Non-linear feedback processes and a latitudinal gradient in the climatic effects determine green spruce aphid outbreaks in the UK

The role of climatic fluctuations in determining the dynamics of insect populations has been a classical problem in population ecology. Here, we use long-term annual data on green spruce aphid populations at nine localities in the UK for determining the importance of endogenous processes, local weather and large-scale climatic factors. We rely on diagnostic and modelling tools from population dynamic theory to analyse these long-term data and to determine the role of the North Atlantic Oscillation (NAO) and local weather as exogenous factors influencing aphid dynamics. Our modelling suggests that the key elements determining population fluctuations in green spruce aphid populations in the UK are the strong non-linear feedback structure, the high potential for population growth and the effects of winter and spring weather. The results indicate that the main effect of the NAO on green spruce aphid populations is operating through the effect of winter temperatures on the maximum per capita growth rate (R_m). In particular, we can predict quite accurately the occurrence of an outbreak by using a simple logistic model with weather as a perturbation effect. However, model predictions using different climatic variables showed a clear geographical signature. The NAO and winter temperature were best for predicting observed dynamics toward the southern localities, while spring temperature was a much better predictor of aphid dynamics at northern localities. Although aphid species are characterized by complex life-cycles, we emphasize the value of simple and general population dynamic models in predicting their dynamics.     

Methods for chaotic dynamics in studies of synchrony in complex natural systems

The wavelet transform modulus maxima method was applied to study pairwise synchrony of irregular fluctuations in insect population size in several localities throughout the United Kingdom. The North Atlantic Oscillation index was shown to affect the population dynamics and spatial pairwise synchrony in the green spruce aphid Elatobium abietinum across the United Kingdom. Synchrony between localities was associated with the relative positions of the localities and climatic effects.     

Climate mediated exogenous forcing and synchrony in populations of the oak aphid in the UK

Contemporary population dynamics theory suggests that animal fluctuations in nature are the result of the combined forces of intrinsic and exogenous factors. Weather is the iconic example of an exogenous force. The common approach for analyzing the relationship between population size and climatic variables is by simple correlation or using the climate as an additive covariable in statistical models. Here, we evaluated different functional forms in which climatic variables could influence population dynamics of the oak aphid Tuberculatus annulatus both in each locality and in relation to synchrony between localities. Results indicate that in at least four of eight aphid populations, climate influences population dynamics by modifying the carrying capacity of the system (lateral effect mediated by winter precipitation). Additionally, path analysis showed that synchrony in population dynamics is highly correlated with synchrony in winter precipitation regime, and the spatial scale of both processes is similar, which suggests that this is an example of the Moran effect. Our results show the key effects of precipitation on intra and inter population processes of this aphid. The methods used, mixing population dynamics modelling and test of synchrony, allowed us to connect the direct and indirect effects of exogenous variables into each population with patterns of synchrony inter populations.     

Main determinants of rodent population fluctuations in managed Central European temperate lowland forests

Whilst studies have shown that climatic (North Atlantic Oscillation (NAO)) and biotic (acorn production) factors influence rodent populations, mechanisms driving temporal and spatial fluctuation of rodent populations are understudied. This study evaluates relationships between the influence of environmental factors (biotic and abiotic) and phenotypic characteristics across two rodent feeding guilds (granivorous and non-granivorous species) represented by four species of rodents in Central Europe. We hypothesise that the relationship between acorn density and population growth rate are indirectly affected by climatic factors (winter NAO) and that these effects differ amongst herbivorous and granivorous species. In addition, we also tested whether effects of weather and competition on individual phenotype characteristic vary amongst mast and non-mast years. Rodent populations were estimated by catching individuals in snap traps during the growing season (from March to November) over a period of 9 years at three sites. The results of the generalised linear model provide evidence that acorn production best explained the population fluctuations. We therefore conclude that the between-year population fluctuations in rodent abundance were governed by density dependence and initiated primarily by acorn mast years. Auto-regressive models also revealed direct density dependence in combination with the direct effects of mast years. Therefore, strong intraspecific competition for food is likely in years following mast years. Our results also showed that abundance of non-granivorous species is mainly influenced by local weather conditions which could regulate food quality and abundance. On the other hand, population dynamics of granivorous species are caused directly by acorn density and indirectly by climatic condition influencing acorn production.     

