Plant hydraulics at the heart of plant, crops and ecosystem functions in the face of climate change.

Plant hydraulics is crucial for assessing the plants' capacity to extract and transport water from the soil up to their aerial organs. Along with their capacity to exchange water between plant compartments and regulate evaporation, hydraulic properties determine plant water relations, water status and susceptibility to pathogen attacks. Consequently, any variation in the hydraulic characteristics of plants is likely to significantly impact various mechanisms and processes related to plant growth, survival and production, as well as the risk of biotic attacks and forest fire behaviour. However, the integration of hydraulic traits into disciplines such as plant pathology, entomology, fire ecology or agriculture can be significantly improved. This review examines how plant hydraulics can provide new insights into our understanding of these processes, including modelling processes of vegetation dynamics, illuminating numerous perspectives for assessing the consequences of climate change on forest and agronomic systems, and addressing unanswered questions across multiple areas of knowledge.

Seed predation-induced Allee effects, seed dispersal and masting jointly drive the diversity of seed sources during population expansion.

The environmental factors affecting plant reproduction and effective dispersal, in particular biotic interactions, have a strong influence on plant expansion dynamics, but their demographic and genetic consequences remain an understudied body of theory. Here, we use a mathematical model in a one-dimensional space and on a single reproductive period to describe the joint effects of predispersal seed insect predators foraging strategy and plant reproduction strategy (masting) on the spatio-temporal dynamics of seed sources diversity in the colonisation front of expanding plant populations. We show that certain foraging strategies can result in a higher seed predation rate at the colonisation front compared to the core of the population, leading to an Allee effect. This effect promotes the contribution of seed sources from the core to the colonisation front, with long-distance dispersal further increasing this contribution. As a consequence, our study reveals a novel impact of the predispersal seed predation-induced Allee effect, which mitigates the erosion of diversity in expanding populations. We use rearrangement inequalities to show that masting has a buffering role: it mitigates this seed predation-induced Allee effect. This study shows that predispersal seed predation, plant reproductive strategies and seed dispersal patterns can be intermingled drivers of the diversity of seed sources in expanding plant populations, and opens new perspectives concerning the analysis of more complex models such as integro-difference or reaction-diffusion equations.

Masting is uncommon in trees that depend on mutualist dispersers in the context of global climate and fertility gradients.

The benefits of masting (volatile, quasi-synchronous seed production at lagged intervals) include satiation of seed predators, but these benefits come with a cost to mutualist pollen and seed dispersers. If the evolution of masting represents a balance between these benefits and costs, we expect mast avoidance in species that are heavily reliant on mutualist dispersers. These effects play out in the context of variable climate and site fertility among species that vary widely in nutrient demand. Meta-analyses of published data have focused on variation at the population scale, thus omitting periodicity within trees and synchronicity between trees. From raw data on 12 million tree-years worldwide, we quantified three components of masting that have not previously been analysed together: (i) volatility, defined as the frequency-weighted year-to-year variation; (ii) periodicity, representing the lag between high-seed years; and (iii) synchronicity, indicating the tree-to-tree correlation. Results show that mast avoidance (low volatility and low synchronicity) by species dependent on mutualist dispersers explains more variation than any other effect. Nutrient-demanding species have low volatility, and species that are most common on nutrient-rich and warm/wet sites exhibit short periods. The prevalence of masting in cold/dry sites coincides with climatic conditions where dependence on vertebrate dispersers is less common than in the wet tropics. Mutualist dispersers neutralize the benefits of masting for predator satiation, further balancing the effects of climate, site fertility and nutrient demands.

Limits to reproduction and seed size-number trade-offs that shape forest dominance and future recovery.

The relationships that control seed production in trees are fundamental to understanding the evolution of forest species and their capacity to recover from increasing losses to drought, fire, and harvest. A synthesis of fecundity data from 714 species worldwide allowed us to examine hypotheses that are central to quantifying reproduction, a foundation for assessing fitness in forest trees. Four major findings emerged. First, seed production is not constrained by a strict trade-off between seed size and numbers. Instead, seed numbers vary over ten orders of magnitude, with species that invest in large seeds producing more seeds than expected from the 1:1 trade-off. Second, gymnosperms have lower seed production than angiosperms, potentially due to their extra investments in protective woody cones. Third, nutrient-demanding species, indicated by high foliar phosphorus concentrations, have low seed production. Finally, sensitivity of individual species to soil fertility varies widely, limiting the response of community seed production to fertility gradients. In combination, these findings can inform models of forest response that need to incorporate reproductive potential.

