Larval parasitism of a forest pest, the common pine sawfly Diprion pini (L.) (Hym., Diprionidae), during an endemic density phase

Abstract: Species composition and impact of larval parasitoids during an endemic density phase of a pine forest pest, the common pine sawfly Diprion pini (L.), were investigated by exposure of laboratory-reared larvae of the sawfly to parasitoid attack in natural habitats. Colony-associated host larvae of different instars were exposed continuously from early June until end of September of the years 1994 and 1995 in two pine stands with a different history of pine sawfly outbreaks. Host larvae were exchanged after a defined time interval and those collected up were examined in the laboratory for parasitism. Parasitoid assemblages were shown to be very species poor at both pine stands. The dominant species were the ichneumonid Olesicampe macellator and the tachinid Drino gilva , which reached attack rates on the collected host larvae of 60 and 94%, respectively, during peak activity. The patterns of temporal attack, of multiparasitism, and of superparasitism of these dominant species were studied. In an additional experiment, the response of these species to different host-patch sizes (number of host larvae per colony) were investigated. No aggregative response to higher host-patch densities could be detected. The role and function of these larval parasitoids in the population dynamics of D. pini are discussed.     

Sex pheromone for monitoring flight periods and population densities of the pine sawfly, Diprion jingyuanensis Xiao et Zhang (Hym., Diprionidae)

Abstract:  The pine sawfly, Diprion jingyuanensis Xiao et Zhang, is a serious pest of Chinese pine ( Pinus tabulaeformis Carr.) in the northern parts of China. The general biology of this recently described sawfly species is little known and in the present study we used a synthetic pheromone to monitor its flight period and to evaluate the possibility to use pheromone traps as a tool for estimating and predicting population densities. The attractant pheromone of D. jingyuanensis is (1 S ,2 R ,6 R )-1,2,6-trimethyldodecyl propionate and in this study we used a four-isomer threo-mixture containing the active attractant. Both doses tested, 1 and 2 mg/trap, revealed the same seasonal flight pattern, the higher dosage demonstrating more clear flight peaks. The first flight peak of D. jingyuanensis occurred in mid-June during all 3 years, 1997–1999, and in 1997 and 1998 a second flight peak was also recorded in mid-July. The flight period was similar in time for populations located at higher (1400 m) and lower altitude (1100 m), from early June to late July or early August. Temperature was the main weather factor-affecting trap catches. Diurnal flight activity began at 9.00, peaked at 13.00 and lasted until 20.00. A series of tests with traps in different positions within stands and trees were conducted and the results demonstrated the importance of standardizing the trapping protocol in population monitoring studies. By using traps with 2 mg baits it is possible to detect sawfly occurrence at very low population densities, not detectable by other means. Strong positive correlations between trap catch and relative population density were found in 2000 and 2001, but not in 1998, when only few sites were monitored and the population was in a decreasing phase. The results are promising for future large-scale use of pheromone-based monitoring of D. jingyuanensis .     

Habitat heterogeneity affects predation of European pine sawfly cocoons

Habitat heterogeneity is thought to affect top‐down control of herbivorous insects and contribute to population stability by providing a more attractive microhabitat for natural enemies, potentially leading to reduced population fluctuations. Identifying the parameters that contribute to habitat heterogeneity promoting top‐down control of herbivorous insects by natural enemies could facilitate appropriate management decisions, resulting in a decreased risk of pest insect outbreaks because of a higher level of predation. In our study, we measured the top‐down pressure exerted by small mammals on the cocoons of a notorious pest insect in pine forests, the European pine sawfly (Neodiprion sertifer), which is known to be regulated by small mammal predation. The forest stands used differed in heterogeneity measured in terms of differences in tree diversity and density, understory vegetation height, presence/absence, and density of dead wood. We found higher predation in more dense spots within forest stands. Further, the effect of dead wood on sawfly cocoon predation depended on the pine proportion in forest stands. The addition of dead wood in a manipulation experiment had a slight positive effect on cocoon predation, while dead wood removal caused a clear decrease in predation rate, and the decrease was more pronounced when the proportion of pine increased. Our results show that habitat heterogeneity affects predation by generalist predators on herbivorous insects. This knowledge could be applied to reduce the risk of insect outbreaks by applying management methods that increase heterogeneity in perennial systems such as forests and orchards, thus decreasing the levels of insect damage.     