Geographical location, climate and land use influences on the phenology and numbers of the aphid, Myzus persicae, in Europe

Aim The purpose of this study was to improve understanding of the relationship between the spatial patterns of an important insect pest, the aphid Myzus persicae, and aspects of its environment. The main objectives were to determine the predominant geographical, climatic and land use factors that are linked with the aphid's distribution, to quantify their role in determining that distribution, including their interacting effects and to explore the ability of artificial neural networks (ANNs) to provide predictive models. Location The study focused on four spatial scales to account for the aphid data base characteristics and available land use data sets: Europe; a broad zone over Europe covering Belgium, Denmark, France, Ireland, Italy, The Netherlands, Scotland, Sweden and Wales (Regio data base coverage); North‐West Europe (i.e. Belgium, France and the United Kingdom); and England with Wales. Methods Multiple linear regression (MLR) was used to identify the variables in the Geographic location, Climate and Land use groups, that explained significant proportions of the variance in M. persicae total annual numbers and Julian date of first capture. A variance partitioning procedure was used to measure the fraction of the variation that can be explained by each environmental factor and of shared variation between the different factors. Finally, ANNs were employed as an alternative modelling approach for the two largest study areas, i.e. Europe and the Regio data base coverage, to determine whether the relationship between aphid and environmental variables was better described by more complex functions as well as their ability to generalize to new data. Results Land use variables are shown to play a significant role in explaining aphid numbers. The area of agricultural crops, in particular oilseed rape, is positively correlated with M. persicae annual numbers. Among the climatic variables, rainfall is negatively correlated with aphid numbers and temperature is positively correlated. The geographical components also explain a significant part of aphid annual numbers. However, the variance partitioning procedure indicates that while each group has an effect, none is dominant. Aphid first capture is mainly explained by climate where rainfall tends to delay migration and warmer conditions tend to advance it. Climate accounts for the greatest part of the variance when considered separately from the other factors. The geographical and land use components also have a significant effect on first capture at each scale, but their direct contribution is negligible. The ability of the ANN models to generalize to new total numbers and phenological data compared with MLR models was less for Europe (9 and 6% increase in the variance accounted for, respectively) than for the Regio data coverage where an increase of 44% in the variance accounted for was observed. Main conclusions This research supports the hypothesis that climate, land use and geographical location play a role in determining patterns of aphid annual numbers and phenology. The ability of ANN models to predict aphid distribution is improved by the inclusion of temporal land use data. However, identification of the processes involved in such relationships is difficult due to numerous interactions between the environmental factors.     

Population synchrony in small-world networks

Network topography ranges from regular graphs (linkage between nearest neighbours only) via small-world graphs (some random connections between nodes) to completely random graphs. Small-world linkage is seen as a revolutionary architecture for a wide range of social, physical and biological networks, and has been shown to increase synchrony between oscillating subunits. We study small-world topographies in a novel context: dispersal linkage between spatially structured populations across a range of population models. Regular dispersal between population patches interacting with density-dependent renewal provides one ecological explanation for the large-scale synchrony seen in the temporal fluctuations of many species, for example, lynx populations in North America, voles in Fennoscandia and grouse in the UK. Introducing a small-world dispersal kernel leads to a clear reduction in synchrony with both increasing dispersal rate and small-world dispersal probability across a variety of biological scenarios. Synchrony is also reduced when populations are affected by globally correlated noise. We discuss ecological implications of small-world dispersal in the frame of spatial synchrony in population fluctuations.     

Common growth signal and spatial synchrony of the chronologies of tree-rings from pines in the Baltic Sea region over the last nine centuries

