Heterogeneity in the distribution of genetically modified and conventional oilseed rape within fields and seed lots

The implementation of co-existence in the commercialisation of GM crops requires GM and non-GM products to be segregated in production and supply. However, maintaining segregation in oilseed rape will be made difficult by the highly persistent nature of this species. An understanding of its population dynamics is needed to predict persistence and develop potential strategies for control, while to ensure segregation is being achieved, the production of GM oilseed rape must be accompanied by the monitoring of GM levels in crop or seed populations. Heterogeneity in the spatial distribution of oilseed rape has the potential to affect both control and monitoring and, although a universal phenomenon in arable weeds and harvested seed lots, spatial heterogeneity in oilseed rape populations remains to be demonstrated and quantified. Here we investigate the distribution of crop and volunteer populations in a commercial field before and during the cultivation of the first conventional oilseed rape (winter) crop since the cultivation of a GM glufosinate-tolerant oilseed rape crop (spring) three years previously. GM presence was detected by ELISA for the PAT protein in each of three morphologically distinguishable phenotypes: autumn germinating crop-type plants (3% GM), autumn-germinating 'regrowths' (72% GM) and spring germinating 'small-type' plants (17% GM). Statistical models (Poisson log-normal and binomial logit-normal) were used to describe the spatial distribution of these populations at multiple spatial scales in the field and of GM presence in the harvested seed lot. Heterogeneity was a consistent feature in the distribution of GM and conventional oilseed rape. Large trends across the field (50 x 400 m) and seed lot (4 x 1.5 x 1.5 m) were observed in addition to small-scale heterogeneity, less than 20 m in the field and 20 cm in the seed lot. The heterogeneity was greater for the 'regrowth' and 'small' phenotypes, which were likely to be volunteers and included most of the GM plants detected, than for the largely non-GM 'crop' phenotype. The implications of the volunteer heterogeneity for field management and GM-sampling are discussed.     

Spatial synchrony in red grouse population dynamics

Red grouse Lagopus lagopus scoticus populations exhibit unstable dynamics that are often characterised by regular periodic fluctuations in abundance. Time-series' of grouse harvesting records collected from 287 management units (moors) across Scotland, England and Wales were analysed to investigate the broad scale patterns of synchrony in these fluctuations. Estimation of the spatial autocorrelation of grouse population dynamics across moors indicates relatively high levels of synchrony between populations on adjacent moors, but that this synchrony declines sharply with increasing inter-moor distance. At distances of greater than 100  km, grouse population time-series exhibit only weakly positive cross-correlation coefficients. Twenty-eight geographical, environmental and other candidate variables were examined to construct a general linear model to explain variation in local synchrony. Grouse moor productivity (average size of shooting bag), distance from the Atlantic coast moving in a north-easterly direction, April and June temperatures, and June rainfall significantly increased the explanatory power of this model. An understanding of the processes underlying synchrony in red grouse population dynamics is a prerequisite to anticipating the effects of large-scale environmental change on regional patterns of grouse distribution and abundance.     

Spatiotemporal dynamics of Floerkea proserpinacoides (Limnanthaceae), an annual plant of the deciduous forest of eastern North America

Because environmental filters are temporally and spatially heterogeneous, there often is a lack of significant relationship between the spatial patterns of successive life stages in plant populations. In this study, we determined the spatiotemporal relationships between different life stages in two populations of an annual plant of the deciduous forests of eastern North America, Floerkea proserpinacoides. Demographic surveys were done over a 4-yr period, and experiments were performed in the field and under controlled conditions to test for the effects of various environmental factors on population dynamics. There was a general lack of relationship between the spatial patterns of seed bank and seedling density, and a lack of similarity between their spatial correlograms. This was related mostly to the effects of spatially variable environmental filters operating on germination and emergence. However, environmental filters acting on plant survival were stable through time and contributed to stabilize the density and spatial patterns of the populations. Despite density-dependent presenescence mortality, spatial patterns of seedlings and mature individuals were similar and their correlograms were alike, suggesting that mortality did not fully compensate for density. Estimated fecundity was negatively correlated with population density over the study period. Although flower production started only 2-3 wk after emergence, seed maturation mostly occurred at the end of the life cycle, just before the onset of plant senescence. Yet, individual fecundity was low for an annual plant, i.e., 3.0 ± 0.5 mature seeds/plant (mean ± 1 SE). Seed predation by vertebrates was not significant. Low soil moisture had little effect on the total number of seeds germinating, although it slowed down the germination process. In quadrats where leaf litter was experimentally doubled, seedling emergence was lower than in control quadrats; in quadrats where leaf litter was completely removed, emergence did not differ from that in control quadrats. Susceptibility to drought stress was higher for seedlings than for mature plants. Although the species does not maintain a long-term persistent soil seed bank, other factors, such as density-dependent fecundity and autogamy, may temper population fluctuations through time and reduce the probability of local extinction.     