Cocoon predation on diprionid sawflies: the effect of forest fertility

Predation by small mammals is thought to be one of the main regulators of outbreaking sawfly species. It has been suggested that predation may be lower in poor and dryish forests, and this is the reason why outbreaks often begin from this type of environment. We studied experimentally how fertility of the forest site affects cocoon predation experienced by two sawfly species, the common pine sawfly Diprion pini (Linnaeus) and the European pine sawfly Neodiprion sertifer (Geoffroy). We applied a fertilization treatment to selected pine-dominated barren forest sites in Finland, and 2–4 years later monitored predation on the sawfly cocoons in fertilized and control areas. The results did not support the idea that forest fertility was related to cocoon predation. We also could not verify that small mammal abundance was related to fertility of the forest. The most obvious pattern we observed was that the two sawfly species differed dramatically in predation experienced. N. sertifer has its cocoon phase in mid-summer and experienced only moderate predation (37%) whereas D. pini, with its cocoon phase in autumn, suffered from very heavy predation (96%). Our observations suggest that if predation is important in controlling the population dynamics of the species, its impact depends more on the sawfly species and season than on the fertility of the forest site. Received: 1 March 1998 / Accepted: 25 May 1998     

Regulation of forest defoliating insects through small mammal predation: reconsidering the mechanisms

Population densities of forest defoliating insects may be regulated by small mammal predation on the pupae. When outbreaks do occur, they often coincide with warm, dry weather and at barren forest sites. A proposed reason for this is that weather and habitat affect small mammal population density (numerical response) and hence pupal predation. We propose an alternative explanation: weather and habitat affect small mammal feeding behaviour (functional response) and hence the outbreak risks of forest pest insects. We report results from laboratory and field-enclosure experiments estimating rates of pupal predation by bank voles (Myodes glareolus) on an outbreak insect, the European pine sawfly (Neodiprion sertifer), at different temperatures (15 and 20 °C), in different microhabitats (sheltered and non-sheltered), and with or without access to alternative food (sunflower seeds). We found that the probability of a single pupa being eaten at 20 °C was lower than at 15 °C (0.49 and 0.72, respectively). Pupal predation was higher in the sheltered microhabitat than in the open one, and the behaviour of the voles differed between microhabitats. More pupae were eaten in situ in the sheltered microhabitat whereas in the open area more pupae were removed and eaten elsewhere. Access to alternative food did not affect pupal predation. The results suggest that predation rates on pine sawfly pupae by voles are influenced by temperature- and habitat-induced variation in the physiology and behaviour of the predator, and not necessarily solely through effects on predator densities as previously proposed.     

Acidic precipitation increases egg survival in Neodiprion sertifer

The effects of simulated acidic precipitation on the egg viability of the European pine sawfly, Neodiprion sertifer Geoffroy (Hymenoptera: Diprionidae) were studied by spraying egg clusters with a mixture of sulphuric and nitric acid (1:1, pH 2, 3, 4, 5, 6, distilled water, natural precipitation) for a month in April–May. Egg viability analyses showed that the proportion of hatched first instar larvae increased with increasing acidity. The better survival of sawfly eggs in pine needles in more acidic conditions may be partly due to changes in needle physiology affecting egg mortality. However, when the eggs were reared without needles in Petri-dishes, the proportion of successfully hatched larvae likewise increased with increasing acidity indicating direct acid-induced changes in the eggs themselves. Acidic precipitation as a stress factor of the host plant apparently changes population dynamics of pine sawflies and may be an important factor in triggering outbreaks.     

Geographic variation in winter freezing susceptibility in the eggs of the European pine sawfly (Neodiprion sertifer)

Abstract 1 The European pine sawfly, Neodiprion sertifer (Geoffroy) (Hymenoptera, Diprionidae), frequently defoliates Scots pine (Pinus sylvestris L.) forests in northern Europe. It overwinters as an egg. It has been proposed that the high egg mortality caused by low winter temperatures limits the occurrence of outbreaks to the southern part of Fennoscandia. 2 In this study, variation in freezing avoidance by egg supercooling between four Finnish populations (originating between latitudes 60°N and 69°N) of N. sertifer was tested by differential thermal analysis. Offspring of 20 females within each population were selected for the study. The freezing avoidance of parasitized eggs was also examined. 3 The northernmost Inari population was found to be the cold hardiest, and the southernmost (Hanko) was the least hardy population. The within‐population variation between females was greatest in the population from Inari, and the next greatest in the one from Hanko. The inland populations in Eastern Finland had the smallest within‐population variation in freezing avoidance. 4 The high variation in freezing avoidance of eggs will enable N. sertifer to adapt to the predicted climate change and to spread its distribution northwards. This may also change the risk for outbreaks in this area. Parasitized eggs froze at higher temperature than healthy eggs. This observation indicates that N. sertifer may experience reduced egg parasitism in certain winter climate conditions.     