The common growth signal and spatial synchrony of nine chronologies of tree rings (Pinus sylvestris L.) from the Baltic Sea region were studied using moving correlations with time windows of 50 and 100 years. Drifts from synchrony to asynchrony and again back to synchrony across several centuries were observed. The chronologies showed higher (or lower) coefficients of correlations and correspondingly gentler (or steeper) declines in spatial synchrony in certain periods. In accordance with research into other ecological phenomena, the origin of the spatial synchrony was found to relate to the intensity of the westerly circulation over the Atlantic and Europe as described by the North Atlantic Oscillation (NAO) index. The tree rings from all of the analysed subregions were found to be wider, identifying faster periods of growth, when there was a positive NAO during the extended winter season from December to March that preceded the period of growth. Previous studies showed that the NAO index can be linked with the growth of tree rings because of factors in the local climate that affect growth, such as temperature and precipitation. Moreover, the spatial synchrony showed multidecadal fluctuations that were correlated with variations of a similar scale in the reconstructed NAO index. However, a high degree of spatial synchrony was observed during the 20^th century in the presence of values of NAO index that were strongly negative. A more detailed view was provided by maps of the correlations in which spatial changes were demonstrated in the growth of pines between the 12^th and the 20^th century. These results could be used as background information in the development of improved proxy-based reconstructions of the NAO index (or other similar meteorological variables), which could also be inferred from tree ring data relating to earlier centuries.     

Interannual dynamics of aerial and arboreal green spruce aphid populations

Partial defoliation of spruce by the green spruce aphid Elatobium abietinum (Walker) is a recurrent event in European and, increasingly, North American forests. The patterns of insect abundance on trees have never been satisfactorily described by a numerical model despite considerable knowledge of endogenous and exogenous factors in the population dynamics of the species. Long-term field population estimates of the aphid on foliage provided the opportunity to evaluate such a model. Unlike comparable models for tree-dwelling aphids, this was also applicable to almost completely independent aphid field data derived from the Rothamsted Insect Survey’s nationwide network of suction traps. Although based on relatively few parameters, the model was robust in its predictions of alate aphids geographically remote from the forest in which the original population was estimated. The population maximum, which causes the greatest forest damage, is reached in early summer and can be predicted from knowledge of winter temperature (chill bouts), spring temperature (thermal sum), and interannual negative feedback (density dependence). The model provides confirmation that alate populations of spruce aphids, upon which a number of other extensive studies have been based, are ultimately influenced by similar endogenous and climatic factors and that they are a reasonable proxy for aphids on trees.     

Increased outbreak frequency associated with changes in the dynamic behaviour of populations of two aphid species

Sudden changes in the variability of natural populations can result in increased likelihood of extinction or in greater frequency and intensity of pest outbreaks. These changes could be associated with changes in some relevant population parameters such as the equilibrium density or the maximum population growth rate. However, changes in these parameters have very different consequences. An increase in equilibrium density results in a higher variance in population fluctuations according to the relationship between mean and variance described by Taylor's power law, but does not modify the stability properties of the system. On the other hand, changes in the maximum growth rate induce changes in the dynamic regimes and stability properties of the population. In this study, using statistical and mathematical methods borrowed from econometrics and engineering, we identify structural changes to the variance in the population dynamics of the sycamore aphid Drepanosiphum platanoidis and the green spruce aphid Elatobium abietinum in the UK. Some localities showed strong changes in their population parameters, such that their dynamic regime changed completely. These changes in the population dynamic regimes increase the expected frequency of outbreaks, which has enormous economic and ecological consequences. Through this study we show the application of methods that could be helpful to pest and wildlife managers in the task of evaluating changes in the risk of outbreaks or extinction of animal populations under changing global environmental scenarios.     

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.     

Climate change and breeding parameters of great and blue tits throughout the western Palaearctic

Increasing evidence suggests that climate change has consequences on avian breeding phenology. Here, variations in laying date and clutch size of great tit Parus major and blue tit Parus caeruleus within and between breeding populations through the western Palaearctic are examined in relation to climatic fluctuations, measured by the winter North Atlantic Oscillation (NAO) index. Within and across breeding sites, laying date was related to winter‐NAO index such that great and blue tit females lay earlier after warmer, moister winters (positive values of winter NAO‐index). The present study shows that for most populations there is an advancement of laying date, but the rate of change with respect to NAO significantly differed geographically across the western Palaearctic and did not differ between species. However, clutch size of great and blue tits was not affected by climatic fluctuations, presumably because the whole season is being shifted, but not in relation to food supplies. These combined analyses for the two species controlled for potentially confounding variables such as latitude, longitude, elevation and habitat of each study site.     