Travelling wave dynamics and self-organization in a spatio-temporally structured population

We analyse spatial population dynamics showing that periodic or period-like chaotic dynamics produce self-organization structures, such as travelling waves. We suggest that self-organized patterns are associated with spatial synchrony patterns that often depend on geographical distance between subpopulations. The population dynamics also show statistical spatial autocorrelation patterns. We contrast our theoretical simulations with empirical data on annual damages in young sapling stands caused by voles. We conclude, on the basis of the periodicity, synchrony, and spatial autocorrelation patterns, and our simulation results, that vole dynamics represent travelling waves in population dynamics. We suggest that because such synchrony patterns are frequently observed in natural populations, spatial self-organization may be more common in population dynamics than reported in the literature.     

Mapping of QTL for seed dormancy in a winter oilseed rape doubled haploid population

Following winter oilseed rape cultivation, considerable numbers of volunteer oilseed rape plants may occur in subsequent years in following crops. The appearance of volunteer oilseed rape plants is based on the capability of the seeds to become secondary dormant and to survive in this stage for many years in the soil. Genetic reduction of secondary seed dormancy in oilseed rape could provide a means to reduce the frequency of volunteer plants and especially the dispersal of transgenic oilseed rape. The objective of the present study was to analyse the inheritance of primary and secondary seed dormancy in a winter oilseed rape doubled haploid population derived from the cross Express 617 × R53 and to study correlations to other seed traits. Field experiments were performed in Germany for 2 years at two locations with two replicates. Seeds harvested from open pollinated plants were used for all analyses, including a laboratory test for seed dormancy. A previously developed molecular marker map of the doubled haploid population was used to map QTL of the relevant traits. For primary, secondary and total seed dormancy, the results showed significant effects of the genotypes and their interactions, with years and locations. Two, four and five QTL were detected for primary, secondary and total seed dormancy which explained 19, 35 and 42 % of the phenotypic variance, respectively. Results show that secondary seed dormancy is a heritable trait and that selection for low secondary seed dormancy is possible.     

Population synchrony and stability in environmentally forced metacommunities

A general prediction from simple metapopulation models is that spatially synchronized forcing can spatially synchronize population dynamics and destabilize metapopulations. In contrast, spatially asynchronous forcing is predicted to decrease population synchrony and promote temporal stability and population persistence, especially in the presence of dispersal. Only recently have studies begun to experimentally address these predictions. Moreover, few studies have experimentally examined how such processes operate in the context of competition communities. Stabilizing processes may continue to operate when placed within a metacommunity context with multiple competing consumers but only at low to intermediate levels of dispersal. High dispersal rates can reverse these predictions and lead to destabilization. We tested this under controlled conditions using an experimental aquatic system composed of three competing species of zooplankton. Metacommunities experienced different levels of dispersal and environmental forcing in the form of spatially synchronous or asynchronous pH perturbations. We found support that dispersal can have contrasting effects on population stability depending on the degree to which population dynamics were synchronized in space. Dispersal under synchronous forcing or no forcing had either neutral of positive effects on spatial population synchrony of all three zooplankton species. In these treatments, dispersal reduced population stability at the local and metapopulation levels for two of three species. In contrast, asynchronously varying environments reduced population synchrony relative to unforced systems, regardless of dispersal level. In these treatments, dispersal enhanced temporal stability and persistence of populations not by reducing population synchrony but by enhancing population minima and spatial averaging of abundances. High dispersal rates under asynchronous forcing reduced the abundance of one species, consistent with increasing regional competition and general metacommunity theory. However, no effects on its stability or persistence were observed. Our work highlights the context‐dependent effects of dispersal on population dynamics in varying environments.     

