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.     

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.     

Frequency and intensity of drought stress alters the population size and dynamics of Elatobium abietinum on Sitka spruce

The green spruce aphid, Elatobium abietinum, is the most important defoliating pest of Sitka spruce, Picea sitchensis, in the UK. Populations are expected to increase in response to predicted climate change, therefore placing Sitka spruce under increased risk of widespread and severe defoliation. The effect of spring–summer drought stress on E. abietinum population dynamics and development over multiple years was assessed in a field experiment under five different drought treatments with differing intensities and frequencies. The impact on host tolerance, in terms of needle retention, was also investigated. No differences in the length of time taken to reach peak population size were observed. Despite this, E. abietinum populations were found to respond positively to low amplitude intermittent stress, with high densities maintained for longer after the peak. Spring peak densities of aphids did not differ among drought levels, although an autumn peak was observed consistently over 2 years on trees subjected to continuous severe drought. This suggested advancement in the onset of Sitka spruce dormancy. Aphid infestation significantly increased percentage needle loss under all drought treatments, although no differences were observed between drought levels. An interaction between aphid presence and drought treatment was, however, observed during a second year of drought where no aphids were present. The study herein presented has contributed to the understanding of E. abietinum population responses under a changing climate. The implications for damage to host Sitka spruce are of relevance to forest management strategies, as an increase in drought events are predicted in the UK.     

Reproduction of an arboreal aphid pest,Elatobium abietinum,is altered under drought stress

Climate change in the UK is predicted to increase both winter temperatures and the frequency of summer drought events. Elatobium abietinum, the green spruce aphid, is the most important defoliating pest of Sitka spruce, Picea sitchensis, a conifer very widely used in British forest industry. This aphid is expected to respond strongly to altered climate, with changes to population densities leading to more frequent serious outbreaks and defoliation. The impact of simulated spring–summer drought on the reproductive performance of E. abietinum was investigated under laboratory conditions. Rates were assessed under five drought treatments of differing frequencies and intensities to characterize the direction of responses under different drought scenarios, and in time‐staggered trials to explore seasonal variation. Variation in the response of reproduction to water deficit was mediated by drought frequency and magnitude. Low‐amplitude, moderate intermittent stress improved reproductive rates, while severe stress, both continuous and high‐amplitude intermittent, had a detrimental impact when compared with observations made on well‐watered controls. Season was also found to modify the response, with improvements to plant nutritional quality under high‐amplitude stress reflected by improving reproduction. Despite this, no differences in rates were found during the autumn, suggesting no advancement in spruce dormancy under drought. Drought stress therefore has the potential to alter E. abietinum population densities, structure and phenology in Sitka spruce plantations, with implications for forest management, damage levels and natural control of the aphid under future altered climate.     

Northern Atlantic Oscillation effects on the temporal and spatial dynamics of green spruce aphid populations in the UK

1. The role of climate variability in determining the spatial and temporal patterns of numerical fluctuations is a central problem in ecology. The influence of the North Atlantic Oscillation (NAO) index on the population dynamics and spatial synchrony of the green spruce aphid Elatobium abietinum across the UK was shown. 2. Fifteen overlapping time series within the UK were analysed; we used nonparametric models for determining the feedback nonlinear structure and the climatic effects. The spatial synchrony of these populations and the relationship between synchrony and NAO was estimated. 3. From the 15 time series across the UK, 11 showed positive and significant NAO effects. In most of the cases the NAO effects were nonlinear showing strong negative effects of low values. The NAO variation improve the explained variance of the first-order feedback models in 14.5%; ranging from 0% to 48%. All data showed strong-nonlinear (concave) feedback structure. In most of the localities the explained variance by the first-order feedback was about 50-60%. 4. The spatial synchrony of the per capita growth rates and residuals is high across long distances for those populations affected by NAO. The correlation function predicts a spatial scale of synchrony of about 350-400 km for NAO influenced populations. 5. We think that simple population theoretical models describing the link between NAO fluctuations and green spruce aphid dynamics may be fundamental for predicting and simulating the consequences of different climatic scenarios of the future.     