Globally, tree fecundity exceeds productivity gradients.

Lack of tree fecundity data across climatic gradients precludes the analysis of how seed supply contributes to global variation in forest regeneration and biotic interactions responsible for biodiversity. A global synthesis of raw seedproduction data shows a 250-fold increase in seed abundance from cold-dry to warm-wet climates, driven primarily by a 100-fold increase in seed production for a given tree size. The modest (threefold) increase in forest productivity across the same climate gradient cannot explain the magnitudes of these trends. The increase in seeds per tree can arise from adaptive evolution driven by intense species interactions or from the direct effects of a warm, moist climate on tree fecundity. Either way, the massive differences in seed supply ramify through food webs potentially explaining a disproportionate role for species interactions in the wet tropics.

Incremental Sampling Methodology for improved characterization of Agent Orange dioxin in Vietnam soil and sediment.

The Danang airport in Vietnam was used heavily by US forces in the 1960s and 1970s. In 2018, the remediation of dioxin contamination at the airport resulting from Agent Orange use and management was completed by the US government. Generation of reliable, defensible, and cost-effective confirmation sampling data over large areas and for large volumes was a significant challenge. Traditional discrete and composite sampling methods were utilized to assess dioxin concentrations and the extent of contamination present at the airport prior to remediation. Confirmation sampling was performed after excavation of contaminated soil and sediment across the entire 28-ha site to demonstrate that remaining concentrations met the Vietnamese standards for the airport site. The volume of excavated soil and sediment was 162,567 m3. Incremental Sampling Methodology (ISM) was used for confirmation sampling to provide a defensible estimate of the mean concentration of dioxin remaining after excavation. The use of ISM revealed that traditional methods underestimated the volume of material requiring treatment. ISM also decreased sampling variability dramatically and provided more reliable estimates of true mean concentrations in an area when compared with traditional methods. The use of ISM: 1) better captured distributional heterogeneity and decreased variability between samples from the same DU by 64%; 2) resulted in low variability between duplicate analyses of the same sample (12%), indicating a reduction in compositional heterogeneity; 3) did not underestimate contaminant levels; and, 4) increased the frequency that excavation boundaries met project goals by 61%, when compared with traditional sampling.

Signature of mid-Pleistocene lineages in the European silver fir (Abies alba Mill.) at its geographic distribution margin.

In a conservation and sustainable management perspective, we identify the ecological, climatic, and demographic factors responsible for the genetic diversity patterns of the European silver fir (Abies alba Mill.) at its southwestern range margin (Pyrenees Mountains, France, Europe). We sampled 45 populations throughout the French Pyrenees and eight neighboring reference populations in the Massif Central, Alps, and Corsica. We genotyped 1,620 individuals at three chloroplast and ten nuclear microsatellite loci. We analyzed within- and among-population genetic diversity using phylogeographic reconstructions, tests of isolation-by-distance, Bayesian population structure inference, modeling of demographic scenarios, and regression analyses of genetic variables with current and past environmental variables. Genetic diversity decreased from east to west suggesting isolation-by-distance from the Alps to the Pyrenees and from the Eastern to the Western Pyrenees. We identified two Pyrenean lineages that diverged from a third Alpine-Corsica-Massif Central lineage 0.8 to 1.1 M years ago and subsequently formed a secondary contact zone in the Central Pyrenees. Population sizes underwent contrasted changes, with a contraction in the west and an expansion in the east. Glacial climate affected the genetic composition of the populations, with the western genetic cluster only observed in locations corresponding to the coldest past climate and highest elevations. The eastern cluster was observed over a larger range of temperatures and elevations. All demographic events shaping the current spatial structure of genetic diversity took place during the Mid-Pleistocene Transition, long before the onset of the Holocene. The Western Pyrenees lineage may require additional conservation efforts, whereas the eastern lineage is well protected in in situ gene conservation units. Due to past climate oscillations and the likely emergence of independent refugia, east-west oriented mountain ranges may be important reservoir of genetic diversity in a context of past and ongoing climate change in Europe.

Is there tree senescence? The fecundity evidence.

