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BackgroundInvasive species threaten the productivity and stability of natural and managed ecosystems. Predicting the spread of invaders, which can aid in early mitigation efforts, is a major challenge, especially in the face of climate change. While ecological niche models are effective tools to assess habitat suitability for invaders, such models have rarely been created for invasive pest species with rapidly expanding ranges. Here, we leveraged a national monitoring effort from 543 sites over 3 years to assess factors mediating the occurrence and abundance of brown marmorated stink bug (BMSB, Halyomorpha halys), an invasive insect pest that has readily established throughout much of the United States.ResultsWe used maximum entropy models to estimate the suitable habitat of BMSB under several climate scenarios, and generalized boosted models to assess environmental factors that regulated BMSB abundance. Our models captured BMSB distribution and abundance with high accuracy, and predicted a 70% increase in suitable habitat under future climate scenarios. However, environmental factors that mediated the geographical distribution of BMSB were different from those driving abundance. While BMSB occurrence was most affected by winter precipitation and proximity to populated areas, BMSB abundance was influenced most strongly by evapotranspiration and solar photoperiod.ConclusionOur results suggest that linking models of establishment (occurrence) and population dynamics (abundance) offers a more effective way to forecast the spread and impact of BMSB and other invasive species than simply occurrence-based models, allowing for targeted mitigation efforts. Implications of distribution shifts under climate change are discussed. © 2022 The Authors. Pest Management Science published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

Drosophila suzukii, or spotted-wing drosophila, is now an established pest in many parts of the world, causing significant damage to numerous fruit crop industries. Native to East Asia, D. suzukii infestations started in the United States a decade ago, occupying a wide range of climates. To better understand invasion ecology of this pest, knowledge of past migration events, population structure, and genetic diversity is needed. In this study, we sequenced whole genomes of 237 individual flies collected across the continental United States, as well as several sites in Europe, Brazil, and Asia, to identify and analyze hundreds of thousands of genetic markers. We observed strong population structure between Western and Eastern US populations, but no evidence of any population structure between different latitudes within the continental United States, suggesting that there are no broad-scale adaptations occurring in response to differences in winter climates. We detect admixture from Hawaii to the Western United States and from the Eastern United States to Europe, in agreement with previously identified introduction routes inferred from microsatellite analysis. We also detect potential signals of admixture from the Western United States back to Asia, which could have important implications for shipping and quarantine policies for exported agriculture. We anticipate this large genomic dataset will spur future research into the genomic adaptations underlying D. suzukii pest activity and development of novel control methods for this agricultural pest.

The olive fruit fly, Bactrocera oleae, has been a key pest of olives in Europe and North America. We conducted the largest exploration for parasitoids associated with the fly across Sub-Saharan Africa (Kenya, Namibia, and South Africa) including some of the fly's adjoining regions (Canary Islands, Morocco, Réunion Island and Tunisia). From Sub-Saharan regions, four braconids were collected: Bracon celer, Psytallia humilis, P. lounsburyi, and Utetes africanus. Results showed that their regional dominance was related to climate niches, with P. humilis dominant in hot semi-arid areas of Namibia, P. lounsburyi dominant in more tropical areas of Kenya, and U. africanus prevalent in Mediterranean climates of South Africa. Psytallia concolor was found in the Canary Islands, Morocco and Tunisian, and the Afrotropical braconid Diachasmimorpha sp. near fullawayi on Réunion Island. Furthermore, we monitored the seasonal dynamics of the fly and parasitoids in Cape Province of South Africa. Results showed that fruit maturity, seasonal variations in climates and interspecific interactions shape the local parasitoid diversity that contribute to the low fly populations. The results are discussed with regard to ecological adaptations of closely associated parasitoids, and how their adaptations impact biocontrol.