Spatial dynamics of Microtus vole populations in continuous and fragmented agricultural landscapes

Small mammal populations often exhibit large-scale spatial synchrony, which is purportedly caused by stochastic weather-related environmental perturbations, predation or dispersal. To elucidate the relative synchronizing effects of environmental perturbations from those of dispersal movements of small mammalian prey or their predators, we investigated the spatial dynamics of Microtus vole populations in two differently structured landscapes which experience similar patterns of weather and climatic conditions. Vole and predator abundances were monitored for three years on 28 agricultural field sites arranged into two 120-km-long transect lines in western Finland. Sites on one transect were interconnected by continuous agricultural farmland (continuous landscape), while sites on the other were isolated from one another to a varying degree by mainly forests (fragmented landscape). Vole populations exhibited large-scale (>120 km) spatial synchrony in fluctuations, which did not differ in degree between the landscapes or decline with increasing distance between trapping sites. However, spatial variation in vole population growth rates was higher in the fragmented than in the continuous landscape. Although vole-eating predators were more numerous in the continuous agricultural landscape than in the fragmented, our results suggest that predators do not exert a great influence on the degree of spatial synchrony of vole population fluctuations, but they may contribute to bringing out-of-phase prey patches towards a regional density level. The spatial dynamics of vole populations were similar in both fragmented and continuous landscapes despite inter-landscape differences in both predator abundance and possibilities of vole dispersal. This implies that the primary source of synchronization lies in a common weather-related environment.     

Weeds in fields with contrasting conventional and genetically modified herbicide-tolerant crops. II. Effects on individual species

We compared the effects of the management of genetically modified herbicide-tolerant (GMHT) and conventional beet, maize and spring oilseed rape on 12 weed species. We sampled the seedbank before and after cropping. During the season we counted plants and measured seed rain and biomass. Ratios of densities were used to calculate emergence, survival, reproduction and seedbank change. Treatments significantly affected the biomass of six species in beet, eight in maize and five in spring oilseed rape. The effects were generally consistent, with biomass lower in GMHT beet and spring oilseed rape and higher in GMHT maize. With few exceptions, emergence was higher in GMHT crops. Subsequent survival was significantly lowered for eight species in beet and six in spring oilseed rape in the GMHT treatments. It was increased for five species in maize and one in spring oilseed rape. Significant effects on seedbank change were found for four species. However, for many species in beet and spring oilseed rape (19 out of 24 cases), seed densities were lower in the seedbank after GMHT cropping. These differences compounded over time would result in large decreases in population densities of arable weeds. In maize, populations may increase.     

Spatial autocorrelation of ecological phenomena

Ecological variables often fluctuate synchronously over wide geographical areas, a phenomenon known as spatial autocorrelation or spatial synchrony. Development of statistical approaches designed to test for spatial autocorrelation combined with the increasing accessibility of long-term, large-scale ecological datasets are now making it possible to document the patterns and understand the causes of spatial synchrony at scales that were previously intractable. These developments promise to foster significant future advances in understanding population regulation, metapopulation dynamics and other areas of population ecology.     

Synchrony in brown trout, Salmo trutta, population dynamics: a 'Moran effect' on early-life stages

Synchrony among populations (i.e. spatial covariation in temporal fluctuations of population size or growth rate) is a common feature to many animals. Both large-scale autocorrelated climatic factors (the 'Moran effect') and dispersal between populations are candidates to explain synchrony, although their relative influence is difficult to assess. Only a few investigations have reported patterns of synchrony among freshwater populations, and even fewer directly related these patterns to an environmental variable. In the present study, we analysed the spatio-temporal patterns of fluctuation of 57 brown trout populations widespread across France, each sampled continuously during 5 years. We compared the respective influence of connectivity and stream distance within basins (i.e. that potentially allow a basin-scale dispersal) and environmental factors (hydrological and air temperature variables, available for 37 sites) on the synchrony of brown trout cohort densities (0+, 1+ and adults). A series of Mantel tests revealed that the degree of synchrony was not related to connectivity or stream distance between sites, suggesting no effect of dispersal at the basin-scale. The degree of synchrony among sites for the 0+ fish was significantly related to the degree of hydrological synchrony (based on high flows during the emergence period). For all three age classes, the synchrony in the temperature patterns did not explain synchrony in trout dynamics. Our results allow us to discuss the respective influence of dispersal and climatic factors on the spatio-temporal patterns of trout dynamics at the basin scale.     

The Pursuit of Population Collapses: Long-Term Dynamics of Semi-Domestic Reindeer in Sweden

Spatiotemporal reindeer population fluctuations are a result of multiple factors that working in concert affecting the structure and functioning of many Arctic and Sub-Arctic ecosystems. We investigated the population dynamics of Swedish semi-domestic reindeer from 1945 to 2012 at the reindeer herding district-level (Sameby) to identify possible population collapses or declines. We also explored spatial trends and synchrony in population dynamics across latitudinal-longitude gradients for two distinct periods: 1945–1965 and 1995–2012 but found no evidence of large-scale reindeer population declines and no visible synchrony across adjacent populations. Our findings were unexpected as both reindeer populations and the pastoral lifestyle face increased habitat loss, predation, fragmentation and climate change.     