Genetic diversity within and among populations of Diprion pini (Hym., Diprionidae) determined by random amplified polymorphic DNA-polymerase chain reaction of haploid males

Abstract: Based on haploid males, random amplified polymorphic DNA (RAPD) markers were used to study genetic variation within and among four French populations and one Finnish outgroup population of the common pine sawfly, Diprion pini (L.), representing a severe European forest pest associated with mass outbreaks. Taking into account that all multilocus haplotypes were detected, a total of 140 individuals were completely discriminated by means of 17 polymorphic markers (present or absent), which were amplified from nine selected random-decamer primers. All populations shared the same genetic types, but pronounced population-specific frequency distributions were found, indicating that on average 84% of the present genetic variation exists within populations and the remaining smaller part counts for interpopulational variation. The haplotype differentiation registered was able to distinguish between the class of lowland populations and the class of highland populations. Reproductive isolation and therefore limited altitudinal gene flow, indicated in case of a French highland population by an enlarged number of fixed markers, and hitchhiking effects with respect to selective processes at certain loci following local adaptation and speciation are discussed to explain the population structures found. A possible selection is indicated by five markers, showing significantly different frequency distributions between the class of highland populations and the class of lowland populations.     

The sawfly <Emphasis Type="Italic">Acantholyda flaviceps</Emphasis> (Hymenoptera,Pamphiliidae) in Russia

Some biological features of the sawfly Acantholyda erythrocephala, previously unknown as the mass pest of the Siberian pine, are given. The focus of this species in Tomsk Province has been active during the last 15–20 years. The population dynamics of sawflies in this focus during 12 years is given. The phenology, distribution of eggs by females, age differences between larvae, and characteristics of the forest stand where an outburst of mass breeding was observed, are described.     

北京地区油松潜在重要害虫松阴吉丁的形态和生物学特性观察

【目的】调查松阴吉丁Phaenops yin KubáňBíly(鞘翅目:几丁甲科)对北京地区油松Pinus tabuliformis Carrière的危害情况,了解该害虫的生物学特性;分析其偶然性暴发成灾的原因及规律,探讨该害虫的综合治理措施。【方法】本研究通过林间采集、解剖受害木段以及实验室罩笼饲养等方法,调查了松阴吉丁的发生现状,包括油松的受害情况和受害症状、天敌生物等,并结合历史文献和北京地区近20年的气象资料分析了该虫偶然暴发的原因。【结果】松树死亡率约为30%,受害株率约为60%,发现两种主要寄生性天敌始刻柄茧蜂Atanycolus initiator(Fabricius)和赤腹深沟茧蜂Iphiaulax impostor(Scopoli)。【结论】该虫在我国北方地区偶然性暴发的主要原因可能与持续高温干旱和降雨量减少导致的油松长势衰弱有关。建议结合气象变化及时预测预报松阴吉丁的发生,通过补水降温、清理枯枝、保护天敌、合理用药等综合治理措施调控该害虫的种群动态。     

Spatial Pattern of Attacks of the Invasive Woodwasp Sirex noctilio,at Landscape and Stand Scales

