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

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

Spatial analysis of gypsy moth populations in Sardinia using geostatistical and climate models

• 1 Spatial fluctuations of the Sardinian population of the gypsy moth Lymantria dispar (L.) (Lepidoptera: Lymantriidae) were characterized using geostatistical and climate models. Data on gypsy moth egg mass abundance recorded at 282 permanent monitoring sites from 1980 to 2004 were incorporated in a geographic information system with the vegetational, geomorphological and pedological features of the sites.
• 2 Statistical analyses revealed that the relative outbreak frequency was related to the predominant host tree, slope and elevation of the monitoring sites, whereas there was no correlation between outbreak frequency and exposure and soil type.
• 3 By using bioclimatic modelling, probability maps of gypsy moth outbreaks were generated. The model identified a probability surface with climatic conditions favourable to gypsy moth outbreaks and thus potentially subject to defoliation. The maps included 92 sites where outbreaks never occurred, suggesting that the Sardinian climate may not be a determinant factor for gypsy moth outbreaks.
• 4 The geostatistical method cokriging with outbreak frequency as a covariate was found to be the most suitable technique to estimate gypsy moth egg mass abundance. Semivariograms showed spatial correlation of egg mass abundance within the range 18.5–53 km. The results obtained were used to create regional gypsy moth distribution maps by cokriging, which demonstrated the outbreak foci and different infestation levels at each monitoring area. These results can help to delimit the treatment areas and develop rational gypsy moth management programmes.

     

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

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

What causes outbreaks of the gypsy moth in North America?

The gypsy moth has been present in North America for more than 100 years, and in many of the areas where it has become established outbreaks occur with varying degrees of periodicity. There also exists extensive spatial synchrony in the onset of outbreaks over large geographic regions. Density-dependent mortality clearly limits high-density populations, but there is little evidence for strong regulation of low-density populations. Predation by small mammals appears to be the major source of mortality affecting low-density populations, but because these are generalist predators and gypsy moths are a less preferred food item, mammals do not appear to regulate populations in a density-dependent fashion. Instead, predation levels appear to be primarily determined by small mammal abundance, which is in turn closely linked to the production of acorns that are a major source of food for overwintering predator populations. Mast production by host oak trees is typically variable among years, but considerable spatial synchrony in masting exists over large geographic areas. Thus, it appears that the temporal and spatial patterns of mast production may be responsible for the episodic and spatially synchronous behavior of gypsy moth outbreaks in North America. This multitrophic relationship among mast, predators, and gypsy moths represents a very different explanation of forest insect outbreak dynamics than the more widely applied theories based upon predator–prey cycles or feedbacks with host foliage quality. Received: September 8, 1999 / Accepted: September 20, 2000     

Lepidoptera‐specific insecticide used to suppress gypsy moth outbreaks may benefit non‐target forest Lepidoptera

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Persistence of invading gypsy moth populations in the United States

Exotic invasive species are a mounting threat to native biodiversity, and their effects are gaining more public attention as each new species is detected. Equally important are the dynamics of exotic invasives that are previously well established. While the literature reports many examples of the ability of a newly arrived exotic invader to persist prior to detection and population growth, we focused on the persistence dynamics of an established invader, the European gypsy moth (Lymantria dispar) in the United States. The spread of gypsy moth is largely thought to be the result of the growth and coalescence of isolated colonies in a transition zone ahead of the generally infested area. One important question is thus the ability of these isolated colonies to persist when subject to Allee effects and inimical stochastic events. We analyzed the US gypsy moth survey data and identified isolated colonies of gypsy moth using the local indicator of spatial autocorrelation. We then determined region-specific probabilities of colony persistence given the population abundance in the previous year and its relationship to a suite of ecological factors. We observed that colonies in Wisconsin, US, were significantly more likely to persist in the following year than in other geographic regions of the transition zone, and in all regions, the abundance of preferred host tree species and land use category did not appear to influence persistence. We propose that differences in region-specific rates of persistence may be attributed to Allee effects that are differentially expressed in space, and that the inclusion of geographically varying Allee effects into colony-invasion models may provide an improved paradigm for addressing the establishment and spread of gypsy moth and other invasive exotic species.     