Despite its importance for forest regeneration, food webs, and human economies, changes in tree fecundity with tree size and age remain largely unknown. The allometric increase with tree diameter assumed in ecological models would substantially overestimate seed contributions from large trees if fecundity eventually declines with size. Current estimates are dominated by overrepresentation of small trees in regression models. We combined global fecundity data, including a substantial representation of large trees. We compared size-fecundity relationships against traditional allometric scaling with diameter and two models based on crown architecture. All allometric models fail to describe the declining rate of increase in fecundity with diameter found for 80% of 597 species in our analysis. The strong evidence of declining fecundity, beyond what can be explained by crown architectural change, is consistent with physiological decline. A downward revision of projected fecundity of large trees can improve the next generation of forest dynamic models.

Effects of Climate and Host Age on Flight Activity, Infestation Percentage, and Intensity by Coptoborus ochromactonus (Coleoptera: Curculionidae: Scolytinae) in Commercial Balsa Plantations of Ecuador.

Coptoborus ochromactonus (Smith and Cognato) is one of the most common and important pests of balsa, Ochroma pyramidale (Cav. Ex Lam. Urb.), an economic pillar of the wood industry in Ecuador. Commercial balsa plantations have been expanded from humid to dry climate areas to limit insect damage, but basic knowledge is still lacking on the interaction of C. ochromactonus activity with variation in climate and plantation age. We investigated the effects of climate and host age on the seasonal flight activity of C. ochromactonus and its infestation rate and intensity, as well as the effect of age and individual infestation intensity on balsa dieback. Experiments were conducted in 1-, 2-, and 3-yr-old commercial balsa plantations located in areas of humid or dry climates. Seasonal flight activity (monitored with baited traps) differed between study sites and seasons. Increased flight activity was significantly correlated with higher relative humidity, higher mean temperature, and reduced precipitation during the dry season in the humid site and with increased mean and minimum temperature and increased precipitation during the rainy season in the dry site. Infestation rates by C. ochromactonus significantly increased with plantation age, especially in the humid site. Intensity of individual infestations (measured as number of successful insect entry holes per tree) was significantly influenced by both climate and plantation age, and it was particularly severe on 3-yr-old trees in the dry site. Percentage of foliage loss significantly increased with infestation intensity. Overall, our results can be relevant for devising preventive measures and suitable management strategies for this emerging pest in Ecuadorian wood plantations.

Spatial and temporal patterns of a pulsed resource dynamically drive the distribution of specialist herbivores.

Patterns and drivers of the spatio-temporal distribution of herbivores are key elements of their ecological and evolutionary impacts on plant populations. Herbivore spatial distributions may be influenced by increased (RCH: resource concentration hypothesis) or decreased (RDH: resource dilution hypothesis) resource densities, but the effect of temporal variations in resource densities on such distributions remains poorly documented. We used a survey of a masting tree species and its seed predators in Southeastern France to address the effect of a host's pulsed resource on the spatio-temporal distributions of highly specialized insect herbivores feeding on seeds. Variations in both resource and seed predator densities were assessed by estimating seed production and seed infestation rates in focus trees during 10 consecutive years. We found increasing seed infestation rates with decreasing host tree densities in years of low seed production, indicating a RDH pattern of seed predators. However, such pattern was not persistent in years of high seed production during which seed infestation rates did not depend on host tree densities. We showed that temporal variations in resource density can lead to transience of seed predator spatial distribution. This study highlights how predictions of plant-herbivore interactions in natural ecosystems may rely on temporal components underlying RCH and RDH hypotheses.

Bark and Ambrosia Beetle (Coleoptera: Curculionidae: Scolytinae) Diversity in Natural and Plantation Forests in Ecuador.

The Scolytinae is highly diversified in tropical forests, but richness and abundance patterns within most Ecuadorian forest habitat types are not yet characterized. In this study, we assessed patterns of variation in Scolytinae richness, abundance, and species composition in a primary and a secondary natural forest, and a commercial balsa plantation in Ecuador. We conducted a 1-yr survey of Scolytinae communities with baited traps and measured associated environmental variables. In total, 18,169 Scolytinae individuals were captured and comprised 85 species, 16 genera, and six tribes. In the natural forests, main indicator species were Xylosandrus morigerus, Xyleborus affinis, Xyleborus sp.02, and Corthylus sp.01, whereas all species of Hypothenemus were indicator species in the balsa plantation. The exotic Premnobius cavipennis (Ipini), Xylosandrus compactus, and Xylosandrus morigerus were indicator species for the natural forests. We provide evidence that commercial balsa plantations provide abundant favorable resources for native and exotic scolytines in Ecuador, and that scolytine communities in natural forest and in plantations are more likely to differ in their species composition than in their cumulated species richness. In all habitats, species composition, abundance, and species richness showed temporal patterns of variation that coincided with seasonal variations in climatic conditions, with highest records during the coldest and driest months in the primary forest and the balsa plantation. We provide new knowledge on the native Ecuadorian scolytine fauna and a foundation for the monitoring for potential scolytine pest species of natural and planted tropical forest ecosystems.