Invasive insect pests are responsible for important damage to native and plantation forests, when population outbreaks occur. Understanding the spatial pattern of attacks by forest pest populations is essential to improve our understanding of insect population dynamics and for predicting attack risk by invasives or planning pest management strategies. The woodwasp Sirex noctilio is an invasive woodwasp that has become probably the most important pest of pine plantations in the Southern Hemisphere. Our aim was to study the spatial dynamics of S. noctilio populations in Southern Argentina. Specifically we describe: (1) the spatial patterns of S. noctilio outbreaks and their relation with environmental factors at a landscape scale; and (2) characterize the spatial pattern of attacked trees at the stand scale. We surveyed the spatial distribution of S. noctilio outbreaks in three pine plantation landscapes, and we assessed potential associations with topographic variables, habitat characteristics, and distance to other outbreaks. We also looked at the spatial distribution of attacked trees in 20 stands with different levels of infestation, and assessed the relationship of attacks with stand composition and management. We found that the spatial pattern of pine stands with S. noctilio outbreaks at the landscape scale is influenced mainly by the host species present, slope aspect, and distance to other outbreaks. At a stand scale, there is strong aggregation of attacked trees in stands with intermediate infestation levels, and the degree of attacks is influenced by host species and plantation management. We conclude that the pattern of S. noctilio damage at different spatial scales is influenced by a combination of both inherent population dynamics and the underlying patterns of environmental factors. Our results have important implications for the understanding and management of invasive insect outbreaks in forest systems.     

The role of prey taxis in biological control: a spatial theoretical model

We study a reaction-diffusion-advection model for the dynamics of populations under biological control. A control agent is assumed to be a predator species that has the ability to perceive the heterogeneity of pest distribution. The advection term represents the predator density movement according to a basic prey taxis assumption: acceleration of predators is proportional to the prey density gradient. The prey population reproduces logistically, and the local population interactions follow the Holling Type II trophic function. On the scale of the population, our spatially explicit approach subdivides the predation process into random movement represented by diffusion, directed movement described by prey taxis, local prey encounters, and consumption modeled by the trophic function. Thus, our model allows studying the effects of large-scale predator spatial activity on population dynamics. We show under which conditions spatial patterns are generated by prey taxis and how this affects the predator ability to maintain the pest population below some economic threshold. In particular, intermediate taxis activity can stabilize predator-pest populations at a very low level of pest density, ensuring successful biological control. However, very intensive prey taxis destroys the stability, leading to chaotic dynamics with pronounced outbreaks of pest density.     

The history and control of the pine beauty moth, Panolis flammea (D. & S.) (Lepidoptera: Noctuidae), in Scotland from 1976 to 2000

1 The pine beauty moth, Panolis flammea, has been a serious pest of lodgepole pine plantations in Scotland since 1976. It historically feeds on native Scots pine throughout Europe but population levels of P. flammea on this host have never been high enough to cause tree mortality in the U.K. 2 This paper reviews recent advances in the biology of the pest and documents control programmes from 1976 to 1999. 3 There has been practically uninterrupted population monitoring of P. flammea from 1977 to the present day in Scottish lodgepole pine plantations. Intervention with chemical spraying has often been necessary. 4 The population data suggest that populations of P. flammea may have had a cyclic pattern over the monitoring period, with outbreaks occurring at regular intervals of between 6 and 7 years. 5 The amplitude of population cycles was large during the 1970s and 1980s, but has dampened in recent years. Natural enemies are believed to contribute to this trend. Fungal disease, specifically, appears to have had a greater effect on pest populations in recent years than in the past and is suggested to have contributed significantly to the population dynamics observed since 1990.     

Influence of plant quality on pine sawfly population dynamics

The contribution of plant quality to the population dynamics of herbivorous insects has been an issue of much controversy. Many studies have documented how variable plant quality differentially influences the survival and fecundity of insect individuals. Whether or not such effects can be translated to the level of insect populations is, however, not clear. In order to test this hypothesis one needs to combine processes at both the level of the individual and the population. This is difficult with an empirical approach, but could be achieved by means of modeling given that appropriate data exist for both levels of organization. In this paper we report on a model developed to analyze whether altered Scots pine (Pinus sylvestris) quality can contribute to the build‐up of populations of the European pine sawfly (Neodiprion sertifer). Experimental data on responses of sawfly larvae to variable plant quality, i.e. needle concentrations of resin acids, were used to parameterize the model. Larval survival and sawfly fecundity are reduced at high resin acid concentrations. However, high resin acid concentrations are, at the same time, beneficial because larval defense against predators is enhanced. In the model, data on individual responses were combined with literature data at the population level; a type III functional response related to cocoon predation was presumed to be the density‐dependent process regulating sawfly populations. The analysis showed that the risk for an outbreak is high when needle resin acid concentration (r) or larval predation pressure (p) is low. When r or p is high there is no risk. By analyzing different scenarios it was found that small changes in r and p can result in the sawfly population moving from low to high outbreak risk. Changes of the same, or larger, magnitude in r have been observed in empirical studies. The role of tritrophic interactions was also considered. This was done by removing the positive effects of resin acids on larval performance in the model. It was found that the anti‐predator defense of N.sertifer makes it prone to outbreak under wider combinations of r and p than an insect without the defense. We conclude that small changes in a density‐independent factor, such as needle chemistry, can have significant effects on herbivore population dynamics because increased fecundity and survival caused by needle quality may allow the population to escape the control of density‐dependent factors, such as cocoon predation.     

