Linking spatiotemporal and demographic patterns in modulating local population outbreak of invasive forest insect

Invasive forest insects can induce tree mortality in two ways: (a) by directly harming trees; or (b) by influencing forest owners to pre‐emptively harvest threatened trees. This study investigates forest owners’ intentions to harvest trees threatened by invasive insects.Our first objective is to identify and characterize agent functional types (AFTs) of family forest owners in the northeastern United States using a set of contingent behaviour questions contained in a mail survey. We establish AFTs as a form of dimension reduction, effectively casting landowners into a typology in which each type (AFT) has distinct probabilities of tree harvesting in response to forest insects. Our analysis identifies three functional types of landowners: ‘Cutters’ (46% of respondents; high intent to harvest trees impacted by invasive forest insects), ‘Responsive Cutters’ (42% of respondents; intent sensitive to insect impact severity), and ‘Non‐cutters’ (12% of respondents; low intent to cut).Our second objective is to model AFT membership to predict the distribution of AFTs across the landscape. Predictors are chosen from a set of survey, geographic and demographic features. Our best AFT‐prediction model has three predictor variables: parcel size (hectares of forest), geographical region, and town‐level forested fraction. Application of the model provides a high‐resolution probability distribution of AFTs across the landscape.By coupling human and insect behaviour, our results allow for holistic assessments of how invasive forest insects disturb forests, inclusive of the management response to these pests.A freePlain Language Summarycan be found within the Supporting Information of this article.

We investigated the oviposition preference and larval performance of Helicoverpa armigera under laboratory conditions to determine if the oviposition preference of individual females on maize, cowpea and cotton correlates with offspring performance on the leaves of the same host plants. The host‐plant preference hierarchy of females did not correlate with their offspring performance. Female moths chose host plants that contributed less to their offspring fitness. Plant effects accounted for the largest amount of variation in offspring performance, while the effects of female (family) was low. The offspring of most females (80%, n = 10) were broadly similar, but 20% (two out of 10), showed marked difference in their offspring performance across the host‐plant species. Similarly, there was no relation between larval feeding preference and performance. However, like most laboratory experiments, our experi‐mental design does not allow the evaluation of ecological factors (for example, natural enemies, host abundance, etc.) that can play an important role in larval performance in the field. Overall, the results highlight the importance of carrying out preference performance analysis on the individual or family level, rather than pooling individuals to obtain average population data.

Higher temperatures projected under current climate change models are generally predicted to exert an overall positive effect on the success of invasive insects through increased survivability, developmental rates and fecundity, and by facilitating geographic range expansion. However, these effects have primarily focused on the shifts in winter temperatures with limited attention to the role that summer heat may play in shaping species ranges or fitness. We examined the thermal ecology of an ecologically important invasive forest insect, the hemlock woolly adelgid (Adelges tsugae), by determining survival during its summer dormancy phase under increasing temperature regimens. From laboratory and field experiments, we documented a positive association between increased temperatures and duration of exposure, and A. tsugae mortality. Adelges tsugae mortality was minimal ( 30 °C). At the warmest, southernmost edge of their range, field mortality of A. tsugae ranged from 8.5 to 81.9% and was strongly correlated with site temperature regimens. Further, we found no significant differences in A. tsugae survival between populations collected from Maine and Georgia, and over a 3-year period within Georgia, indicating that A. tsugae may not be acclimating to heat. These results highlight the importance of including summer temperatures in studies regarding increased temperatures on insect dynamics, and may alter historical predictions of climate change impacts on invasive insects and the conservation of forest ecosystems.

