Measuring mast seeding behavior: relationships among population variation,individual variation and synchrony

Mast seeding, or masting, is the variable production of flowers, seeds, or fruit across years more or less synchronously by individuals within a population. A critical issue is the extent to which temporal variation in seed production over a collection of individuals can be viewed as arising from a combination of individual variation and synchrony among individuals. Studies of masting typically quantify such variation in terms of the coefficient of variation (CV). In this paper we examine mathematically how the population CV relates to the mean individual CV and synchrony, concluding that the relationship is a complex one which cannot isolate an overall measure of synchrony, and involves additional factors, principally the number of plants sampled and the mean productivity per plant. Our development suggests some simple approximate relationships of population CV to individual variability, synchrony and the number of individuals. These were found to fit quite well when applied to data from 59 studies which included seed production at the individual level.     

Mechanisms inducing spatially extended synchrony in mast seeding: The role of pollen coupling and environmental fluctuation

Trees in mature forests often show intermittent reproduction. Intensive flowering and seed production occur only once in several years (mast seeding), often synchronized over a long distance. Recently, coupled map models for the dynamics of individual energy reserves have been adopted to explain the phenomena. Even in a constant environment, the trees show a large between-year fluctuation in seed crops and the reproduction can be synchronized over the whole forest if the fruit production is limited by the availability of outcross pollen (pollen coupling). The model with local coupling in which trees are coupled by pollen exchange only with the neighbors shows that a strong synchronization of tree reproduction can develop over the whole forest that may be orders of magnitude larger than the distance of direct pollen exchange between trees. However, their fluctuation is close to the period-two oscillation, and is unable to explain observed intermittent reproduction of a longer interval between mast years. Finally the effect of common environmental fluctuation experienced by different individuals is studied, when the annual productivity and the reproductive threshold of trees fluctuate between years. In the absence of pollen limitation, environmental fluctuation correlated strongly between individuals (Moran effect) failed to produce a high positive correlation in seed production between individuals. If both pollen limitation and correlated environmental fluctuation are at work, a significantly large correlation was maintained. Hence, both pollen coupling and common environmental fluctuation are needed to explain synchronized reproduction with intervals longer than 2years.     

Saccharomyces paradoxus and Saccharomyces cerevisiae are associated with exudates of North American oaks

Genetic hybridization and karyotypic analyses revealed the biological species Saccharomyces paradoxus and Saccharomyces cerevisiae in exudates from North American oaks for the first time. In addition, two strains collected from elm flux and from Drosophila by Phaff in 1961 and 1952 were reidentified as S. paradoxus. Each strain studied showed a unique profile of chromosomal hybridization with a probe for the retrotransposable element Ty1. The wild distribution of natural Saccharomyces sensu stricto yeasts is discussed.     

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.     

Mechanisms of synchrony in the North American fireflyPhotinus carolinus (Coleoptera: Lampyridae)

Photometric recordings combined with computer stimulation, acquisition and analysis were used to study synchrony in the North American fireflyPhotinus carolinus. A computer-generated burst of simulatedP. carolinus flashes was used to trigger a firefly flash burst. We found that the first triggered firefly flashes occurred after the second or third flashes in the stimulus burst, that there were fewer flashes in a triggered burst than a spontaneous burst, and that extending the stimulus flashes into the firefly's interburst interval inhibited firefly flashing. When the stimulus flash interval (389–560 ms) was changed, no change was seen in the interflash interval. When the stimulus flash interval was changed, the average time between stimulus flash and firefly flash (flash delay) changed as if the firefly interflash interval was constant. Thus, interflash interval inP. carolinus does not change its length, making it similar to the Southeast Asian synchronizerPteroptyx cribellata and different fromPteroptyx malaccae, which can change its interval. We suspect that the time between bursts is functionally analogous to the time between flashes in Southeast Asian synchronizers.     