我国甜菜夜蛾间歇性暴发的非均衡性循环波动

[摘要] 为进一步了解我国甜菜夜蛾Spodoptera exigua ( Hbner)间歇性暴发成灾的规律,作者应用时间序列分析技术研究了我国甜菜夜蛾间歇性暴发的时间序列波动规律。结果表明,在1949—2008年的60a中,我国甜菜夜蛾大尺度暴发总年频次为120次,各年度暴发次数(年频次)存在着非均衡性循环波动特征,并且在其波动过程中呈明显的上升趋势。按“年频次”强度,可将我国1949—2008年60a中甜菜夜蛾发生过程,划分成两个明显不同的阶段。第一阶段（1949—1984年）为平稳低发阶段,36a总年频次量为4次,年平均仅0.11次;第二阶段（1985—2008年）则为波浪式上升性高发阶段,24a总年频次为116次,年平均4.83次（为第一阶段的43.91倍））。按第二阶段24a（1985—2008）数据进行自相关系数和频谱图分析,结果表明,我国甜菜夜蛾大尺度暴发存在2.8a和11.2a2种不同长度的循环周期,其暴发趋势指数对滞后1a和滞后4a的影响为正相关,而对滞后5a和滞后6a的影响则为负相关。本文根据甜菜夜蛾暴发指数的非均衡周期性特征,建立了以时间序列为自变量的甜菜夜蛾暴发指数非均衡周期性预测模型,经回代结果检验,理论值与实测值之间无显著差异。     

Monitoring the European pine sawfly with pheromone traps in maturing Scots pine stands

Abstract

• 1 During 1989–93, field studies were conducted in Finland to develop a method based on pheromone traps to monitor and forecast population levels of the European pine sawfly (Neodiprion sertifer Geoffr.) and tree defoliation.
• 2 Three traps per site were baited with 100 µg of the N. sertifer sex pheromone, the acetate ester of (2S,3S,7S)‐3,7‐dimethyl‐2‐pentadecanol (diprionol), in maturing pine stands in southern and central Finland. In addition, three different dosages (1, 10 and 100 µg) of the pheromone were tested in 1991–92.
• 3 The highest number of males was observed in traps baited with the highest dose. On average, there was a 10‐fold increase in trap catch between lure doses.
• 4 Density of overwintering eggs was used to evaluate the effectiveness of pheromone traps in predicting sawfly populations. The proportion of healthy overwintering eggs was determined each year. A model based on the number of current shoots on sample trees, diameter at breast height and tree height was formulated to estimate eggs per hectare.
• 5 Linear regression analysis produced high coefficients of determination between number of males in traps and density of total eggs in the subsequent generation, when populations were at peak densities. The relationships were not significant for low population densities. The results indicate a risk of moderate defoliation when the seasonal trap catch is 800–1000 males per trap or higher.

     

Larch- and pine-feeding host races of the larch bud moth (Zeiraphera diniana) have cyclic and synchronous population fluctuations

Population cycles of many forest-defoliating insects often show synchronous fluctuations at both intra-specific (spatial synchrony) and inter-specific levels. However, population dynamics of different host-associated biotypes of the same species, such as those of the larch budmoth (LBM), Zeiraphera diniana (Lepidoptera: Tortricidae), have never been compared. This species causes extensive defoliation of larch forests every 8 to 9 years in the Alps, but it consists of two genetically-differentiated host races, the first one developing on European larch, Larix decidua , and the other one developing on Swiss stone pine, Pinus cembra . The dynamics of Zeiraphera populations have been extensively studied on larch, whereas little is known about larval density and possible population fluctuations on sympatric pines. A larval census on Swiss stone pine was conducted in the Swiss Alps intermittently between 1958 to 2004 and in the French Alps from 1992 to 2004. Population density of Zeiraphera on pine varied up to 5000-fold and showed periodic oscillations, with five peaks in Switzerland and one in France. Because the feeding activity of the pine race is restricted to the elongating shoot of the current year, no conspicuous defoliation of pine trees was noted during years of high larval densities. Zeiraphera populations on pine oscillated in significant synchrony with larch-associated populations, and peak densities were observed either the same year or shifted by±one year. Our results did not allow any explanation for cyclic fluctuations of LBM on pine, but the synchrony with the larch race's cycle suggests that studies on genetics as well as on parasitism should be intensified.     