Oviposition preferences of a herbivore, the wheat midge Sitodiplosis mosellana (Gehin) (Diptera: Cecidomyiidae), on wheat were investigated in relation to two hypotheses: female preferences are adapted to offspring performance; plants may evolve independent defenses that both deter oviposition and reduce offspring performance. Variation in egg density and larval performance were measured for three plant genotypes: a preferred, susceptible wheat; a less preferred, susceptible wheat; a less preferred wheat defended against larval feeding. Oviposition peaked 10–11 d after emergence of the inflorescences and then declined sharply on all three wheat genotypes, although the inflorescences of the genotypes developed at different rates. On the preferred, susceptible wheat, larval performance was high for oviposition that occurred until pollination and low later. On a less preferred wheat, larval performance was high when eggs were laid before or after pollination. On a defended wheat, larval performance was always low. Oviposition preference was associated with larval performance that varied with plant developmental stage, but imperfectly, possibly because females do not detect cues for seed development. Females deposited eggs further from larval feeding sites when ovipositing on less preferred wheats, regardless of whether larval performance on the wheat was high or low. A low preference in combination with a shift in oviposition site supports the hypothesis that some wheats have evolved a defense that deters oviposition. This defense against oviposition is independent of a defense that reduces larval performance, which causes an apparent failure in the expected preference-performance relationship.

The high diversity of phytophagous insects has been explained by the tendency of the group towards specialization; however, generalism may be advantageous in some environments. The cerambycid Apagomerella versicolor exhibits intraspecific geographical variation in host use. In northern Argentina it is highly specialized on the herb Pluchea sagittalis (Asteraceae), while in central and southern areas it uses seven Asteraceae species. To study host species geographical variation from ecological and evolutionary perspectives, we investigated field host availability and use across a wide latitudinal range, and performed laboratory studies on insect oviposition preference and larval performance and mitochondrial DNA (mtDNA) variation in a phylogeographical framework. Geographic variation in host use was unrelated to host availability but was highly associated with laboratory oviposition preference, larval performance, and mtDNA variation. Genetic studies revealed three geographic races of A. versicolor with gene flow restriction and recent geographic expansion. Trophic generalism and oligophagy within A. versicolor seem to have evolved as adaptations to seasonal and spatial unavailability of the preferred host P. sagittalis in cooler areas of the species' geographic range. No single genotype is successful in all environments; specialization may be advantageous in environments with uniform temporal and spatial host availability, while being a trophic generalist may provide an adaptive advantage in host-constrained environments.

The preference–performance hypothesis predicts that when female herbivorous insects determine where to position offspring of low mobility, they will select sites that maximize development and survival of those offspring. How this critical relationship responds to variation in climatic and habitat conditions remains untested, however, despite its important consequences for population and evolutionary dynamics. Here we report on 13 years of data totaling 1348 egg clusters of the montane Gillette's checkerspot butterfly Euphydryas gillettii (Lepidoptera: Nymphalidae). We used these data to test the hypothesis that, in environments with climatic and habitat heterogeneity, the oviposition behavior–offspring performance relationship should vary in both space and time. Orientation of egg clusters for maximum morning sun exposure is known to affect developmental rate. We therefore predicted female preference for morning sun orientation to be variable and a function of climatic and habitat conditions. We found that preference for egg cluster orientation on the leaf tracked the phenology of the start of the female flight season but that seasonal temperatures drove most of the variation in egg cluster development time. The relationship between behavior and performance was also dependent upon the climatic effects on survival; sun‐oriented egg clusters had higher survivorship in the coldest year of the four years for which measurements were made. We also examined how conifer cover affected larval survival and female oviposition behavior in one year. Females selected oviposition sites in more open habitat. However, when egg clusters were oriented to intercept morning sun, conifer cover increased survivorship to diapause. Finally, we found that predator activity was lower for morning sun‐oriented egg clusters suggesting that predation patterns may further influence habitat selection for oviposition. This study exemplifies how the relationship between oviposition behavior and offspring performance is context‐dependent: habitat and climate interact to determine preference–performance outcomes.