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.     

A reappraisal of flight regulation in the green spruce aphid, Elatobium abietinum

Alate formation and its importance in the population dynamics of Elatobium abietinum is examined. It is concluded that alate formation is related to high soluble nitrogen levels in Sitka spruce needles. In N.E. Scotland the proportion of the population developing as alatae was related to aphid density. The apparently differing status of alatae in S. England may be related to differences in the physiology of the host caused by heat input. In Scotland alate formation may cause a decline in spruce aphid populations when densities are very high but, at the densities normally encountered in Scotland and Germany only small proportions of the populations become alate. In these areas the population collapses in late June or early July occur in response to low soluble nitrogen levels in the spruce needles. The role of alate adults in the formation of new colonies following migration to uncolonised hosts is discussed.     

Influence of the green spruce aphid on defoliation and radial stem growth of Sitka spruce

Premature abscission of older Sitka spruce foliage on trees within forests in Northern Ireland was compared with green spruce aphid population levels 2 to 3 months earlier. Each aphid has a lower relative impact on needle loss at higher aphid population density; nevertheless, more aphids on a tree results in greater rates of needle loss. Evidence from horizontal stem increment sequences suggested that aphid feeding inhibited incremental growth, and that this inhibition might be delayed for up to two field seasons. However, the effect of aphids on stem growth in most years and on most trees seems relatively small compared with the potential of other tree-or year-specific processes.     

Genetic diversity of the green spruce aphid (Elatobium abietinum Walker) in north-west Europe

Abstract 1 The Random Amplified Polymorphic DNA (RAPD) method was used to investigate genetic diversity of anholocyclic populations of the green spruce aphid, Elatobium abietinum Walker, in north‐west Europe. 2 The results showed that the aphid in this region was divided genetically into three major groups. Aphids from the British Isles and north‐west France comprised the first group, the second group consisted of aphids from Denmark and Iceland, and the third group consisted of aphids from Norway. 3 The results indicated a significant level of gene flow within and between sites and geographical regions, especially in the British Isles and north‐west France. Lateral migration of the aphid and/or sexual reproduction is likely to have facilitated the gene flow. 4 The implications of these findings on management of the green spruce aphid are discussed.     

Combined effect of ENSO and SAM on the population dynamics of the invasive yellowjacket wasp in central Chile

The population dynamics of the yellowjacket wasp (Vespula germanica Fabricus) in central Chile were analyzed for the first time. Using a simple Ricker logistic model and adding the effects of local weather variables (temperature and precipitation) and large-scale climate phenomena as El Niño Southern Oscillation (ENSO) and the Southern Annular Mode (SAM), we modeled the interannual fluctuations in nest density. The best model according to the Bayesian information criterion (BIC) included 1-year-lag negative feedback combined with the positive additive effects of ENSO and SAM. According to this model, yellowjacket nest density was favored by warm and dry winters, which probably influenced the survival of overwintering queens. Large-scale climatic variables [Southern Oscillation Index (SOI) and SAM] described the effect of exogenous factors in wasp fluctuations better than local weather variables did. Our results emphasize the usefulness of climate indices and simple theoretical-based models in insect ecological research.     

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.     

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.     

Temperature change and complex dynamics

Density-dependent factors, such as population growth rate and migration, influence dynamic behaviour in ecological models. Temperature, an abiotic and density-independent factor, is also an important determinant of insect population growth. We investigated the endogenous dynamics of a density-dependent response-surface model that included temperature, based on time series for two aphid species. We investigated the effects of temperature and random noise on the model dynamics. In most cases, an increase in temperature resulted in a higher predicted equilibrium density; it could induce complex dynamics. Noise at the level of the natural variation in temperature resulted in extinctions in some models. Our results from these models indicate that aphid populations might become more abundant, and less stable in some circumstances, if there is climate warming. Received: 25 November 1996 / Accepted: 30 June 1997     