Top-down pressure by generalist and specialist natural enemies in relation to habitat heterogeneity and resource availability

Habitat heterogeneity might promote the abundance and richness of natural enemies potentially leading to higher top-down pressure on herbivorous insects. Heterogeneous habitats could provide natural enemies with more abundant and alternative resources and a greater variety of micro-habitats. Natural enemies with different searching behaviours, e.g. generalists and specialists, could be affected in different ways by habitat heterogeneity, thus affecting their pressure on herbivorous insects.To understand how top-down pressure on herbivorous insects is promoted by habitat heterogeneity, it is crucial to investigate which parameters contributing to habitat heterogeneity affect not only the abundance and richness but also the searching behaviour of different natural enemies. We investigated the relationship between heterogeneity in forest habitats and the top-down pressure exerted by generalist predators and specialist parasitoids on larvae of the European pine sawfly (Neodiprion sertifer).We used forest stands with endemic or epidemic densities of resident sawfly populations. Within each stand we selected experimental trees to create variation in tree species diversity and density in their surrounding area, i.e. habitat heterogeneity. We found that a higher tree density increased the predation by generalists on sawfly larvae in stands with endemic sawfly densities. Parasitoids were less successful in stands with endemic sawfly densities. Total mortality depended on stand character and the proportion of pine around experimental trees.The explained variation in the response variables by the models is relatively low, indicating that other measures of heterogeneity, like understory vegetation and presence of dead wood could contribute to the observed variation. Also, interference between generalist and specialist enemies could affect the realized mortality pressure. Thus, the effect of tree species diversity in combination with these other measures of heterogeneity needs to be recognized to promote the presence and the activity of natural enemies in managed habitats.     

First use of pheromones to detect phenology patterns and density relationships of pine sawflies in German forests

1 Monitoring studies of pine sawflies with pheromone traps were performed for the first time in Germany. Pheromone traps baited with species‐specific pheromone substances were installed in pine forests at different locations in Bavaria, Brandenburg and Lower Saxony during two years. 2 It was possible to track the flight phenology of Diprion pini, Gilpinia pallida and Neodiprion sertifer reliably and to get information about the number of generations of these species in 1997 and 1998. 3 A clear relationship between trap catch and population density could not be detected, but qualitative changes in trap catch caused by different density levels were observed. 4 For D. pini, trap catches were different among endemic populations of different forest types. Furthermore, catches of males reflected the results from the regular cocoon collections by foresters during the previous winter. 5 For N. sertifer, trap catches in endemic populations were well separated from trap catches on sites with higher sawfly densities. However, no significant correlation between trap catch and sawfly density or defoliation level could be found. These results suggest that the efficacy of the pheromone traps probably varied with biological features (sex ratio, density level, immigration) of the particular population.     

Outbreaks, gene flow and effective population size in the migratory locust, Locusta migratoria: a regional-scale comparative survey

The potential effect of population outbreaks on within and between genetic variation of populations in pest species has rarely been assessed. In this study, we compare patterns of genetic variation in different sets of historically frequently outbreaking and rarely outbreaking populations of an agricultural pest of major importance, the migratory locust, Locusta migratoria . We analyse genetic variation within and between 24 populations at 14 microsatellites in Western Europe, where only ancient and low-intensity outbreaks have been reported (non-outbreaking populations), and in Madagascar and Northern China, where frequent and intense outbreak events have been recorded over the last century (outbreaking populations). Our comparative survey shows that (i) the long-term effective population size is similar in outbreaking and non-outbreaking populations, as evidenced by similar estimates of genetic diversity, and (ii) gene flow is substantially larger among outbreaking populations than among non-outbreaking populations, as evidenced by a fourfold to 30-fold difference in F _ST values. We discuss the implications for population dynamics and the consequences for management strategies of the observed patterns of genetic variation in L. migratoria populations with contrasting historical outbreak frequency and extent.