The vine mealybug (VMB), Planococcus ficus (Hemiptera: Pseudococcidae), is a key insect pest of vineyards, and improvements in sustainable control of this pest are needed to meet increasing consumer demand for organically farmed products. One promising option is mating disruption. In a series of experiments conducted from 2004 to 2007, we tested the effects of mating disruption on trap captures of Pl. ficus males in pheromone-baited traps, on Pl. ficus numbers and age structure on vines, and on damage to grape clusters. From 2004 to 2005, the effects of dispenser load (mg active ingredient per dispenser) were also assessed, and dispensers were compared to a flowable formulation. Across all trials, mating disruption consistently reduced pheromone trap captures and often reduced mealybug numbers on vines and/or crop damage, regardless of the pheromone dose that was applied. Reductions in Pl. ficus densities in mating disruption plots were not accompanied by clear effects on mealybug population age structure; however, production of non-viable ovisacs by unmated females may have obscured differences in proportional representation of ovisacs. Pheromone trap captures were never lowered to zero (often called trap shut down), possibly because trials were conducted in vineyards with unusually high Pl. ficus densities. Trap-capture patterns in both treated and control plots commonly began low in April-May, increased in mid-July or August, and often decreased in September-October when post-harvest insecticides were applied. During the four-year trial, the release rate from plastic sachet dispensers was improved by industry cooperators as pheromone was released too quickly (2004) or not completely released during the season (2005-2006). The flowable formulation performed slightly better than dispensers at the same application dose. Results over all years suggest season-long coverage or late-season coverage may be as or more important than dose per hectare. Development of a dispenser with optimized season-long pheromone emission or targeted seasonal periods should be a future goal.

A major challenge to the area-wide management of Drosophila suzukii is understanding the fly's host use and temporal dynamics, which may dictate local movement patterns. We determined D. suzukii's seasonal host use in California's San Joaquin Valley by sampling common crop and non-crop fruits in a temporal sequence of fruit ripening. We then evaluated the suitability of selected fruits as hosts. Drosophila suzukii emerged from both intact and damaged cherries during the cooler, early season period. Fly density remained low through the hot spring-summer period and re-surged as temperatures lowered in fall when the fly did not cause damage to intact peach, nectarine, plum, pear, grape, pomegranate, apple, persimmon and citrus (in order of ripening) but did emerge from the damaged fruits of these crops. The fly also emerged from two ornamental fruits (loquats and cactus) but was not found on wild plum and two endemic wild fruits (buckthorn and bitter berry). Drosophila suzukii completed development (egg to adult) on cactus, mandarin carpel, pomegranate seed, wild plum and buckthorn at survival rates similar to cherry (51.2-68.8%), whereas it had a lower survival rate on bitter cherry (33.2%), table grape (31.5%), raisin grape (26.5%), and wine grape (4.5%). The high acidity levels of grapes negatively affected the fly's fitness. Among 10 cherry cultivars, survival rate was not affected by sugar content, but it decreased with increasing egg density per gram of fruit. Results suggest that in California's San Joaquin Valley, the early season crops are most vulnerable, summer fruits ripen during a period of low pest pressure, and late season fruits, when damaged, serve to sustain D. suzukii's populations in this region.

Legalization of cannabis production has daylighted a unique and highly valuable crop in California agriculture. State and regulatory agencies must now address the ecological, social and agricultural effects of cannabis production, but little is known about how growers produce this crop. Using an online survey, we gathered information from growers in July 2018 on their production practices. According to responses from about 100 growers, most cannabis was produced outdoors or in greenhouses, relied primarily on groundwater and used biologically based inputs for pest management. Many farms employed seasonal workers paid at fixed piece rates. Regulatory compliance varied according to farm size. Beginning to document growing practices will help scientists formulate key environmental, social and agronomic questions and develop relevant research and extension programs to promote best management practices and minimize negative environmental impacts of production.

H. Bodwitch is Postdoctoral Fellow, Department of Natural Resource Sciences, McGill University, Quebec, Canada; J. Carah is Senior Freshwater Ecologist, The Nature Conservancy, San Francisco; K.M. Daane is UC Cooperative Extension Specialist, Department of Environmental Science, Policy, and Management, UC Berkeley; C. Getz is Associate Cooperative Extension Specialist, Department of Environmental Science, Policy, and Management, UC Berkeley; T.E. Grantham is Assistant Cooperative Extension Specialist and Adjunct Professor, Department of Environmental Science, Policy, and Management, UC Berkeley; G.M. Hickey is Associate Professor, Department of Natural Resource Sciences, McGill University, Quebec, Canada; H. Wilson is Assistant Cooperative Extension Specialist, Department of Entomology, UC Riverside.