Spatial analysis of harmonic oscillation of gypsy moth outbreak intensity

Outbreaks of many forest-defoliating insects are synchronous over broad geographic areas and occur with a period of approximately 10 years. Within the range of the gypsy moth in North America, however, there is considerable geographic heterogeneity in strength of periodicity and the frequency of outbreaks. Furthermore, gypsy moth outbreaks exhibit two significant periodicities: a dominant period of 8–10 years and a subdominant period of 4–5 years. In this study, we used a simulation model and spatially referenced time series of outbreak intensity data from the Northeastern United States to show that the bimodal periodicity in the intensity of gypsy moth outbreaks is largely a result of harmonic oscillations in gypsy moth abundance at and above a 4 km² scale of resolution. We also used geographically weighted regression models to explore the effects of gypsy moth host-tree abundance on the periodicity of gypsy moths. We found that the strength of 5-year cycles increased relative to the strength of 10-year cycles with increasing host tree abundance. We suggest that this pattern emerges because high host-tree availability enhances the growth rates of gypsy moth populations.     

Geographic variation in density-dependent dynamics impacts the synchronizing effect of dispersal and regional stochasticity

Explanations for the ubiquitous presence of spatially synchronous population dynamics have assumed that density-dependent processes governing the dynamics of local populations are identical among disjunct populations, and low levels of dispersal or small amounts of regionalized stochasticity (Moran effect) can act to synchronize populations. In this study we used historical spatially referenced data on gypsy moth (Lymantria dispar) outbreaks to document that density-dependent processes can vary substantially across geographical landscapes. This variation may be due in part to geographical variation in habitat (e.g., variation in forest composition). We then used a second-order log-linear stochastic model to explore how inter-population variation in density-dependent processes affects synchronization via either synchronous stochastic forcing or dispersal. We found that geographical variation in direct density-dependence (first order) greatly diminishes synchrony caused by stochasticity but only slightly decreases synchronization via dispersal. Variation in delayed density-dependence (second order) diluted synchrony caused by regional stochasticity to a lesser extent than first-order variation, but it did not have any influence on synchrony caused by dispersal. In general, synchronization caused by dispersal was primarily dependent upon the instability of populations and only weakly, if at all, affected by similarities in density-dependence among populations. We conclude that studies of synchrony should carefully consider both the nature of the synchronizing agents and the pattern of local density-dependent processes, including how these vary geographically.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

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.     

Interpopulation quality in gypsy moths with implications for success of two pupal parasitoids: Brachymeria intermedia (Nees) and Coccygomimus turionellae (L.)*

Abstract.

• 1 Gypsy moth egg masses were collected from innocuous, release and outbreak populations and reared in the laboratory on synthetic diet under identical conditions.
• 2 Outbreak population gypsy moths hatched sooner, were smaller and less fecund than innocuous or release gypsy moths, but had a higher concentration of total carbohydrates in their haemolymph.
• 3 Pupae from each population source were submitted to parasitization by two pupal parasitoids. Emerging B.intermedia, an established parasitoid of the gypsy moth associated with outbreak populations, were largest on outbreak source gypsy moths. C. turionellae, not a usual parasitoid of the gypsy moth, were largest when emerging from innocuous or release population gypsy moths. Implications for population dynamics of the gypsy moth are discussed.

     

Natural enemies and environmental factors affecting the population dynamics of the gypsy moth

The population densities of the gypsy moth (Lymantria dispar; Lepidoptera: Lymantriidae) may reach outbreak levels that pose considerable economic and environmental impacts to forests in Europe, Asia, Africa and North America. Compared with the situation in its native European range feeding damage by gypsy moth is often found to be more severe in North America and other parts of the world. Thus, the release from natural enemies can be interpreted as an important cause for high feeding damages. Natural enemies, especially parasitoids, can cause delayed density‐dependent mortality, which may be responsible for population cycles. In North America where only few parasitoids have been introduced and the parasitism rates are considerably lower than in Europe, generalist predators play a larger role than in Europe. Many other factors seem to influence the population dynamics of the gypsy moth such as the host plants and weather. Nevertheless, much of the variability in population densities of the gypsy moth may be attributed to interacting effects of weather conditions and attack by natural enemies. In spite of the considerable number of studies on the ecology and population dynamics of the gypsy moth and the impact of their natural enemies, more quantitative information is required to predict the population dynamics of this pest species and to control its economic and ecologic impact.     