The ecology of predispersal insect herbivory on tree reproductive structures in natural forest ecosystems.

Plant-insect interactions are key model systems to assess how some species affect the distribution, the abundance, and the evolution of others. Tree reproductive structures represent a critical resource for many insect species, which can be likely drivers of demography, spatial distribution, and trait diversification of plants. In this review, we present the ecological implications of predispersal herbivory on tree reproductive structures by insects (PIHR) in forest ecosystems. Both insect's and tree's perspectives are addressed with an emphasis on how spatiotemporal variation and unpredictability in seed availability can shape such particular plant-animal interactions. Reproductive structure insects show strong trophic specialization and guild diversification. Insects evolved host selection and spatiotemporal dispersal strategies in response to variable and unpredictable abundance of reproductive structures in both space and time. If PIHR patterns have been well documented in numerous systems, evidences of the subsequent demographic and evolutionary impacts on tree populations are still constrained by time-scale challenges of experimenting on such long-lived organisms, and modeling approaches of tree dynamics rarely consider PIHR when including biotic interactions in their processes. We suggest that spatially explicit and mechanistic approaches of the interactions between individual tree fecundity and insect dynamics will clarify predictions of the demogenetic implications of PIHR in tree populations. In a global change context, further experimental and theoretical contributions to the likelihood of life-cycle disruptions between plants and their specialized herbivores, and to how these changes may generate novel dynamic patterns in each partner of the interaction are increasingly critical.

Host-Parasite Interactions from the Inside: Plant Reproductive Ontogeny Drives Specialization in Parasitic Insects.

Host plant interactions are likely key drivers of evolutionary processes involved in the diversification of phytophagous insects. Granivory has received substantial attention for its crucial role in shaping the interaction between plants and their seed parasites, but fine-scale mechanisms explaining the role of host plant reproductive biology on specialization of seed parasites remain poorly described. In a comparative approach using plant histological techniques, we tested the hypotheses that different seed parasite species synchronize their life cycles to specific stages in seed development, and that the stage they target depends on major differences in seed development programs. In a pinaceous system, seed storage products are initiated before ovule fertilization and the wasps target the ovule's nucellus during megagametogenesis, a stage at which larvae may benefit from the by-products derived from both secreting cells and dying nucellar cells. In a cupressaceous system, oviposition activity peaks later, during embryogenesis, and the wasps target the ovule's megagametophyte where larvae may benefit from cell disintegration during embryogenesis. Our cytohistological approach shows for the first time how, despite divergent oviposition targets, different parasite species share a common strategy that consists of first competing for nutrients with developing plant structures, and then consuming these developed structures to complete their development. Our results support the prediction that seed developmental program is an axis for specialization in seed parasites, and that it could be an important parameter in models of their ecological and taxonomic divergence. This study provides the basis for further investigating the possibility of the link between plant ontogeny and pre-dispersal seed parasitism.

Predictors for dioxin accumulation in residents living in Da Nang and Bien Hoa, Vietnam, many years after Agent Orange use.

Agent Orange (AO) was the main defoliant used by the US in Vietnam from 1961 to 1971; AO was contaminated with dioxin (2,3,7,8-tetrachlorodibenzo-p-dioxin, or TCDD). Three major dioxin "hot spots" remain from previous AO storage and use at former US bases at Bien Hoa, Da Nang, and Phu Cat, posing potential health risks for Vietnamese living on or near these hot spots. We evaluated potential risk factors contributing to serum TCDD levels in Vietnamese residents at and near contaminated sites in Da Nang and Bien Hoa, Vietnam. We used multiple linear regression to analyze possible associations of blood dioxin concentrations with demographic, socioeconomic, lifestyle, and dietary risk factors for residents living on or near these hot spots. For the Da Nang study, fish farming on the site, living on property flooded from monsoon rains, and age were among the factors showing significant positive associations with serum TCDD concentrations. For the Bien Hoa study, fish farmers working at this site and their immediate family members had significantly higher serum TCDD concentrations. Our results suggest that water-related activities, especially fish-farming, at the hot spots increased the risk of exposure to dioxin.