Studies of spatiotemporal dynamics are central to efforts to characterize the epidemiology of infectious disease, such as mechanism of pathogen spread and pathogen or vector sources in the landscape, and are critical to the development of effective disease management programs. To that end, we conducted a multi-year study of 20 vineyard blocks in coastal northern California to relate the dynamics of a mealybug vector, Pseudococcus maritimus (Ehrhorn) (Hemiptera: Pseudococcidae), to incidence of grapevine leafroll disease (GLD). In each vineyard block, a subset of vines were scored visually for relative mealybug abundance, disease was quantified by visual assessment, and virus presence was verified using standard laboratory molecular assays. GLD incidence was analyzed with a classification and regression tree, and with a hierarchical model that also captured variability among blocks and heterogeneity within blocks. Both analyses found strong interannual variability in incidence, with the hierarchical model also capturing substantial between- and within-block heterogeneity, but with significant contributions of vector abundance and pathogen supply (prior disease incidence) to the frequency of newly diseased vines. These results strengthen further the conclusion that mealybug vectors are causally related to pathogen spread in this system and are therefore an important target for management. Moreover, they are consistent with relatively efficient secondary spread of the pathogen, suggesting an important role for the removal of diseased vines as a tool to mitigate further damage.

Determining the most likely source of an invasive pest species might help to improve their management by establishing efficient quarantine measures and heading the search of efficient biological control agents. Planococcus ficus is an invasive mealybug pest of vineyards in Argentina, California, Mexico, Peru and South Africa. This mealybug pest had a previously known geographic distribution spanning southern Europe, the Middle East, and parts of northern Africa. In North America, Pl. ficus was first discovered in the early 1990s and soon thereafter in Mexico. To determine the origin of invasive populations in North America, Pl. ficus from California and Mexico were compared with material throughout its presumptive native range in the Mediterranean region, as well as material collected from an older invasion in South Africa and recently invaded Argentina. From each sample location, genomic DNA was sequenced for the nuclear internal transcribed spacer one (ITS1) and the mitochondrial cytochrome c. oxidase one (CO1). Phylogenetic analyses of CO1, ITS1 and concatenated CO1 and ITS1 data-sets using Bayesian and neighbor-joining analysis support two major divisions: a European grouping (Europe, Tunisia, Turkey) and a Middle Eastern grouping (Israel and Egypt). The invasive populations in Argentina and South Africa align with the European group and the invasive populations in North America align with the Middle Eastern group, with one Israel sample aligning closely with the North American clade, suggesting that Israel was the origin of those populations.

In recent years, there has been an increase in the adventive establishment and spread of parasitoid wasps outside of their native range. However, lack of taxonomic tools can hinder the efficient screening of field-collected samples to document the establishment and range expansion of parasitoids on continent-wide geographic scales. Here we report that Leptopilina japonica (Hymenoptera, Figitidae), a parasitoid of the globally invasive fruit pest Drosophila suzukii (Diptera, Drosophilidae), is now widespread in much of North America despite not having been intentionally introduced. Surveys in 2022 using a variety of methods detected L. japonica in 10 of 11 surveyed USA States and one Canadian Province where it was not previously known to occur. In most surveys, L. japonica was the most common species of D. suzukii parasitoid found. The surveys also resulted in the detection of Ganaspis cf. brasiliensis (Hymenoptera, Figitidae), the recently-released biological control agent of D. suzukii, in six USA States where it had not previously been found. These new detections are likely a result of intentional biological control introductions rather than spread of adventive populations. A species-specific multiplex PCR assay was developed as a rapid, accurate and cost-effective method to distinguish L. japonica, G. cf. brasiliensis, the closely-related cosmopolitan parasitoid Leptopilina heterotoma (Hymenoptera, Figitidae) and other native parasitoid species. This dataset and the associated molecular tools will facilitate future studies of the spread and ecological impacts of these introduced parasitoids on multiple continents. [ABSTRACT FROM AUTHOR]

