and two species that produce dormant seeds (Fagaceae) during mast and non‐mast years in Central China from 2007–2010. Consistent with the food perishability hypothesis that squirrels make decisions to minimize the perishability of their caches, non‐dormant seeds were hoarded at a lower rate, but had a higher probability of embryo removal than dormant seeds. Compared with non‐mast years, a lower proportion of seeds were harvested during mast years, but a higher proportion of the harvested seeds were hoarded. In addition, the probability of embryo removal in the hoarded seeds was higher during mast years. Squirrels also dispersed seeds at shorter distances during mast years. Moreover, the interactions between mast seeding and seed germination schedule had significant effects on squirrel foraging decisions, including embryo removal. This study is the first to demonstrate that squirrels show greater sensitivity to seed germination potential when seeds are abundant.     

A mast‐seeding desert shrub regulates population dynamics and behavior of its heteromyid dispersers

Granivorous rodent populations in deserts are primarily regulated through precipitation‐driven resource pulses rather than pulses associated with mast‐seeding, a pattern more common in mesic habitats. We studied heteromyid responses to mast‐seeding in the desert shrub blackbrush (Coleogyne ramosissima), a regionally dominant species in the Mojave–Great Basin Desert transition zone. In a 5‐year study at Arches National Park, Utah, USA, we quantified spatiotemporal variation in seed resources in mast and intermast years in blackbrush‐dominated and mixed desert vegetation and measured responses of Dipodomys ordii (Ord's kangaroo rat) and Perognathus flavescens (plains pocket mouse). In blackbrush‐dominated vegetation, blackbrush seeds comprised >79% of seed production in a mast year, but 0% in the first postmast year. Kangaroo rat abundance in blackbrush‐dominated vegetation was highest in the mast year, declined sharply at the end of the first postmast summer, and then remained at low levels for 3 years. Pocket mouse abundance was not as strongly associated with blackbrush seed production. In mixed desert vegetation, kangaroo rat abundance was higher and more uniform through time. Kangaroo rats excluded the smaller pocket mice from resource‐rich patches including a pipeline disturbance and also moved their home range centers closer to this disturbance in a year of low blackbrush seed production. Home range size for kangaroo rats was unrelated to seed resource density in the mast year, but resource‐poor home ranges were larger (P < 0.001) in the first postmast year, when resources were limiting. Blackbrush seeds are higher in protein and fat but lower in carbohydrates than the more highly preferred seeds of Indian ricegrass (Achnatherum hymenoides) and have similar energy value per unit of handling time. Kangaroo rats cached seeds of these two species in similar spatial configurations, implying that they were equally valued as stored food resources. Blackbrush mast is a key resource regulating populations of kangaroo rats in this ecosystem.     

Masting in ponderosa pine: comparisons of pollen and seed over space and time

Many plant species exhibit variable and synchronized reproduction, or masting, but less is known of the spatial scale of synchrony, effects of climate, or differences between patterns of pollen and seed production. We monitored pollen and seed cone production for seven Pinus ponderosa populations (607 trees) separated by up to 28?km and 1,350?m in elevation in Boulder County, Colorado, USA for periods of 4?C31?years for a mean per site of 8.7?years for pollen and 12.1 for seed cone production. We also analyzed climate data and a published dataset on 21?years of seed production for an eighth population (Manitou) 100?km away. Individual trees showed high inter-annual variation in reproduction. Synchrony was high within populations, but quickly became asynchronous among populations with a combination of increasing distance and elevational difference. Inter-annual variation in temperature and precipitation had differing influences on seed production for Boulder County and Manitou. We speculate that geographically variable effects of climate on reproduction arise from environmental heterogeneity and population genetic differentiation, which in turn result in localized synchrony. Although individual pines produce pollen and seed, only one-third of the covariation within trees was shared. As compared to seed cones, pollen had lower inter-annual variation at the level of the individual tree and was more synchronous. However, pollen and seed production were similar with respect to inter-annual variation at the population level, spatial scales of synchrony and associations with climate. Our results show that strong masting can occur at a localized scale, and that reproductive patterns can differ between pollen and seed cone production in a hermaphroditic plant.     