Herbivorous insects use plant metabolites to inform their host plant selection for oviposition. These host-selection behaviours are often consistent with the preference-performance hypothesis; females oviposit on hosts that maximize the performance of their offspring. However, the metabolites used for these oviposition choices and those responsible for differences in offspring performance remain unknown for ecologically relevant interactions. Here, we examined the host-selection behaviours of two sympatric weevils, the Datura (Trichobaris compacta) and tobacco (T.mucorea) weevils in field and glasshouse experiments with transgenic host plants specifically altered in different components of their secondary metabolism. Adult females of both species strongly preferred to feed on D.wrightii rather than on N.attenuata leaves, but T.mucorea preferred to oviposit on N.attenuata, while T.compacta oviposited only on D.wrightii. These oviposition behaviours increased offspring performance: T.compacta larvae only survived in D.wrightii stems and T.mucorea larvae survived better in N.attenuata than in D.wrightii stems. Choice assays with nicotine-free, JA-impaired, and sesquiterpene-over-produced isogenic N.attenuata plants revealed that although half of the T.compacta larvae survived in nicotine-free N.attenuata lines, nicotine did not influence the oviposition behaviours of both the nicotine-adapted and nicotine-sensitive species. JA-induced sesquiterpene volatiles are key compounds influencing T.mucorea females' oviposition choices, but these sesquiterpenes had no effect on larval performance. We conclude that adult females are able to choose the best host plant for their offspring and use chemicals different from those that influence larval performance to inform their oviposition decisions.

Abstract1 Outbreaks of herbivorous insects tend to be spatially restricted, possibly because of demographic differences between inside and outside the outbreak area. In some cases, the margin of the outbreak area is distinct, allowing comparisons of adjacent areas that may identify factors leading to such differences in abundance. The northern pine processionary mothThaumetopoea pinivorapresently occurs at outbreak densities within a well‐defined area of approximately 3000 ha on the island of Gotland, south Sweden. We investigated whether cohorts of young larvae (first and second instar) had higher growth rate and survival inside or outside the outbreak area.2 Group‐feeding appears to promote outbreaks in certain insect groups. BecauseT. pinivoralarvae are highly social, we also compared larval performance between groups of different sizes inside and outside of the outbreak area: ‘small’ (33 eggs/group) and ‘normal’ (100 eggs/group).3 Averaged over group size, whole colony mortality through the first two instars was two‐fold higher in the non‐outbreak area compared with the outbreak area. Mortality of individual larvae in the surviving colonies, however, did not differ between the two areas. There were only small differences in food quality (toughness, nitrogen content) between the areas, with no detectable effects on larval performance.4 Larval relative growth rate did not differ between reduced and normal‐sized groups, which is surprising given that growth rate is known to increase with group size in other group‐feeding lepidopterans.5 Reduced group size negatively affected larval survival, particularly in the outbreak area; by contrast, normal‐sized groups survived equally well in the two areas. Wood ants (Formicaspp.) were more common outside the outbreak area, and appeared to be the main cause of colony mortality at low larval density. A different result was observed with regard to per‐capita mortality, which was higher in the outbreak area. We speculate that this could have been due to solitary predators being locally specialized onT. pinivorain the high‐density area.

Within a population of the butterfly Euphydryas editha that oviposits predominantly on two host species, heritable variation in postalighting oviposition preference was found. In a separate experiment, oviposition preference of adult females was found to be correlated with offspring performance (growth). There was a significant tendency for offspring to perform better on the host species that their female parent preferred. Analysis of the data showed that no single factor, neither maternal preference nor the host species on which the offspring were raised, accounted for any significant variation in larval performance. However, the effect of the interaction between host species and maternal preference on offspring performance was highly significant. These findings imply specialization in both oviposition preference and offspring performance by individuals within a single population. With present evidence, this preference-performance correlation is likely to be genetic. However, as in previous studies, other interpretations cannot be excluded.