Linking niche theory to ecological impacts of successful invaders: insights from resource fluctuation-specialist herbivore interactions.

Theories of species coexistence and invasion ecology are fundamentally connected and provide a common theoretical framework for studying the mechanisms underlying successful invasions and their ecological impacts. Temporal fluctuations in resource availability and differences in life-history traits between invasive and resident species are considered as likely drivers of the dynamics of invaded communities. Current critical issues in invasion ecology thus relate to the extent to which such mechanisms influence coexistence between invasive and resident species and to the ability of resident species to persist in an invasive-dominated ecosystem. We tested how a fluctuating resource, and species trait differences may explain and help predict long-term impacts of biological invasions in forest specialist insect communities. We used a simple invasion system comprising closely related invasive and resident seed-specialized wasps (Hymenoptera: Torymidae) competing for a well-known fluctuating resource and displaying divergent diapause, reproductive and phenological traits. Based on extensive long-term field observations (1977-2010), we developed a combination of mechanistic and statistical models aiming to (i) obtain a realistic description of the population dynamics of these interacting species over time, and (ii) clarify the respective contributions of fluctuation-dependent and fluctuation-independent mechanisms to long-term impact of invasion on the population dynamics of the resident wasp species. We showed that a fluctuation-dependent mechanism was unable to promote coexistence of the resident and invasive species. Earlier phenology of the invasive species was the main driver of invasion success, enabling the invader to exploit an empty niche. Phenology also had the greatest power to explain the long-term negative impact of the invasive on the resident species, through resource pre-emption. This study provides strong support for the critical role of species differences in interspecific competition outcomes within animal communities. Our mechanistic-statistical approach disentangles the critical drivers of novel species assemblages resulting from intentional and non-intentional introductions of non-native species.

Public health assessment of dioxin-contaminated fish at former US airbase, Bien Hoa, Vietnam.

Ponds at the former US airbase at Ben Hoa, Vietnam are contaminated with Agent Orange. The ponds had been used for aquaculture, and in all likelihood, fish from those ponds have been sold to the public. We assessed human exposure to 2,3,7,8-tetrachloro-dibenzo-dioxin (2,3,7,8-TCDD) in fish samples from the ponds. For on-base tilapia, muscle concentrations 2,3,7,8-TCDD ranged from 1.4 to 32.7 pg/g. Fat concentrations ranged from 73.3 to 3990 pg/g. Estimated human exposure doses exceed international guidelines and exceed 2,3,7,8-TCDD's lowest adverse effect levels. The Bien Hoa fishponds are a completed human pathway for TCDD exposure.

Reconstruction of a windborne insect invasion using a particle dispersal model, historical wind data, and Bayesian analysis of genetic data.

Understanding how invasive species establish and spread is vital for developing effective management strategies for invaded areas and identifying new areas where the risk of invasion is highest. We investigated the explanatory power of dispersal histories reconstructed based on local-scale wind data and a regional-scale wind-dispersed particle trajectory model for the invasive seed chalcid wasp Megastigmus schimitscheki (Hymenoptera: Torymidae) in France. The explanatory power was tested by: (1) survival analysis of empirical data on M. schimitscheki presence, absence and year of arrival at 52 stands of the wasp's obligate hosts, Cedrus (true cedar trees); and (2) Approximate Bayesian analysis of M. schimitscheki genetic data using a coalescence model. The Bayesian demographic modeling and traditional population genetic analysis suggested that initial invasion across the range was the result of long-distance dispersal from the longest established sites. The survival analyses of the windborne expansion patterns derived from a particle dispersal model indicated that there was an informative correlation between the M. schimitscheki presence/absence data from the annual surveys and the scenarios based on regional-scale wind data. These three very different analyses produced highly congruent results supporting our proposal that wind is the most probable vector for passive long-distance dispersal of this invasive seed wasp. This result confirms that long-distance dispersal from introduction areas is a likely driver of secondary expansion of alien invasive species. Based on our results, management programs for this and other windborne invasive species may consider (1) focusing effort at the longest established sites and (2) monitoring outlying populations remains critically important due to their influence on rates of spread. We also suggest that there is a distinct need for new analysis methods that have the capacity to combine empirical spatiotemporal field data, genetic data, and environmental data to investigate dispersal and invasion.

Temporal population genetics of time travelling insects: a long term study in a seed-specialized wasp.