Geographical variation in the spatial synchrony of a forest-defoliating insect: isolation of environmental and spatial drivers

Despite the pervasiveness of spatial synchrony of population fluctuations in virtually every taxon, it remains difficult to disentangle its underlying mechanisms, such as environmental perturbations and dispersal. We used multiple regression of distance matrices (MRMs) to statistically partition the importance of several factors potentially synchronizing the dynamics of the gypsy moth, an invasive species in North America, exhibiting outbreaks that are partially synchronized over long distances (approx. 900 km). The factors considered in the MRM were synchrony in weather conditions, spatial proximity and forest-type similarity. We found that the most likely driver of outbreak synchrony is synchronous precipitation. Proximity played no apparent role in influencing outbreak synchrony after accounting for precipitation, suggesting dispersal does not drive outbreak synchrony. Because a previous modelling study indicated weather might indirectly synchronize outbreaks through synchronization of oak masting and generalist predators that feed upon acorns, we also examined the influence of weather and proximity on synchrony of acorn production. As we found for outbreak synchrony, synchrony in oak masting increased with synchrony in precipitation, though it also increased with proximity. We conclude that precipitation could synchronize gypsy moth populations directly, as in a Moran effect, or indirectly, through effects on oak masting, generalist predators or diseases.     

Quantitative methods for defining mast‐seeding years across species and studies

Abstract: Although there is a quantitative method that is commonly used for identifying mast‐seeding behaviour of a plant population based on the coefficient of variation (i.e. CV is standard deviation/mean>1), there is no general quantitative method for delineating “mast” as opposed to “non‐mast” years. Mast years are, however, described qualitatively as years when “large”, “unusually large” and “high” seed production occurs. The use of a consistent and generally applicable method for delineating mast years across species and plant populations is important for synthesizing knowledge of the causes and consequences of mast seeding, which could be confounded by using different methods among studies. We examine six quantitative methods for identifying mast years: four methods from the literature and two methods developed here. We use 36 seed production datasets covering a variety of species with ≥10 years of data to test the performance of these six methods. For each method, we quantify the percentage of the datasets to which the method could be successfully applied, the magnitude of the mast year relative to the mean, the frequency of mast years and the occurrence of consecutive mast years. The majority of the methods failed to meet the criteria for a suitable method. The best method used the number of standard deviates (standardized deviate method) of the annual mean seed production from the long‐term mean of the dataset to identify mast‐seeding years. General results from the standardized deviate method include that the occurrence of mast‐seeding years is largely unrelated to plant population CV, but similar across species and data collection methods.     

湖北三峡大老岭自然保护区光叶水青冈群落种子雨10年观测: 种子雨密度、物种构成及其与群落的关系

种子雨是植物种子扩散的起点, 对群落更新及种群动态起着关键作用。该文以三峡大老岭自然保护区内一片面积为1.3 hm ²的光叶水青冈(Fagus lucida)群落固定样地为研究对象, 运用分层随机设计, 在10个不同的地形部位放置了100个种子雨收集框, 自2001年起进行种子雨观测, 对该群落种子雨的数量与物种多样性的年际动态、种子雨和群落物种构成的关系等进行了统计分析。结果表明: 1)过去10年间, 共收集到来自48种木本植物的60 926粒种子, 种子雨的多年平均密度为(82.9 ± 61.5) seeds·m^-2·a^-1(mean ± SD), 平均物种丰富度为(16.7 ± 5.5) species·a^-1(mean ± SD)。2)种子生产的种间差异极为显著, 种子量排名前三的植物贡献了累计种子雨总量的70%。3)群落种子雨的密度和物种丰富度在10年中基本同步, 均呈现显著的周期性波动, 并出现了3个大年。乔木和灌木种子雨密度的年际波动无显著相关性, 但物种数变化显著正相关; 4)种子雨与样地群落共有种为23种, 分别占种子雨和群落中木本植物种数的47.92%和54.76%, 但这些共有种贡献了种子总量的96.22%, 表明扩散限制在研究群落中十分显著。与国内其他森林群落种子雨研究结果相比, 该研究群落的种子雨密度明显较低。     

Energy or information? The role of seed availability for reproductive decisions in edible dormice