Individual growth rates do not predict aphid population densities under altered atmospheric conditions

• 1 Altered atmospheric composition, associated with climate change, can modify herbivore population dynamics through CO₂ and/or O₃‐mediated changes in plant quality.
• 2 Although pea aphid Acyrthosiphon pisum genotypes exhibit intraspecific variation in population growth in response to atmospheric composition, the proximate mechanisms underlying this variation are largely unknown.
• 3 By rearing single (green, pink) and mixed (green + pink) pea aphid genotypes on red clover Trifolium pratense at the Aspen Free Air CO₂ and O₃ Enrichment (Aspen FACE) site, we assessed whether: (i) elevated CO₂ and/or O₃ concentrations alter aphid growth and development and (ii) individual aphid growth rates predict aphid population densities.
• 4 We showed that growth and development of individual green and pink aphids were not influenced by CO₂ and/or O₃ concentrations when reared as individual or mixed genotypes. Individual growth rates, however, did not predict population densities.
• 5 Reared as a single genotype, green pea aphid populations decreased in response to elevated CO₂ concentrations, but not in response to elevated CO₂ + O₃ concentrations. Pink pea aphid populations reared as a single genotype were unaffected by augmented CO₂ or O₃. Populations of mixed genotypes, however, were reduced under elevated CO₂ concentrations, irrespective of O₃ concentrations.
• 6 Herbivore population sizes may not readily be predicted from growth rates of individual organisms under atmospheric conditions associated with global climate change.

     

Climate change and the fate of cereal aphids in Southern Britain

Climate change will drive dramatic changes in the abundance and distribution of species. Assessing the impacts of climate change on our agricultural systems is essential for mitigation planning. Here, I combine projections from the UK Hadley Centre's HadRM03 climate change model for Southern Britain with a general mechanistic model of the interaction between climate, temperate grass physiology and cereal aphid population dynamics. Aphids are one of the largest and most important groups of crop pests and disease vectors worldwide but particularly in temperate regions. The model predicts an increasingly dramatic decline in cereal aphid abundance from the 1961 to 1990 baseline with increasing CO₂ emissions: low emissions=?5%, medium low=?12%, medium high=?61%, and high=?92%. Of the six climate variables used in the model, changes in temperature and rainfall were the most important across all emissions scenarios and, counter‐intuitively, the direct impact of elevated CO₂ actually declines as emissions increase. The results suggest that the pest status of cereal aphids in Southern Britain will significantly decline by the end of this century.     

Assessment of the relative impact of different natural enemies on population dynamics of the grain aphid Sitobion avenae in the field

1. A detailed population dynamics model was devised to provide a tool for integrated pest management against the cereal aphid Sitobion avenae on winter wheat. 2. This model allowed investigation of the relative impact of different natural enemies on aphid population dynamics. 3. The output of the model was compared with a set of data collected in the western part of France from 1976 to 1986. 4. Fungal diseases accounted for 75% of the reduction in peak aphid density and were the key factor acting on aphid dynamics in this region. 5. This study highlights the importance of detailed population dynamics modelling of keystone species, like aphids, for elucidation of the relations between the keystone species and other species associated in the ecosystem.     

A population dynamics perspective on effect of irrigation regimes and plant densities on greenbug abundance in grain sorghum using regression

• 1 A dataset generated from previous experiments on greenbug Schizaphis graminum (Rondani) (Hemiptera: Aphididae) response to irrigation and plant density in grain sorghum was reanalyzed using a recently‐developed mechanistic ecological model for describing aphid population density curves. The model was used to estimate seven response variables: observed peak aphid abundance, predicted peak aphid abundance, time of peak abundance, per capita birthrate, death rate coefficient, final cumulative density and duration of substantial aphid infestation across three irrigation regimes and five plant densities.
• 2 Using regression, the observed peak aphid abundance, predicted peak aphid abundance, per capita birthrate and final cumulative abundance were shown to decrease significantly, whereas the death rate coefficient and duration of the infestation were shown to increase significantly for each 100 000 plant/ha increase.
• 3 Although significant results were found for a number of variables generated from the specific data set used in the analyses, of perhaps greater importance is the potential use of these equations in future predictions of aphid population dynamics. An example of projecting population curves based on estimated peak and cumulative counts and an example of projecting population curves based on estimated birth and death rate coefficients are provided.