The occurrence of synchrony in the North American fireflyPhotinus carolinus (Coleoptera: Lampyridae)

Synchronous flashing has been described for some Southeast Asian fireflies but has rarely been reported in North American fireflies. Our field observations indicated thatPhotinus carolinus flash synchronously. The flash pattern of individualP. carolinus was characterized by a burst of five to eight flashes over a period of approximately 4 s. These flash bursts were repeated about every 12 s. Groups of fireflies comprised individuals exhibiting this species-specific flash pattern. Remarkably, members of the group flashed synchronously: Flash bursts started and stopped at the same time and the flashes among individuals occurred at the same time as well. We used low-light level videography to examine this behavior in caged groups of (3 or 10) fireflies for synchronic flashing. The occurrence of concurrent rhythmic group flashing satisfies the criteria for synchrony as defined by Buck (1988). The intermittent nature of the bursts of flashes shown byP. carolinus makes this a discontinuous synchrony. The mechanisms underlying discontinuous synchrony are not known.     

Dispersal and spatial scale affect synchrony in spatial population dynamics

A large body of theoretical studies has shown that synchrony among populations is critical for the long-term persistence of species in fragmented habitats. Although the effects of dispersal and environmental factors on synchrony have been investigated theoretically, empirical studies of these relationships have been lacking. We explored the interplay between environmental and demographic factors (fecundity, survival, dispersal) on population synchrony for 53 species of birds. We show that the interspecific differences in mean synchrony were determined by global environmental factors whose action was probably mediated by the abundance of each species. After removing the effect of these global factors on synchrony, the residual synchrony was strongly correlated with dispersal distance. The relationship between dispersal and synchrony was stronger for the species nesting in wet habitats than for those nesting in dry habitats. Our results indicate that different factors synchronize bird populations at different spatial scales, thus highlighting the role of scale in understanding spatial population dynamics and extinction risks.     

Local variations in spatial synchrony of influenza epidemics

Background

Understanding the mechanism of influenza spread across multiple geographic scales is not complete. While the mechanism of dissemination across regions and states of the United States has been described, understanding the determinants of dissemination between counties has not been elucidated. The paucity of high resolution spatial-temporal influenza incidence data to evaluate disease structure is often not available.

Methodology and Findings

We report on the underlying relationship between the spread of influenza and human movement between counties of one state. Significant synchrony in the timing of epidemics exists across the entire state and decay with distance (regional correlation = 62%). Synchrony as a function of population size display evidence of hierarchical spread with more synchronized epidemics occurring among the most populated counties. A gravity model describing movement between two populations is a stronger predictor of influenza spread than adult movement to and from workplaces suggesting that non-routine and leisure travel drive local epidemics.

Conclusions

These findings highlight the complex nature of influenza spread across multiple geographic scales.     

A five-year study of mast seeding inPinus densiflora

I examined the annual pattern of seed production in three Korean populations ofPinus densiflora from 1998 to 2002. During the fall season, data were collected on the numbers of cones per tagged tree, as well as seed counts, their mass, and seed set following dehiscence. Dbh was measured for individual trees during each spring. Overall, 80.3% of the trees produced seed cones in even-numbered years compared with 51.6% in odd-numbered years. Moreover, trees that matured seed cones in even years produced 12 times more seeds per tree than those in odd years. Tree diameters generally increased significantly during the summers after odd-numbered years, although the trees at one site showed a consistent growth rate each year after 2000. Annual variations in individual seed mass, seed set, and the proportion of damaged seed cones occurred largely within odd or even years. Based on these results, the following conclusions were made: 1) seed-cone production ofP. densiflora is somewhat synchronized, but is not an all-or-nothing phenomenon, 2)P. densiflora tends to undergo mast seeding at two-year intervals, and 3) this alternate-year excess in seed crop may have evolved to increase fitness, though the exact evolutionary mechanism is unclear.     

Predicting spatial occurrence of beetles and pseudoscorpions in hollow oaks in southeastern Sweden

We modelled presence/absence per tree of beetles and pseudoscorpions living in tree hollows in relation to trunk circumference, habitat openness, and connectivity (= density of hollow oaks in the surrounding area), using data from 281 oaks. The presence/absence models were then used to predict species’ occurrences in a county (11,600 km²) in southeastern Sweden. For eight of the nine species, the most parsimonious occupancy model included a positive relationship with connectivity and at least one tree characteristic. Occupancy underestimates from occurrence records—the ratio of the area of occupancy based on our predictive model to the area of occupancy based on occurrence records—varied between 3 and 83 among species when using occurrence records up to 1993, with significantly larger underestimates for smaller beetle species. Today (after extensive surveys), underestimation has decreased to 1.3–25, confirming that calculations solely based on species occurrence records greatly underestimate the area of occupancy. We suggest this should be taken into account to a greater extent and in a clearer way than today when constructing red lists. The radius of the connectivity measure that generated the best fit varied between 135 and 2,857 m among species, with longer distances for more threatened species. Consequently, preservation of the most threatened species (Elater ferrugineus and Tenebrio opacus) requires conservation efforts at larger spatial scales than required to protect Osmoderma eremita, which frequently has been used as an indicator and umbrella species.     