Baseline susceptibility and insecticide resistance monitoring in European populations of Meligethes aeneus and Ceutorhynchus assimilis collected in winter oilseed rape

Pollen beetle, Meligethes aeneus (Fabricius) (Coleoptera: Nitidulidae), and cabbage seed weevil, Ceutorhynchus assimilis (Paykull) (Coleoptera: Curculionidae), are important pests in the production of European winter oilseed rape, Brassica napus L. (Brassicaceae), which is grown on several million hectares in Europe. Insecticide treatments are common to control both pests once they exceed economic damage thresholds; however, not many chemical classes are available for their control in European oilseed rape. Particularly pollen beetles recently developed high levels of pyrethroid resistance impairing field control at recommended rates in many countries, whereas no resistance is yet reported to another important insecticide, thiacloprid. The major objective of this study was to investigate the spatio‐temporal susceptibility status of pollen beetle against the recently introduced insecticide thiacloprid. From 2009 to 2012 more than 630 populations of pollen beetle collected in 13 countries were monitored for resistance to thiacloprid by using an adult vial test. No shifting to lower susceptibility of pollen beetle to thiacloprid has been observed between 2009 and 2012. Furthermore, we were able to show that pollen beetle larvae are extremely susceptible to thiacloprid, whereas within strains larvae are significantly more resistant than adults to pyrethroids such as lambda‐cyhalothrin. Dose‐response data for thiacloprid against cabbage seed weevil populations collected in 2011 in Germany, Sweden, and Ukraine showed a 10‐fold higher intrinsic sensitivity compared to pollen beetle, and showed only a low variation in response. In addition, we also tested 17 cabbage seed weevil populations collected in five countries against lambda‐cyhalothrin with low variation in response (three‐fold), suggesting full baseline susceptibility and no resistance to pyrethroids. The implications of the data presented for resistance management in coleopteran pests in winter oilseed rape are discussed.     

Similarity in seasonal flow regimes,not regional environmental classifications explain synchrony in brown trout population dynamics in France

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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.     

Are bark beetle outbreaks less synchronous than forest Lepidoptera outbreaks?

Comparisons of intraspecific spatial synchrony across multiple epidemic insect species can be useful for generating hypotheses about major determinants of population patterns at larger scales. The present study compares patterns of spatial synchrony in outbreaks of six epidemic bark beetle species in North America and Europe. Spatial synchrony among populations of the Eurasian spruce bark beetle Ips typographus was significantly higher than for the other bark beetle species. The spatial synchrony observed in epidemic bark beetles was also compared with previously published patterns of synchrony in outbreaks of defoliating forest Lepidoptera, revealing a marked difference between these two major insect groups. The bark beetles exhibited a generally lower degree of spatial synchrony than the Lepidoptera, possibly because bark beetles are synchronized by different weather variables that are acting on a smaller scale than those affecting the Lepidoptera, or because inherent differences in their dynamics leads to more cyclic oscillations and more synchronous spatial dynamics in the Lepidoptera.     

Spatially autocorrelated disturbances and patterns in population synchrony

Spatially synchronous population dynamics have been documented in many taxa. The prevailing view is that the most plausible candidates to explain this pattern are extrinsic disturbances (the Moran effect) and dispersal. In most cases disentangling these factors is difficult. Theoretical studies have shown that dispersal between subpopulations is more likely to produce a negative relationship between population synchrony and distance between the patches than perturbations. As analyses of empirical data frequently show this negative relationship between the level of synchrony and distance between populations, this has emphasized the importance of dispersal as a synchronizing agent. However, several weather patterns show spatial autocorrelation, which could potentially produce patterns in population synchrony similar to those caused by dispersal. By using spatially extended versions of several population dynamic models, we show that this is indeed the case. Our results show that, especially when both factors (spatially autocorrelated perturbations and distance-dependent dispersal) act together, there may exist groups of local populations in synchrony together but fluctuating asynchronously with some other groups of local populations. We also show, by analysing 56 long-term population data sets, that patterns of population synchrony similar to those found in our simulations are found in natural populations as well. This finding highlights the subtlety in the interactions of dispersal and noise in organizing spatial patterns in population fluctuations.     