Distribution-wide effects of climate on population densities of a declining migratory landbird

1. Increases in global temperatures have created concern about effects of climatic variability on populations, and climate has been shown to affect population dynamics in an increasing number of species. Testing for effects of climate on population densities across a species' distribution allows for elucidation of effects of climate that would not be apparent at smaller spatial scales. 2. Using autoregressive population models, we tested for effects of the North Atlantic Oscillation (NAO) and the El Ni?o Southern Oscillation (ENSO) on annual population densities of a North American migratory landbird, the yellow-billed cuckoo Coccyzus americanus, across the species' breeding distribution over a 37-year period (1966-2002). 3. Our results indicate that both the NAO and ENSO have affected population densities of C. americanus across much of the species' breeding range, with the strongest effects of climate in regions in which these climate systems have the strongest effects on local temperatures. Analyses also indicate that the strength of the effect of local temperatures on C. americanus populations was predictive of long-term population decline, with populations that were more negatively affected by warm temperatures experiencing steeper declines. 4. Results of this study highlight the importance of distribution-wide analyses of climatic effects and demonstrate that increases in global temperatures have the potential to lead to additional population declines.     

Weather and regional crop composition variation drive spatial synchrony of lepidopteran agricultural pests

1. Spatial synchrony, the tendency for temporal population fluctuations to be correlated across multiple locations at regional scales, is common and contributes to the severity of outbreaks and epidemics, but is little studied in agricultural pests. 2. This study analysed spatial synchrony from 1974 to 2008 in 16 lepidopteran agricultural pests in Maryland, U.S.A., and investigated whether pest synchrony is driven by interannual variability in seasonal weather and the areas planted in different crop types. 3. Lepidopteran pests exhibited high degrees of spatial synchrony, which was driven by environmental variation, a phenomenon known as the Moran effect. Region-wide variation in the areas planted in major crops drove spatially synchronous abundance fluctuations in more than half of studied species. The combination of weather and crop composition explained large fractions of synchrony in black cutworm, corn earworm, European corn borer, and spotted cutworm populations. Other pests, including forage looper and variegated cutworm, displayed a high degree of spatial synchrony, but without dependence on the tested drivers. 4. The study finding that synchronous variation in the area planted in different crop types contributed to synchronous pest abundance fluctuations suggests that strategies to reduce synchrony in changes in crop type across a region could reduce the severity of pest outbreaks and enhance the stability of agricultural systems.     

Interannual population dynamics of the green spruce aphid Elatobium abietinum (Walker) in France

The hypothesis that similar processes govern interannual dynamics of green spruce aphid in the UK and France, is generally supported by the application of a general discrete model. A simple model based on relatively few parameters was able to closely characterise interannual population dynamics from completely independent aerial and arboreal samples of aphids. Long-term field population estimates of the green spruce aphid Elatobium abietinum (Walker) in France have provided the opportunity to select and evaluate the generality of a model, which was developed in the UK to explain the year-to-year variations in peak abundance of the aphid. The objective was to observe the influence of the local climates and disturbing climate factors on the population densities of the insect in two regions of France. The model uses climate variables and aphid population data from regular samples in the two regions that were investigated. A general discrete model was used to predict aphid population densities. The model performed well in tracking the interannual patterns of population but was less likely to predict absolute population density. To improve predictions, further account would need to be taken of additional site-specific climate variables and the strength of overcompensating density dependence. Nevertheless, it is clear that broadly similar processes are at work in the population dynamics of this insect across its biogeographical range.     

Anisotropic patterned population synchrony in climatic gradients indicates nonlinear climatic forcing

Although climatic forcing has been suspected to be the most common cause of spatial population synchrony owing to the Moran effect, it has proved difficult to disentangle the impact of climate from other possible causes of synchrony based on population survey data. Nonlinear population responses to climatic variation may be a part of this difficulty, but they can also provide an opportunity to highlight the climate impacts through targeted survey designs. In particular, when species distribution ranges encompass consistent spatial gradients in climate (e.g. according to latitude or altitude), such gradients can be strategically included in the spatial design of population surveys as to facilitate comparisons of spatial synchrony patterns across and along the gradient. In that case, we predict that nonlinear impacts of climatic variation on population growth rates will result in anisotropic (direction specific) synchrony patterns in the sense that synchrony will drop faster with distance along the climatic gradient than across it. We provide an empirical case study to exemplify survey design and analyses. Of two sympatric species of geometrids, inhabiting an altitudinal gradient in subarctic birch forest, one (Operophtera brumata L.) showed anisotropic synchrony consistent with a strongly nonlinear sensitivity to climatic variation, whereas the other (Epirrita autumnata Bkh.) did not. These results are interpreted in light of the biological characteristics of the species.     