The sugarcane borer, Diatraea saccharalis (Fabricius) (Lepidoptera: Crambidae), is a major agronomic pest in the southeastern USA. This study was conducted to investigate the performance of D. saccharalis on eight rice cultivars widely grown in Louisiana and to assess the relationship between oviposition preference and offspring performance. In a previous study, oviposition preference of sugarcane borers was found to vary on the same eight rice cultivars. In this study, variation in larval performance on the eight cultivars was observed. The overall performance of larvae in terms of relative growth rate and boring success of larvae was highest on Priscilla and Cocodrie. Larval performance was lower on the hybrid XL723, the Clearfield (herbicide‐resistant) variety CL161, and the medium grain Bengal. Also, larvae took more time to enter into the stem of the hybrid XL723 compared with Cocodrie. Significant positive correlations were observed among relative growth rate and boring success and between larval performance and oviposition preference. Significant positive correlations among measures of larval performance and oviposition preference suggest the operation of a common resistance mechanism. Results in laboratory and greenhouse extended into the field, where Cocodrie and Cheniere were the most damaged cultivars, whereas CL161 and the medium grain Jupiter were least injured in terms of average number of stem borer entry/exit holes per plant. Results from this study could contribute to the use of resistant cultivars in an integrated management program for stem borers and the resistant cultivars identified in this study could be used as sources of resistance in breeding programs for resistance against stem borers.

A cross-sectional survey was conducted in West Kalimantan (Borneo), Indonesia to geographically profile hepatitis E virus (HEV) prevalence in the riverine areas recognized as the foci of epidemic HEV transmission in 1987. Additionally, a contiguous, although distinct, population with no identifiable historical exposure to epidemic HEV was surveyed downstream for comparative purposes. Eight hundred eighty-five sera were assayed by enzyme immunoabsorbent assay for anti-HEV IgG and anti-hepatitis A virus (HAV) IgG markers. A very high percent (90%) of both the outbreak and comparison populations was anti-HAV IgG positive by the age of nine years. In contrast, the prevalence of anti-HEV IgG in the outbreak area (50%) was significantly higher than in the comparison area (23%) (P < 0.0001). In both the outbreak and comparison areas, anti-HEV IgG prevalence increased with age ( < 0.0001), except for the group > or = 50 years of age. The prevalence (53%) of antibody to HEV in the population > or = seven years of age from the outbreak area (alive during the actual 1987 outbreak) was significantly (P < 0.0001) greater than among the children < seven years of age (born after the outbreak) (15%). However, anti-HEV IgG prevalence among the population from the comparison area did not differ significantly between the > or = seven- (23%) and < seven- (20%) year-old age groups. The percentage of anti-HEV IgG-positive individuals among males (47%) from the outbreak area was lower (P < 0.05) compared with females (55%). While overall usage of river water for drinking purposes was not universal, dependence on river water as a primary source was significantly higher (P < 0.001) in households from the outbreak area (60%) compared with the comparison area (30%). This study indicates persistence of an anti-HEV IgG response in a large percentage of the population seven years after an epidemic of HEV infections. Also, the relatively high prevalence (15%) of anti-HEV in children < seven years of age from the outbreak area reflects continuing, sporadic infections.

Preface. Acknowledgement. 1. Are island forests vulnerable to invasive defoliators? M.K. Kay. 2. Changing forest communities: role of tree resistance to insects in insect invasions and tree introductions F. Lieutier. 3. Southern hemisphere exotic pine plantations threatened by insect pests and their associated fungal pathogens M.J. Wingfield et al. 4. Native insects colonizing introduced tree species - patterns and potential risks P. Dalin and C. Bjorkman. 5. Biological pest control in mix and match forests R.G. van Driesche. 6. Impacts of insects in forest landscapes: implications for forest health management R.N. Coulson and F.M. Stephen. 7. Insect populations in relation to environmental change in forests of temperate Europe A. Battisti. 8. Synecology of Wasmannia auropunctata, an invasive ant species (Hymenoptera: Formicidae), in continuous and fragmented areas in the Brazilian Atlantic forest C.R.F. Brandao and R.R. Silva. 9. Changing relationships among biodiversity, management, and biosecurity in managed and unmanaged forests L.M. Hanks. 10. Changing the mix: new rules in regulating herbivore populations T.D. Paine. Keyword index. Species index.