The edible dormouse is a specialized seed predator which is highly adapted to the fluctuations of food availability caused by mast seeding of beech and oak trees. Dormice produce young just in time with maximum food availability, and can completely skip reproduction in years with a lack of seeding. Because their decision to reproduce or not in any particular year is made long before the ripe seeds are available, it seems that dormice can anticipate the upcoming mast situation. We tested the hypothesis that the presence of high caloric food in spring affects their reproductive decision. Therefore, we supplementary fed dormice in a field experiment from spring to early summer with sunflower seeds, which also contain a high amount of energy. Supplemental feeding caused significant increases in the proportion of reproducing females and reproductively active males. These results suggest that edible dormice may use the occurrence of an energy rich food resource to predict the autumnal mast situation. Further, our data indicate that the decision to reproduce was not the result of an increased body mass due to the consumption of surplus food, but that sufficient seed abundance acts as an environmental signal to which dormice adjust their reproduction.     

The community and ecosystem consequences of intraspecific diversity: a meta‐analysis

Understanding the relationships between biodiversity and ecosystem functioning has major implications. Biodiversity–ecosystem functioning relationships are generally investigated at the interspecific level, although intraspecific diversity (i.e. within‐species diversity) is increasingly perceived as an important ecological facet of biodiversity. Here, we provide a quantitative and integrative synthesis testing, across diverse plant and animal species, whether intraspecific diversity is a major driver of community dynamics and ecosystem functioning. We specifically tested (i) whether the number of genotypes/phenotypes (i.e. intraspecific richness) or the specific identity of genotypes/phenotypes (i.e. intraspecific variation) in populations modulate the structure of communities and the functioning of ecosystems, (ii) whether the ecological effects of intraspecific richness and variation are strong in magnitude, and (iii) whether these effects vary among taxonomic groups and ecological responses. We found a non‐linear relationship between intraspecific richness and community and ecosystem dynamics that follows a saturating curve shape, as observed for biodiversity–function relationships measured at the interspecific level. Importantly, intraspecific richness modulated ecological dynamics with a magnitude that was equal to that previously reported for interspecific richness. Our results further confirm, based on a database containing more than 50 species, that intraspecific variation also has substantial effects on ecological dynamics. We demonstrated that the effects of intraspecific variation are twice as high as expected by chance, and that they might have been underestimated previously. Finally, we found that the ecological effects of intraspecific variation are not homogeneous and are actually stronger when intraspecific variation is manipulated in primary producers than in consumer species, and when they are measured at the ecosystem rather than at the community level. Overall, we demonstrated that the two facets of intraspecific diversity (richness and variation) can both strongly affect community and ecosystem dynamics, which reveals the pivotal role of within‐species biodiversity for understanding ecological dynamics.     

Snow tussocks, chaos, and the evolution of mast seeding

One hitherto intractable problem in studying mast seeding (synchronous intermittent heavy flowering by a population of perennial plants) is determining the relative roles of weather, plant reserves, and evolutionary selective pressures such as predator satiation. We parameterize a mechanistic resource-based model for mast seeding in Chionochloa pallens (Poaceae) using a long-term individually structured data set. Each plant's energy reserves were reconstructed using annual inputs (growing degree days), outputs (flowering), and a novel regression technique. This allowed the estimation of the parameters that control internal plant resource dynamics, and thereby allowed different models for masting to be tested against each other. Models based only on plant size, season degree days, and/or climatic cues (warm January temperatures) fail to reproduce the pattern of autocovariation in individual flowering and the high levels of flowering synchrony seen in the field. This shows that resource-matching or simple cue-based models cannot account for this example of mast seeding. In contrast, the resource-based model pulsed by a simple climate cue accurately describes both individual-level and population-level aspects of the data. The fitted resource-based model, in the absence of environmental forcing, has chaotic (but often statistically periodic) dynamics. Environmental forcing synchronizes individual reproduction, and the models predict highly variable seed production in close agreement with the data. An evolutionary model shows that the chaotic internal resource dynamics, as predicted by the fitted model, is selectively advantageous provided that adult mortality is low and seeds survive for more than 1 yr, both of which are true for C. pallens. Highly variable masting and chaotic dynamics appear to be advantageous in this case because they reduce seed losses to specialist seed predators, while balancing the costs of missed reproductive events.