Differing contributions of density dependence and climate to the population dynamics of three eruptive herbivores

1. Although both endogenous and exogenous processes regulate populations, the current understanding of the contributions from density dependence and climate to the population dynamics of eruptive herbivores remains limited. 2. Using a 17‐year time series of three cereal aphid species [Rhopalosiphum padi L., Metopolophium dirhodum (Walker), and Diuraphis noxia (Kurdumov)] compiled from a trapping network spanning the northwestern U.S.A., temporal and spatial patterns associated with population fluctuations, and modelled density dependence in aphid abundances were tested. These models were used to analyse correlations between climate and aphid abundances in the presence and absence of residual variance as a result of density‐dependent effects. 3. The temporal dynamics of aphid population fluctuations indicated periodicity, with no clear evidence for a spatial pattern underlying population fluctuations. 4. Aphid abundances oscillated in a manner consistent with delayed density dependence for all three aphid species, although the strength of these feedbacks differed among species. 5. Diuraphis noxia abundances were negatively correlated with increasing temperatures in the absence of density‐dependent effects, whereas M. dirhodum abundances were positively correlated with increasing cumulative precipitation in the presence of density‐dependent effects; yet, R. padi abundances were unrelated to climate variables irrespective of population feedbacks. 6. Our analysis suggests that endogenous feedbacks differentially regulate aphid populations in the northwestern U.S.A., and these feedbacks may operate at an expansive spatial scale. It is concluded that the contributions of density dependence and climate to aphid population dynamics are species‐specific in spite of similar ecological niches, with implications for assessing species responses to climate variability.     

Searching for mechanisms of synchrony in spatially structured gamebird populations

1.  Time series data on five species of gamebird from the Dolomitic Alps were used to examine the relative importance of dispersal and common stochastic events in causing synchrony between spatially structured populations.
2.  Cross-correlation analysis of detrended time series was used to describe the spatial pattern of fluctuations in abundance, while standardized time series were used to describe both fluctuations and the trend in abundance. There were large variations in synchrony both within and between species and only weak negative relationships with distance.
3.  Species in neighbouring habitats were more likely to be in synchrony than species separated by several habitats. Species with similar density-dependent structure were more likely to be in synchrony.
4.  In order to estimate the relative importance of dispersal and environmental stochasticity, we modelled the spatial dynamics of each species using two different approaches. First, we used estimating functions and bootstrapping of time series data to calculate the relative importance of dispersal and stochastic effects for each species. Second, we estimated the intensity of environmental stochasticity from climatic records during the breeding season and then modelled the dispersal rate and dispersal distance for each species. The two models exhibited similar results for rock ptarmigan, black grouse, hazel grouse and rock partridge, while contrasting patterns were observed for capercaillie.
5.  The results suggest that environmental stochasticity plays the dominant role in synchronizing the fluctuations of these galliform species, although there will also be some dispersal between populations.     

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.     

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.     

Long-term responses in arctic ungulate dynamics to changes in climatic and trophic processes

 Following predictions from climatic general circulation models, the effects of perturbations in global climate are expected to be most pronounced in the Northern Hemisphere. Elaborating on a recently developed plant–herbivore–climate model, we explore statistically how different winter climate regimes and density-dependent processes during the past century have affected population dynamics of two arctic ungulate species. Our analyses were performed on the dynamics of six muskox and six caribou populations. In muskoxen, variation in winter climate, mediated through the North Atlantic Oscillation (NAO), explained up to 24% of the variation in interannual abundance, whereas in caribou up to 16% was explained by the NAO. Muskoxen responded negatively following warm and snowy winters, whereas caribou responded negatively to dry winters. Direct and delayed density dependence was recorded in most populations and explained up to 32% and 90% of variations in abundance of muskoxen and caribou, respectively. Received: November 19, 2001 / Accepted: May 28, 2002