Bootstrap methods for estimating spatial synchrony of fluctuating populations

We describe and examine methods for estimating spatial correlations used in population ecology. We base our analyses on a hypothetical example of a species that has been censured at 30 different locations for 20 years. We assume that the population fluctuations can be described by a simple linear model on logarithmic scale. Stochastic simulations is utilized to check how seven different ways of resampling perform when the goal is to find nominal 95% confidence intervals for the spatial correlation in growth rates at given distances. It turns out that resampling of locations performs badly, with true coverage level as low as 30–40%, especially for small correlations at long distances. Resampling of timepoints performs much better, with coverage varying from 80 to 90%, depending on the strength of density regulation and whether the spatial correlation is estimated for the response variable or for the error terms in the model. Assuming that the underlying model is known, the best results are achieved for parametric bootstrapping, a result that strongly emphasize the importance of defining and estimating a proper population model when studying spatial processes.     

Arthropods in tropical oaks: differences in their spatial distributions within tree crowns

The arthropod activity in the upper and the lower zone of nine emergent oak trees (genus Quercus) of a submontane forest in Sabah, Borneo, Malaysia, was investigated by using flight interception traps, yellow colour traps, and arboreal pitfall traps. Number of arthropods did not differ significantly between the upper and the lower region of the tree crowns. Nevertheless, the results reveal significant differences in the abundances of several arthropod groups between the two zones. In the lower part of the oak crowns Homoptera and ants (mostly alates) were sampled in higher numbers, whereas the upper zone was dominated by Diptera and Hymenoptera (ants excluded). Beetles and most other groups showed no significant differences in abundance between the lower and upper zone of the oak trees. Thus, the arthropods of the investigated community are not evenly distributed. The arthropod community in tree crowns, which often has been treated as a homogenous unit, is in fact divided into subunits formed by differently composed arthropod assemblages.     

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.     

Generalizations of the Moran effect explaining spatial synchrony in population fluctuations

The Moran effect for populations separated in space states that the autocorrelations in the population fluctuations equal the autocorrelation in environmental noise, assuming the same linear density regulation in all populations. Here we generalize the Moran effect to include also nonlinear density regulation with spatial heterogeneity in local population dynamics as well as in the effects of environmental covariates by deriving a simple expression for the correlation between the sizes of two populations, using diffusion approximation to the theta-logistic model. In general, spatial variation in parameters describing the dynamics reduces population synchrony. We also show that the contribution of a covariate to spatial synchrony depends strongly on spatial heterogeneity in the covariate or in its effect on local dynamics. These analyses show exactly how spatial environmental covariation can synchronize fluctuations of spatially segregated populations with no interchange of individuals even if the dynamics are nonlinear.     

Alternative strategies of seed predator escape by early-germinating oaks in Asia and North America

Early germination of white oaks is widely viewed as an evolutionary strategy to escape rodent predation; yet, the mechanism by which this is accomplished is poorly understood. We report that chestnut oak Quercus montana (CO) and white oak Q. alba (WO) (from North America), and oriental cork oak Q. variabilis (OO) and Mongolian oak Q. mongolica (MO) (from Asia) can escape predation and successfully establish from only taproots. During germination in autumn, cotyledonary petioles of acorns of CO and WO elongate and push the plumule out of the cotyledons, whereas OO and MO extend only the hypocotyls and retain the plumule within the cotyledons. Experiments showed that the pruned taproots (>6 cm) of CO and WO acorns containing the plumule successfully germinated and survived, and the pruned taproots (≥12 cm) of OO and MO acorns without the plumule successfully regenerated along with the detached acorns, thus producing two seedlings. We argue that these two distinct regeneration morphologies reflect alternative strategies for escaping seed predation.     