What selection pressures drive the evolution of offspring size? Answering this fundamental question for any species requires an understanding of the relationship between offspring size and offspring fitness. A major goal of evolutionary ecologists has been to estimate this critical relationship, but for organisms with complex lifecycles, logistical constraints restrict most studies to early life-history stages only. Here, we examine the relationship between offspring size and offspring performance in the field across multiple life-history stages and across generations in a marine invertebrate .We then use these data to parameterise a simple optimality model to generate predictions of optimal offspring size and determined whether these predictions depended on which estimate of offspring performance was used. We found that offspring size had consistently positive effects on performance (estimated as post-metamorphic growth, fecundity and reproductive output). We also found that manipulating the experience of offspring during the larval phase changed the way in which offspring size affects performance: offspring size affected post-metamorphic growth when larvae were allowed to settle immediately but offspring size affected survival when larvae were forced to swim prior to settlement. Despite finding consistently positive effects of offspring size, early measures of the effect of offspring size resulted in the systematic underestimation of optimal offspring size. Surprisingly, the amount of variation in offspring performance that offspring size explained decreased with increasing time in the field but the steepness of the relationship between offspring size and performance actually increased. Our results suggest caution should be exercised when empirically examining offspring size effects - it may not be appropriate to assume that early measures are a good reflection of the actual relationship between offspring size and fitness. © 2009 Oikos.

The effects of irrigation and simulated acid rain on the growth performance of pine sawfly larvae (Neodiprion sertifer) and on the oviposition behaviour and fecundity of adult females were studied in a subarctic area. Two anatomical traits and resin acid concentrations of pine needles from experimental trees were measured and bioassays undertaken to determine their effect on herbivores. Twice as many females oviposited on trees receiving only ambient rain (dry control) as on irrigated trees. Larval growth performance was also slightly better on dry control trees. However, larval survival, relative larval growth rate, female fecundity and oviposition preference did not differ between acid-treated (pH 3 ; both H 2 SO 4 and HNO 3 ) and irrigated control (pH 6) trees. So, the oviposition preference of European pine sawfly is modified more by the amount of rainfall than by its pH. The concentrations of measured total and some individual resin acids showed significant negative correlations with larval survival and relative growth rate. There was a weak correlation between resin duct area, expressed as a proportion of needle cross-sectional area, and female oviposition choice. Our results suggest that the oviposition preference expressed by this eruptive species is a more sensitive indicator of host plant quality than larval performance. Secondly, for N. sertifer, needle resin acid content and related structural characteristics modify larval performance and female oviposition behaviour more than needle surface thickness.

Biological invasions in forest ecosystems are recognised as a global scale challenge. However, our current knowledge of invasive alien forest insect species (IAFIS) in Romania is still lacking and rather insufficient to support clear policies. The aim of this study is to contribute to a better understanding of the distribution of IAFIS in the Muntenia and Dobrogea provinces of Romania. For this purpose, direct observations and insect collections were made two consecutive years, in 76 locations. Of these, 67 sampling locations forming a long field observation route were visited only once, and nine sampling locations were fixed monitoring points with primed traps used in first year for regular collection of insects at 1-2-week intervals. Seventeen non-native forest insect species from five orders and 11 families were detected: Aproceros leucopoda, Cameraria ohridella, Corythucha arcuata, Cydalima perspectalis, Dasineura gleditchiae, Eopineus strobus, Euura tibialis, Gilletteella cooleyi, Hyphantria cunea, Ips duplicatus, Macrosaccus robiniella, Neoclytus acuminatus, Obolodiplosis robiniae, Parectopa robiniella, Phyllonorycter issikii, Prociphilus fraxinifolii and Xylosandrus germanus. Eleven IAFIS species originate from North America, four from Asia and two from Europe. All detected species were previously reported from Romania, but eight species had not been recorded in these two provinces. Seven species live on native tree species, while 10 thrive on exotic trees or shrubs. The highest number of IAFIS was recorded in urban areas. Except for E. tibialis, all found IAFIS have a significant ecological or economic impact across invaded habitats. The risks posed by biological invasions need more thorough consideration that deserves greater attention and scientific support.