Synchrony and geographical variation of the spruce bark beetle (<Emphasis Type="Italic">Ips typographus</Emphasis>) during a non-epidemic period

Spatio-temporal analyses of non-epidemic bark beetle populations may provide insight in dynamics predisposing for outbreaks. The present article presents a spatio-temporal analysis of the population dynamics of Ips typographus based on pheromone trap data from southeast and mid-Norway in the post-epidemic period 1979–2002. The analyses include regression analyses, hierarchical cluster analysis, and analysis of spatial synchrony of beetle time series and climatic data by means of nonparametric spatial covariance functions. The mean abundance of beetles declined linearly with latitude. In addition, the time series means were higher in areas with high forest productivity and rocky soils predisposed to drought. The time series patterns differed significantly between northern and southern study areas. The regional synchrony of the time series was fairly high (0.38), indicating that some large-scale climatic factor may influence the dynamics. Windfelling was the external variable showing the most parallel pattern of correlation to the beetle dynamics. We thus posit that large windfall events may be a major instigator and synchronizer of beetle outbreaks in areas subjected to regionalized weather systems.     

Measuring and modelling anthropogenic secondary seed dispersal along roadverges for feral oilseed rape

Seed dispersal of feral crop plants along roadverges is likely to be influenced by numerous anthropogenic vectors in the agroecosystem. Within the context of introducing genetically modified (GM) cultivars, long-distance dispersal of feral seeds associated with the growth of GM feral populations (via a selective advantage due to transgene expression) could make these populations become invasive. Their expansion could then favour the spread of transgenes and modify the composition of roadverge plant communities. Because quantitative data on anthropogenic seed dispersal along roadverges were few, we estimated effective secondary dispersal for oilseed rape, the seeds of which are not adapted to dispersal by wind or biotic agents. A seed deposition experiment showed that secondary dispersal did not systematically occur along roadverges, was correlated with traffic intensity and was local. Low traffic intensity and anthropogenic disturbances (covering of seeds by mown grasses and burial by farming machinery) prevented dispersal on three of the experimental sites. Along a road with higher traffic, secondary dispersal occurred (d_max=21.5 m), probably induced by wind turbulence behind vehicles. The best-fitting dispersal kernel was a mixture of two components: 20% of seeds dispersing over a few metres on average and 80% remaining in the original place. Expansion rates of feral populations of GM herbicide-tolerant oilseed rape were computed using an invasion model and this kernel. They were low (1–4 m yr⁻¹) when only ballistic and/or secondary dispersal were included but higher (4–20 m yr⁻¹) when theoretically rare events of long-distance dispersal by verge mowers were added. This study suggests that secondary seed dispersal is unlikely to have a significant impact on the spread of GM feral oilseed rape populations in highly disturbed and dynamic habitats such as roadverges. Detecting long-distance dispersal events induced by other vectors (e.g. mowers) would require integrative approaches based on genetic and spatial data.     

Estimation of Plant Demographic Parameters from Stage‐Structured Censuses

Summary This article presents some statistical methods for estimating the parameters of a population dynamics model for annual plants. The model takes account of reproduction, immigration, seed survival in a seed bank, and plant growth. The data consist of the number of plants in several developmental stages that were measured in a number of populations for a few consecutive years; they are incomplete since seeds could not be counted. It is assumed that there are no measurement errors or that measurement errors are binomial and not frequent. Some statistical methods are developed within the framework of estimating equations or Bayesian inference. These methods are applied to oilseed rape data.     

The extended Moran effect and large-scale synchronous fluctuations in the size of great tit and blue tit populations

1. Synchronous fluctuations of geographically separated populations are in general explained by the Moran effect, i.e. a common influence on the local population dynamics of environmental variables that are correlated in space. Empirical support for such a Moran effect has been difficult to provide, mainly due to problems separating out effects of local population dynamics, demographic stochasticity and dispersal that also influence the spatial scaling of population processes. Here we generalize the Moran effect by decomposing the spatial autocorrelation function for fluctuations in the size of great tit Parus major and blue tit Cyanistes caeruleus populations into components due to spatial correlations in the environmental noise, local differences in the strength of density regulation and the effects of demographic stochasticity. 2. Differences between localities in the strength of density dependence and nonlinearity in the density regulation had a small effect on population synchrony, whereas demographic stochasticity reduced the effects of the spatial correlation in environmental noise on the spatial correlations in population size by 21.7% and 23.3% in the great tit and blue tit, respectively. 3. Different environmental variables, such as beech mast and climate, induce a common environmental forcing on the dynamics of central European great and blue tit populations. This generates synchronous fluctuations in the size of populations located several hundred kilometres apart. 4. Although these environmental variables were autocorrelated over large areas, their contribution to the spatial synchrony in the population fluctuations differed, dependent on the spatial scaling of their effects on the local population dynamics. We also demonstrate that this effect can lead to the paradoxical result that a common environmental variable can induce spatial desynchronization of the population fluctuations. 5. This demonstrates that a proper understanding of the ecological consequences of environmental changes, especially those that occur simultaneously over large areas, will require information about the spatial scaling of their effects on local population dynamics.