The diet of adult parasitoid wasps is vital for their survival and reproduction. However, the availability of food resources, such as plant nectar, can vary widely in cropping systems, potentially affecting parasitoid fitness and thereby biological control of pests. The egg parasitoid Hadronotus pennsylvanicus (Ashmead) (Hymenoptera: Scelionidae) is a potential biological control agent of the pistachio pest Leptoglossus zonatus (Dallas) (Heteroptera: Coreidae). While H. pennsylvanicus is known to attack L. zonatus eggs in California, USA, parasitism rates in orchards are highly variable. Floral resource provisioning has the potential to enhance parasitoid longevity and thus improve parasitism rates, leading to reduced pest densities. Here, a combination of field and laboratory studies was used to assess the influence of flowering groundcovers on the reproductive fitness of H. pennsylvanicus and the abundance of L. zonatus. Evaluated groundcovers included oat (Avena sativa L.), cowpea (Vigna unguiculata L.), white mustard (Sinapis alba L.), and buckwheat (Fagopyrum esculentum Moench). Under laboratory conditions, buckwheat and mustard provided the greatest benefit to female H. pennsylvanicus longevity. However, females provided a buckwheat diet produced the greatest number of offspring over the course of their lifetime. In field trials, flowering groundcovers did not influence the abundance of H. pennsylvanicus nor parasitism rates on L. zonatus. While the availability of floral resources can improve the reproductive fitness of H. pennsylvanicus, the use of groundcovers in pistachio did not enhance biological control of L. zonatus. [ABSTRACT FROM AUTHOR]

Grapevine leafroll-associated virus 3 (GLRaV-3) is associated with the economically damaging grapevine leafroll disease, and is transmitted in a semi-persistent manner by several mealybug species. We performed the first controlled field study of vector-mediated inoculations with GLRaV-3 in a commercial vineyard with previously asymptomatic vines, and monitored the vines during four growing seasons. We then compared the outcome of vector-mediated inoculations in the field study to an analogous laboratory study. In the vineyard, about half of all inoculated plants became infected with GLRaV-3, fewer than in the controlled laboratory inoculations. Mealybugs had lower settling and feeding success in the field than in the laboratory inoculations. Our study suggests that laboratory studies may overestimate transmission efficiency. All successfully inoculated vineyard plants first became symptomatic one growing season after inoculations, and berry quality declined within one year after inoculations. Vineyard plants were effective virus sources one year after inoculations. Our findings generally indicate that newly symptomatic vines in commercial vineyards probably became infected during the previous growing season, and a decline in berry quality can be expected during the same year in which symptoms appear.

Drosophila suzukii causes economic damage to berry and stone fruit worldwide. Laboratory-generated datasets were standardized and combined on the basis of degree days (DD), using Gompertz and Cauchy curves for survival and reproduction. Eggs transitioned to larvae at 20.3 DD; larvae to pupae at 118.1 DD; and pupae to adults at 200 DD. All adults are expected to have died at 610 DD. Oviposition initiates at 210 DD and gradually increases to a maximum of 15 eggs per DD at 410 DD and subsequently decreases to zero at 610 DD. These data were used as the basis for a DD cohort-level population model. Laboratory survival under extreme temperatures when DD did not accumulate was described by a Gompertz curve based on calendar days. We determined that the initiation of the reproductive period of late dormant field-collected female D. suzukii ranged from 50 to 800 DD from January 1. This suggests that D. suzukii females can reproduce early in the season and are probably limited by availability of early host plants. Finally, we used the DD population model to examine hypothetical stage-specific mortality effects of IPM practices from insecticides and parasitoids at the field level. We found that adulticides applied during the early season will result in the largest comparative population decrease. It is clear from model outputs that parasitism levels comparable to those found in field studies may have a limited effect on population growth. Novel parasitoid guilds could therefore be improved and would be valuable for IPM of D. suzukii.

Some diseases manifest as one characteristic set of symptoms to the host, but can be caused by multiple pathogens. Control treatments based on plant symptoms can make it difficult to effectively manage such diseases, as the biology of the underlying pathogens can vary. Grapevine leafroll disease affects grapes worldwide, and is associated with several viral species in the family Closteroviridae. Whereas some of the viruses associated with this disease are transmitted by insect vectors, others are only graft-transmissible. In three regions of California, we surveyed vineyards containing diseased vines and screened symptomatic plants for all known viral species associated with grapevine leafroll disease. Relative incidence of each virus species differed among the three regions regions, particularly in relation to species with known vectors compared with those only known to be graft-transmitted. In one region, the pathogen population was dominated by species not known to have an insect vector. In contrast, populations in the other surveyed regions were dominated by virus species that are vector-transmissible. Our survey did not detect viruses associated with grapevine leafroll disease at some sites with characteristic disease symptoms. This could be explained either by undescribed genetic diversity among these viruses that prevented detection with available molecular tools at the time the survey was performed, or a misidentification of visual symptoms that may have had other underlying causes. Based on the differences in relative prevalence of each virus species among regions and among vineyards within regions, we expect that region and site-specific management strategies are needed for effective disease control.