Predators ofLymantria dispar (Lep. lymantriidae) egg masses: Spatio-temporal variation of their impact during the 1988–89 pest generation in the mamora cork oak forest (Morocco)

In the Mamora cork oak forest (Morocco), oophagous predators of the gypsy mothLymantria dispar (L.) (Lepidoptera: Lymantriidae) are one of the main mortality factors influencing pest population dynamics. The predators destroy egg masses more by disrupting their cohesiveness than by predation. From 1987 to 1990, the impact of oophagous enemies significantly contributed to the collapse of a localized gypsy moth outbreak. Variation of egg mass dislocation intensity was greater between trees than among the different strata of a tree. The number of gypsy moth egg masses and egg predator attacks increased when oaks were large and unhealthy. Forest degradation probably explains why egg mass destruction rates were so high (60 to 90% of the eggs) in the infested forest. Oophagous predators find food and shelter under the dehiscent bark of unhealthy cork oaks.     

Long-distance dispersal of the gypsy moth (Lepidoptera: Lymantriidae) facilitated its initial invasion of Wisconsin

Gypsy moth (Lymantria dispar L.) spread is dominated by stratified dispersal, and, although spread rates are variable in space and time, the gypsy moth has invaded Wisconsin at a consistently higher rate than in other regions. Allee effects, which act on low-density populations ahead of the moving population that contribute to gypsy moth spread, have also been observed to be consistently weaker in Wisconsin. Because a major cause of an Allee effect in the gypsy moth is mate-finding failure at low densities, supplementing low-density populations with immigrants that arrive through dispersal may facilitate establishment and consequent spread. We used local indicator of spatial autocorrelation methods to examine space-time gypsy moth monitoring data from 1996 to 2006 and identify isolated, low-density colonies that arrived through dispersal. We measured the distance of these colonies from the moving population front to show that long-distance dispersal was markedly present in earlier years when Wisconsin was still mainly uninfested. Recently, however, immigrants arriving through long-distance dispersal may no longer be detected because instead of invading uninfested areas, they are now supplementing high-density colonies. In contrast, we observed no temporal pattern in the distance between low-density colonies and the population front in West Virginia and Virginia. We submit that long-distance dispersal, perhaps facilitated through meteorological mechanisms, played an important role in the spread dynamics of the initial Wisconsin gypsy moth invasion, but it currently plays a lesser role because the portion of Wisconsin most susceptible to long-distance immigrants from alternate sources is now heavily infested.     

Geographically partitioned spatial synchrony among cyclic moth populations

Many species of forest lepidopterans exhibit regular population cycles, which culminate in outbreak densities at approximately ten-year intervals. Population peaks and mass outbreaks typically occur synchronously and may lead to extensive forest damages over large geographic areas. Here, we report patterns of spatial synchrony among cyclic autumnal moth ( Epirrita autumnata ) populations across Fennoscandia, as inferred from 24 long-term (10–33 years) data sets. The study provides the first formal analysis of spatial synchrony of this pest species which damages mountain birch ( Betula pubescens ssp. czerepanovii ) forests in the sub Arctic. We detected positive cross-correlations in population growth rates between the time series, indicating overall spatial synchrony. However, we found the strongest degree of synchrony within geographically and climatically distinct regional clusters, into which time series were partitioned using cluster analyses. Within regional clusters, moth populations were exposed to the synchronizing effects of common, spatially autocorrelated environmental conditions, i.e. a Moran effect. Consequently, we conclude that a geographically and climatically restricted Moran effect, perhaps interacting with dispersal, is the most likely explanation for the regionally partitioned pattern of synchrony among autumnal moth populations in Fennoscandia. Our results emphasize that high amounts of environmental variation may result in a clear structuring of spatial synchrony at unexpectedly small scales.     

The potential for Entomophaga maimaiga to regulate gypsy moth Lymantria dispar (L.) (Lepidoptera: Erebidae) in Europe

Gypsy moth, Lymantria dispar L., is one of the most important pests of deciduous trees in Europe. In regular cycles, it causes large‐scale defoliation mostly of oak, Quercus spp., forests. Government authorities in the most infested countries in Europe conduct large‐scale applications of pesticides against gypsy moth. In 1999, a new natural enemy, the entomopathogenic fungus Entomophaga maimaiga, was successfully introduced into a gypsy moth population in Bulgaria. Recent investigations suggest that now E. maimaiga is quickly spreading in Europe. Herein, past studies are reviewed regarding this fungus with special emphasis on its potential for becoming an important factor regulating gypsy moth populations in Europe, focusing on the host's population dynamics in relation to the fungus, the influence of environmental conditions on fungal activity, the influence of E. maimaiga on the native entomofauna, including other gypsy moth natural enemies, and spread of the fungus. Based on this analysis, the potential of E. maimaiga for providing control in European gypsy moth populations is estimated.     