Using mechanistic models to understand synchrony in forest insect populations: the North American gypsy moth as a case study

In many forest insects, subpopulations fluctuate concurrently across large geographical areas, a phenomenon known as population synchrony. Because of the large spatial scales involved, empirical tests to identify the causes of synchrony are often impractical. Simple models are, therefore, a useful aid to understanding, but data often seem to contradict model predictions. For instance, chaotic population dynamics and limited dispersal are not uncommon among synchronous forest defoliators, yet both make it difficult to achieve synchrony in simple models. To test whether this discrepancy can be explained by more realistic models, we introduced dispersal and spatially correlated stochasticity into a mechanistic population model for the North American gypsy moth Lymantria dispar. The resulting model shows both chaotic dynamics and spatial synchrony, suggesting that chaos and synchrony can be reconciled by the incorporation of realistic dynamics and spatial structure. By relating alterations in model structure to changes in synchrony levels, we show that the synchrony is due to a combination of spatial covariance in environmental stochasticity and the origins of chaos in our multispecies model.     

Experimental investigation on weather cues for mast seeding of Fagus crenata

Although a recent study has suggested that the minimum temperature from late April to mid-May in the year preceding flowering causes mast seeding in Fagus crenata, no direct evidence is available to support this finding. The aim of the present investigation was, therefore, to test – in a field experiment – whether the minimum temperature determines mast seeding in F. crenata. We examined the effect of nighttime temperatures on flower-bud initiation in F. crenata by enclosing fruit-bearing branches in heated bags at night, thereby maintaining average nighttime temperatures of approximately 2°C above the ambient temperatures. Heating was applied at night from 21 April to 20 May, 21 May to 19 June, and 21 April to 19 June in 2001. Female inflorescence initiation was inhibited by the nocturnal heating in the period 21 April to 20 May and 21 May to 19 June. However, nocturnal heating from 21 April to 20 May was the more important based on the odds ratio of the former being much lower than that of the latter in a logistic regression model. Male inflorescence initiation was also inhibited by nocturnal heating from 21 April to 20 May. We therefore conclude that flower-bud initiation in F. crenata was controlled by nighttime temperatures between 21 April and 20 May.     

Genetic variation across VNTR loci in central North American Taraxacum surveyed at different spatial scales

Dandelions (Taraxacum officinale Weber(sensu lato); Asteraceae) have been introduced to NorthandSouth America with human migration from Europe. While potential sourcepopulations have both sexually and obligate asexually (agamospermous)reproducing lineages, apparently only the latter have successfully colonizedtheAmericas. The consequences of obligate agamospermy on dandelion populationgenetic diversity in North America remain little explored. Here we use fourdifferent synthetic DNA probes that reveal genetic markers at multiplevariable-number-tandem-repeat (VNTR) loci to examine patterns of geneticvariation among plants collected along three different central North Americantransects with plants (21 to 22 individuals per transect) separated by: 1) >2 m and < 60 m (short transect); 2) > 5km and < 30 km (medium transect); and 3) > 30km and < 340 km (long transect). The mean numberofVNTR markers revealed per plant was 59.3. Co-clonal individuals (proportion ofbands shared exceeding 90%) were found in each transect, with the indexof clonality (the percent of co-clonal individuals detected in a transect)ranging from 34.12% for the short transect to 18.65% for the longtransect. Co-clonal individuals were separated by up to 200 km.With redundant examples of co-clonal individuals removed, mean similarity(proportion of band sharing) of distinct genotypes within transects was 0.426,and no statistical differences in level of similarity between transects, norindication of genetic differentiation between transects, was detected (meanF_st between transect levels with all individuals included =0.05). These results indicate: 1) that dandelion genetic diversity ofcolonizinglineages in central North America is moderately high and does not reflectextreme bottleneck effects shown by some colonizing species; and 2) thatdandelion seed dispersal can be very effective in maintaining similar levels ofgenetic diversity at the different scales of sampling in this study, withcertain clones maintaining numerous, widespread individuals. Evidence that VNTRmutation is detectable within dandelion clonal lineages is presented,demonstrating that clonal families with lines increasinglydifferentiated from one another will continually evolve, and that Muller'srachet is, in all likelihood, turning for asexual lines.