This study evaluated how the proportional area of natural habitat surrounding a vineyard (i.e. landscape diversity) worked in conjunction with crop vigor, cultivar and rootstock selection to influence biological control of the western grape leafhopper (Erythroneura elegantula Osborn). The key natural enemies of E. elegantula are Anagrus erythroneurae S. Trjapitzin & Chiappini and A. daanei Triapitsyn, both of which are likely impacted by changes in landscape diversity due to their reliance on non-crop habitat to successfully overwinter. Additionally, E. elegantula is sensitive to changes in host plant quality which may influence densities on specific cultivars, rootstocks and/or vines with increased vigor. From 2010-2013, data were collected on natural enemy and leafhopper densities, pest parasitism rates and vine vigor from multiple vineyards that represented a continuum of landscape diversity. Early in the season, vineyards in more diverse landscapes had higher Anagrus spp. densities and lower E. elegantula densities, which led to increased parasitism of E. elegantula. Although late season densities of E. elegantula tended to be lower in vineyards with higher early season parasitism rates and lower total petiole nitrogen content, they were also affected by rootstock and cultivar. While diverse landscapes can support higher natural enemy populations, which can lead to increased biological control, leafhopper densities also appear to be mediated by cultivar, rootstock and vine vigor.

Temperature-dependent fecundity and survival data was integrated into a matrix population model to describe relative Drosophila suzukii Matsumura (Diptera: Drosophilidae) population increase and age structure based on environmental conditions. This novel modification of the classic Leslie matrix population model is presented as a way to examine how insect populations interact with the environment, and has application as a predictor of population density. For D. suzukii, we examined model implications for pest pressure on crops. As case studies, we examined model predictions in three small fruit production regions in the United States (US) and one in Italy. These production regions have distinctly different climates. In general, patterns of adult D. suzukii trap activity broadly mimicked seasonal population levels predicted by the model using only temperature data. Age structure of estimated populations suggest that trap and fruit infestation data are of limited value and are insufficient for model validation. Thus, we suggest alternative experiments for validation. The model is advantageous in that it provides stage-specific population estimation, which can potentially guide management strategies and provide unique opportunities to simulate stage-specific management effects such as insecticide applications or the effect of biological control on a specific life-stage. The two factors that drive initiation of the model are suitable temperatures (biofix) and availability of a suitable host medium (fruit). Although there are many factors affecting population dynamics of D. suzukii in the field, temperature-dependent survival and reproduction are believed to be the main drivers for D. suzukii populations.

Grapevine leafroll disease (GLD) is caused by a complex of vector-borne virus species in the family Closteroviridae. GLD is present in all grape-growing regions of the world, primarily affecting wine grape varieties. The disease has emerged in the last two decades as one of the major factors affecting grape fruit quality, leading to research efforts aimed at reducing its economic impact. Most research has focused on the pathogens themselves, such as improved detection protocols, with limited work directed toward disease ecology and the development of management practices. Here we discuss the ecology and management of GLD, focusing primarily on Grapevine leafroll-associated virus 3, the most important virus species within the complex. We contextualize research done on this system within an ecological framework that forms the backbone of the discussion regarding current and potential GLD management strategies. To reach this goal, we introduce various aspects of GLD biology and ecology, followed by disease management case studies from four different countries and continents (South Africa, New Zealand, California-USA, and France). We review ongoing regional efforts that serve as models for improved strategies to control this economically important and worldwide disease, highlighting scientific gaps that must be filled for the development of knowledge-based sustainable GLD management practices.