     

Population responses of a conifer-dwelling aphid to seasonal changes in its host

Abstract.  1. Current evidence suggests that seasonal changes in spruce needle sap nutrients have a decisive influence on green spruce aphid ( Elatobium abietinum ) population density, but the mechanisms of population change, the roles of development rate, fertility and mortality, and the existence of density-dependent processes, are not clearly understood.
2. Experimental studies of aphid populations were conducted in controlled environments to estimate seasonal patterns in aphid mean relative growth rate, prenatal development, fertility, and mortality. Studies were also made of the effect of aphid crowding on vital rates.
3. Independent of the degree of aphid crowding, seasonal changes in the amino acid concentration of needle sap were tracked by aphid growth rate, fertility (and adult size), but not by rates of aphid mortality. The most pronounced change in vital rates, and the one most likely to drive seasonal population change, was in fertility. Prenatal development time actually became shorter in periods when nutrients were scarce, but the resulting adult aphids were smaller and less fertile than during periods of improved nutrition.
4. Density dependence of vital rates was only observed during mid-summer when nutrients were least available. Mortality, growth rate, and prenatal development were the most strongly density-dependent processes. In contrast, there was no evidence that fertility rates were likely to respond to crowding.
5. There were no important differences between populations reared on small, potted spruce trees and those on plantation trees aged 25 years. This gives confidence that demographic data from a variety of field and laboratory sources could be used to compile data appropriate for population models.     

Shifting daylength regimes associated with range shifts alter aphid‐parasitoid community dynamics

With climate change leading to poleward range expansion of species, populations are exposed to new daylength regimes along latitudinal gradients. Daylength is a major factor affecting insect life cycles and activity patterns, so a range shift leading to new daylength regimes is likely to affect population dynamics and species interactions; however, the impact of daylength in isolation on ecological communities has not been studied so far. Here, we tested for the direct and indirect effects of two different daylengths on the dynamics of experimental multitrophic insect communities. We compared the community dynamics under “southern” summer conditions of 14.5‐hr daylight to “northern” summer conditions of 22‐hr daylight. We show that food web dynamics indeed respond to daylength with one aphid species (Acyrthosiphon pisum) reaching much lower population sizes at the northern daylength regime compared to under southern conditions. In contrast, in the same communities, another aphid species (Megoura viciae) reached higher population densities under northern conditions. This effect at the aphid level was driven by an indirect effect of daylength causing a change in competitive interaction strengths, with the different aphid species being more competitive at different daylength regimes. Additionally, increasing daylength also increased growth rates in M. viciae making it more competitive under summer long days. As such, the shift in daylength affected aphid population sizes by both direct and indirect effects, propagating through species interactions. However, contrary to expectations, parasitoids were not affected by daylength. Our results demonstrate that range expansion of whole communities due to climate change can indeed change interaction strengths between species within ecological communities with consequences for community dynamics. This study provides the first evidence of daylength affecting community dynamics, which could not be predicted from studying single species separately.     

Trophic regulation of postdiapause ovariole maturation inCoccinella septempunctata [Col.: Coccinellidae]

In 1977 the vernal maturation ofCoccinella septempunctata L. females was investigated in relation to the variation in aphid population density on alfalfa and cereals in central Bohemia. The ovarioles do not ripen unless the aphid population density reaches a certain threshold. This leads to considerable variability in the time of vernal reproductive activity (up to 1.5 months in the same geographic locality) among subpopulations living on crops with different aphid densities. This mechanism enables the reproduction of the aphidophagous insect to synchronize with the population development of the aphid.