Geographical variation in the periodicity of gypsy moth outbreaks

The existence of periodic oscillations in populations of forest Lepidoptera is well known. While information exists on how the periods of oscillations vary among different species, there is little prior evidence of variation in periodicity within the range of a single Lepidopteran species. The exotic gypsy moth is an introduced foliage-feeding insect in North America. Its populations are characterized by high amplitude oscillations between innocuously low densities and outbreak levels during which large regions of forests are defoliated. These outbreaks are recognized to arise periodically with considerable synchrony across much of the gypsy moth's North American range. Our analysis indicates that gypsy moth outbreaks in North America are periodic but they exhibit two dominant periodicities: a primary period of 8–10 yr (as previously reported) and a secondary period of 4–5 yr (a new finding in this study). The outbreak periodicity varied geographically and this variation was associated with forest type. We found that in the most susceptible forest types, those on xeric sites where oak is often mixed with pines, outbreak periodicity had a more dominant 5-yr period while in forest types characteristic of more mesic sites where oak was mixed with maples and other species, cycles were clearly operating on a 10-yr period.     

Spatial patterns at host and forest stand scale and population regulation of the pine processionary moth Thaumetopoea pityocampa

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Pathogen-Driven Outbreaks in Forest Defoliators Revisited: Building Models from Experimental Data

Models of outbreaks in forest-defoliating insects are typically built from a priori considerations and tested only with long time series of abundances. We instead present a model built from experimental data on the gypsy moth and its nuclear polyhedrosis virus, which has been extensively tested with epidemic data. These data have identified key details of the gypsy moth-virus interaction that are missing from earlier models, including seasonality in host reproduction, delays between host infection and death, and heterogeneity among hosts in their susceptibility to the virus. Allowing for these details produces models in which annual epidemics are followed by bouts of reproduction among surviving hosts and leads to quite different conclusions than earlier models. First, these models suggest that pathogen-driven outbreaks in forest defoliators occur partly because newly hatched insect larvae have higher average susceptibility than do older larvae. Second, the models show that a combination of seasonality and delays between infection and death can lead to unstable cycles in the absence of a stabilizing mechanism; these cycles, however, are stabilized by the levels of heterogeneity in susceptibility that we have observed in our experimental data. Moreover, our experimental estimates of virus transmission rates and levels of heterogeneity in susceptibility in gypsy moth populations give model dynamics that closely approximate the dynamics of real gypsy moth populations. Although we built our models from data for gypsy moth, our models are, nevertheless, quite general. Our conclusions are therefore likely to be true, not just for other defoliator-pathogen interactions, but for many host-pathogen interactions in which seasonality plays an important role. Our models thus give qualitative insight into the dynamics of host-pathogen interactions, while providing a quantitative interpretation of our gypsy moth-virus data.     

Distribution of the entomopathogenic fungus Entomophaga maimaiga (Entomophthorales: Entomophthoraceae) at the northern edge of its range in Europe

The gypsy moth Lymantria dispar is a serious economic pest in European broadleaf forests. However, the entomopathogenic fungus Entomophaga maimaiga, which has a great potential to regulate gypsy moth numbers, has recently spread in the Central and Eastern European area of the moth's range. In the current study, 39 plots in oak forests in the Slovak Republic and the Czech Republic were monitored for E. maimaiga from 2014 to 2016. These plots were located along the northern edge of the E. maimaiga range where gypsy moth outbreaks have occurred in the past. The fungus was detected in 16 of the 39 plots. The results thus confirm that E. maimaiga is quite widespread along the northern edge of its range in Europe and can be considered to be established in that area.     

The Allee effect, stochastic dynamics and the eradication of alien species

Previous treatments of the population biology of eradication have assumed that eradication can only be achieved via 100% removal of the alien population. However, this assumption appears to be incorrect because stochastic dynamics and the Allee effect typically contribute to the extinction of very low‐density populations. We explore a model that incorporates Allee dynamics and stochasticity to observe how these two processes contribute to the extinction of isolated populations following eradication treatments of varying strength (percentage mortality). As a case study, we used historical data on the dynamics of isolated gypsy moth, Lymantria dispar, populations to fit parameters to this model. The parameterized model was then used in simulations that evaluated the efficacy of various eradication strategies. The results indicated that eradication of isolated gypsy moth populations could be easily achieved following a treatment of >80% mortality as long as populations were relatively low (indicated by <100 males captured in pheromone traps).