Grapevine leafroll disease (GLD) is caused by a complex of several virus species (grapevine leafroll-associated viruses, GLRaV) in the family Closteroviridae. Because of its increasing importance, it is critical to determine which species of GLRaV is predominant in each region where this disease is occurring. A structured sampling design, utilizing a combination of RT-PCR based testing and sequencing methods, was used to survey GLRaVs in Napa Valley (California, USA) vineyards (n = 36). Of the 216 samples tested for GLRaV-1, -2, -3, -4, -5, and -9, 62% (n = 134) were GLRaV positive. Of the positives, 81% (n = 109) were single infections with GLRaV-3, followed by GLRaV-2 (4%, n = 5), while the remaining samples (15%, n = 20) were mixed infections of GLRaV-3 with GLRaV-1, 2, 4, or 9. Additionally, 468 samples were tested for genetic variants of GLRaV-3, and of the 65% (n = 306) of samples positive for GLRaV-3, 22% were infected with multiple GLRaV-3 variants. Phylogenetic analysis utilizing sequence data from the single infection GLRaV-3 samples produced seven well-supported GLRaV-3 variants, of which three represented 71% of all GLRaV-3 positive samples in Napa Valley. Furthermore, two novel variants, which grouped with a divergent isolate from New Zealand (NZ-1), were identified, and these variants comprised 6% of all positive GLRaV-3 samples. Spatial analyses showed that GLRaV-3a, 3b, and 3c were not homogeneously distributed across Napa Valley. Overall, 86% of all blocks (n = 31) were positive for GLRaVs and 90% of positive blocks (n = 28) had two or more GLRaV-3 variants, suggesting complex disease dynamics that might include multiple insect-mediated introduction events.

Olive fruit fly commonly infests olives in California’s Central Valley. Field studies indicate that trap counts for olive fruit fly adults in pesticide-free sites decrease in mid- and late summer and then rebound from September to November. Part of this decline is associated with heat stress that the flies experience in mid-July and August. Studies have shown that adult flies will die within a few days if they cannot access adequate amounts of water and carbohydrates. Flight ability is dramatically reduced when resources are unavailable. Olive fruit fly adults may use black scale honeydew as a carbohydrate source to help them survive hot periods. Heat also affects the fly’s reproduction and immature stages within olive fruit. Geographic information system (GIS) maps may be useful for predicting the risk of olive fruit fly infestation.

The widespread and rapid establishment of the olive fruit fly in California required immediate changes in integrated pest management (IPM) programs for olives. After finding that resident natural enemies did not provide adequate control, researchers began a worldwide search for parasitoids, with exploration in the Republic of South Africa, Namibia, India, China and other countries. Parasitoids were shipped to California, and most were studied in quarantine to determine the best species for release. Two parasitoid species — Psyttalia lounsburyi and Psyttalia humilis — are now being released throughout the state’s olive-growing regions, and researchers are studying their effectiveness.

Invasions have often been linked to reduced biodiversity, but the role of non-native species in the decline of native species is ambiguous. In a 2003 survey of four California vineyard regions, exotic spiders (Cheiracanthium spp.) were more dominant in vineyards with lower spider species diversity and reduced spider abundance. There was no evidence for the role of species interactions in the invasion of Cheiracanthium spiders, however, as native spiders from the same feeding guild were most abundant in regions with high Cheiracanthium levels. Comparison with a survey conducted 10 years earlier indicated that the recent invader C. mildei simply represented an addition to the spider community, with no apparent change in proportions of the congener C. inclusum. Invasion success is discussed with respect to agricultural habitat, as results suggest that disturbed conditions in many vineyards may favor Cheiracanthium spp. and native wandering spiders while decreasing overall spider diversity.

Mealybugs have become increasingly important vineyard pests — a result of their direct damage to the vine, their role in transmitting grapevine leafroll viruses, and the costs for their control. Numerous mealybug species are found in vineyards, and each has different biological traits that affect sustainable control options. We review the mealybug pests and their natural enemies to provide some clarification about current trends in biological control tactics and needed directions for future work.

Liquid ant baits are an alternative to broad-spectrum insecticide sprays conventionally used to control Argentine ants. We review the development of liquid ant baits, which capitalize on the ants’ sugar-feeding requirements and social structure to deliver small doses of toxicant throughout the colony. The ant bait program described here, developed for commercial vineyards, also has the potential to facilitate the use of biological controls for mealybug and scale pests. The implementation of an Argentine ant bait program will enable grape growers to target other pests more selectively with insecticides, further contributing to their sustainable viticulture practices.

Our studies illustrate the dynamic nature of both the availability of split nuts and the prevalence of NOW after harvest, as well as illustrate the low density of moths needed to begin the next generation in the spring. Additionally, our work provides some insight into the relative importance of ground mummies compared to tree mummies. Although mortality was greater in ground mummies, they were more important because of their number.