Many animal species experiencing spatial or interannual fluctuations of their environment are capable of prolonged diapause, a kind of dormancy that extends over more than one year. Such a prolonged diapause is commonly perceived as a temporal demographic refuge in stochastic environments, but empirical evidence is still lacking of its consequences on temporal population genetic structures. In this long-term study, we investigated how a particular pattern of prolonged diapause may influence the temporal population genetics of the invasive seed-specialized wasp Megastigmus schimitscheki (Hymenoptera: Torymidae) in southeastern France. We characterized the diapause strategy of M. schimitscheki using records of emergence from diapause in 97 larval cohorts, and we conducted a temporal population genetic study on a natural invasive wasp population sampled during ten consecutive years (1999-2008) using polymorphic microsatellite markers. We found that M. schimitscheki can undergo a prolonged diapause of up to five years and displays two main adult emergence peaks after two and four years of diapause. Such a bimodal and atypical pattern did not disrupt temporal gene flow between cohorts produced in even and in odd years during the period of the study. Unexpectedly, we found that this wasp population consisted of two distinct genetic sub-populations that strongly diverged in their diapause strategies, with very few admixed individuals. One of the sub-populations displayed both short and prolonged diapause (2 and 4 years respectively) in equal proportions, whereas the other sub-population displayed mainly short diapause. This study provided empirical evidence that prolonged diapause phenotypes can substantially contribute to reproduction and impact temporal genetic structures. Prolonged diapause is likely to act as both demographic and genetic refuges for insect populations living in fluctuating environments.

Insecticide resistance may enhance the response to a host-plant volatile kairomone for the codling moth, Cydia pomonella (L.).

The behavioral and electroantennographic responses of Cydia pomonella (L.) to the ripe pear volatile ethyl (2E,4Z)-2,4-decadienoate (Et-E,Z-DD), were compared in insecticide-susceptible and -resistant populations originating from southern France. A dose-response relationship to this kairomonal attractant was established for antennal activity and did not reveal differences between susceptible and resistant strains. Conversely, males of the laboratory strains expressing metabolic [cytochrome P450-dependent mixed-function oxidases (mfo)] or physiological (kdr-type mutation of the sodium-channel gene) resistance mechanisms exhibited a significantly higher response to Et-E,Z-DD than those of the susceptible strain in a wind tunnel experiment. No response of the females to this kairomone could be obtained in our wind-tunnel conditions. In apple orchards, mfo-resistant male moths were captured at significantly higher rates in kairomone-baited traps than in traps baited with the sex pheromone of C. pomonella. Such a differential phenomenon was not verified for the kdr-resistant insects, which exhibited a similar response to both the sex pheromone and the kairomonal attractant in apple orchards. Considering the widespread distribution of metabolic resistance in European populations of C. pomonella and the enhanced behavioral response to Et-E,Z-DD in resistant moths, the development of control measures based on this kairomonal compound would be of great interest for the management of insecticide resistance in this species.

Modelling the interactions between phenology and insecticide resistance genes in the codling moth Cydia pomonella.

In the codling moth Cydia pomonella (L), insecticide resistance genes have been associated with pleiotropic effects affecting phenology. In this paper, we investigated whether an increase in the frequency of insecticide resistance in field populations of C pomonella was likely to entail significant divergences in the temporal occurrence of both susceptible and insecticide-resistant individuals. For this purpose, we built a phenological model that provided suitable predictions of the distinct and diverging seasonal evolutions of populations of a susceptible and two insecticide-resistant (at two and three loci) homozygous genotypes of C pomonella. Model simulations for each genotype were further compared with pheromone trap catches recorded in a field insecticide-treated population over an 8-year period (from 1992 to 2000), which reflected the progressive annual increase in the frequency of resistance in southeastern France. We found a significant delay in field adult emergence relative to those predicted by the homozygous susceptible model, and the magnitude of such a delay was positively correlated with increasing frequencies of insecticide resistance in the sampled field population of C pomonella. Adult emergence predicted in the theoretical population that was homozygous for resistance at two loci converged with those recorded in the field during the investigated 8-year period. This suggested that the pleiotropic effects of resistance were likely to result in a significant phenological segregation of insecticide-resistant alleles in the field. The results of this study emphasized the potential for pest populations exposed to chemical selection to evolve qualitatively with respect to phenology. This may raise critical questions regarding the use of phenological modelling as a forecasting tool for appropriate resistance management strategies that would take into account the diverging seasonal evolutions of both insecticide resistance and susceptibility.