Predicting bee community responses to land-use changes: Effects of geographic and taxonomic biases.

Land-use change and intensification threaten bee populations worldwide, imperilling pollination services. Global models are needed to better characterise, project, and mitigate bees' responses to these human impacts. The available data are, however, geographically and taxonomically unrepresentative; most data are from North America and Western Europe, overrepresenting bumblebees and raising concerns that model results may not be generalizable to other regions and taxa. To assess whether the geographic and taxonomic biases of data could undermine effectiveness of models for conservation policy, we have collated from the published literature a global dataset of bee diversity at sites facing land-use change and intensification, and assess whether bee responses to these pressures vary across 11 regions (Western, Northern, Eastern and Southern Europe; North, Central and South America; Australia and New Zealand; South East Asia; Middle and Southern Africa) and between bumblebees and other bees. Our analyses highlight strong regionally-based responses of total abundance, species richness and Simpson's diversity to land use, caused by variation in the sensitivity of species and potentially in the nature of threats. These results suggest that global extrapolation of models based on geographically and taxonomically restricted data may underestimate the true uncertainty, increasing the risk of ecological surprises.

Pollen mixing in pollen generalist solitary bees: a possible strategy to complement or mitigate unfavourable pollen properties?

Generalist herbivorous insects, which feed on plant tissue that is nutritionally heterogeneous or varies in its content of secondary metabolites, often benefit from dietary mixing through more balanced nutrient intake or reduced exposure to harmful secondary metabolites. Pollen is similarly heterogeneous as other plant tissue in its content of primary and secondary metabolites, suggesting that providing their offspring with mixed pollen diets might be a promising strategy for pollen generalist bees to complement nutrient imbalances or to mitigate harmful secondary metabolites of unfavourable pollen. In the present study, we compared larval performance of the pollen generalist solitary bee species Osmia cornuta (Megachilidae) on five experimental pollen diets that consisted of different proportions of unfavourable pollen diet of Ranunculus acris (Ranunculaceae) and favourable pollen diet of Sinapis arvensis (Brassicaceae). In addition, we microscopically analysed the pollen contained in the scopal brushes of field-collected females of O. cornuta and three closely related species to elucidate to what degree these pollen generalist bees mix pollen of different hosts in their brood cells. In striking contrast to a pure Ranunculus pollen diet, which had a lethal effect on most developing larvae of O. cornuta, larval survival, larval development time and adult body mass of both males and females remained nearly unaffected by the admixture of up to 50% of Ranunculus pollen diet to the larval food. Between 42% and 66% of all female scopal pollen loads analysed contained mixtures of pollen from two to six plant families, indicating that pollen mixing is a common behaviour in O. cornuta and the three related bee species. The present study provides the first evidence that the larvae of pollen generalist bees can benefit from the nutrient content of unfavourable pollen without being negatively affected by its unfavourable chemical properties if such pollen is mixed with favourable pollen. We conclude that the widespread pollen mixing by females of pollen generalist bees should also be considered as a possible strategy to exploit flowers with unfavourable pollen and to optimize larval food quality.

High genetic diversity and structured populations of the oriental fruit moth in its range of origin.

The oriental fruit moth Grapholita ( = Cydia) molesta is a key fruit pest globally. Despite its economic importance, little is known about its population genetics in its putative native range that includes China. We used five polymorphic microsatellite loci and two mitochondrial gene sequences to characterize the population genetic diversity and genetic structure of G. molesta from nine sublocations in three regions of a major fruit growing area of China. Larval samples were collected throughout the season from peach, and in late season, after host switch by the moth to pome fruit, also from apple and pear. We found high numbers of microsatellite alleles and mitochondrial DNA haplotypes in all regions, together with a high number of private alleles and of haplotypes at all sublocations, providing strong evidence that the sampled area belongs to the origin of this species. Samples collected from peach at all sublocations were geographically structured, and a significant albeit weak pattern of isolation-by-distance was found among populations, likely reflecting the low flight capacity of this moth. Interestingly, populations sampled from apple and pear in the late season showed a structure differing from that of populations sampled from peach throughout the season, indicating a selective host switch of a certain part of the population only. The recently detected various olfactory genotypes in G. molesta may underly this selective host switch. These genetic data yield, for the first time, an understanding of population dynamics of G. molesta in its native range, and of a selective host switch from peach to pome fruit, which may have a broad applicability to other global fruit production areas for designing suitable pest management strategies.

Worldwide population genetic structure of the oriental fruit moth (Grapholita molesta), a globally invasive pest.

BACKGROUND: Invasive pest species have large impacts on agricultural crop yields, and understanding their population dynamics is important for ensuring food security. The oriental fruit moth Grapholita molesta is a cosmopolitan pest of stone and pome fruit species including peach and apple, and historical records indicate that it has invaded North and South America, Europe, Australia and Africa from its putative native range in Asia over the past century. RESULTS: We used 13 microsatellite loci, including nine newly developed markers, to characterize global population structure of G. molesta. Approximately 15 individuals from each of 26 globally distributed populations were genotyped. A weak but significant global pattern of isolation-by-distance was found, and G. molesta populations were geographically structured on a continental scale. Evidence does not support that G. molesta was introduced to North America from Japan as previously proposed. However, G. molesta was probably introduced from North America to The Azores, South Africa, and Brazil, and from East Asia to Australia. Shared ancestry was inferred between populations from Western Europe and from Brazil, although it remains unresolved whether an introduction occurred from Europe to Brazil, or vice versa. Both genetic diversity and levels of inbreeding were surprisingly high across the range of G. molesta and were not higher or lower overall in introduced areas compared to native areas. There is little evidence for multiple introductions to each continent (except in the case of South America), or for admixture between populations from different origins. CONCLUSIONS: Cross-continental introductions of G. molesta appear to be infrequent, which is surprising given its rapid worldwide expansion over the past century. We suggest that area-wide spread via transport of fruits and other plant materials is a major mechanism of ongoing invasion, and management efforts should therefore target local and regional farming communities and distribution networks.

A host-plant-derived volatile blend to attract the apple blossom weevil Anthonomus pomorum - the essential volatiles include a repellent constituent.

BACKGROUND: Plant volatiles are promising cues for trapping pest insects. This study started with a recently identified complex blend released by prebloom apple trees and aimed to reduce the number of compounds in the blend while maintaining the attraction of the target pest, the apple blossom weevil Anthonomus pomorum. An evaluation was made to determine whether attraction to plant volatiles is a general feature in this species. RESULTS: Laboratory-based bioassays with field-collected weevils demonstrated repellency by volatiles from the non-host walnut, indicating that preference for plant odours is not a general feature in this species. By a subtractive bioassay approach, the original number of compounds in the apple-plant-released blend was stepwise reduced from 12 to 6 while maintaining weevil attraction. This resulting blend was as attractive as the full blend and as a blossom-bud-carrying apple twig. It was found to be composed of two synergistically interacting constituents, of which the first containing benzenoids was behaviourally inactive, and the second comprising the remaining compounds was even repellent. CONCLUSIONS: This study enhances knowledge of the interaction of behaviourally effective constituents in complex odour blends and contributes to the development of an efficient monitoring system involving plant volatiles for the apple blossom weevil.

Molecular genetics and genomics generate new insights into invertebrate pest invasions.

Invertebrate pest invasions and outbreaks are associated with high social, economic, and ecological costs, and their significance will intensify with an increasing pressure on agricultural productivity as a result of human population growth and climate change. New molecular genetic and genomic techniques are available and accessible, but have been grossly underutilized in studies of invertebrate pest invasions, despite that they are useful tools for applied pest management and for understanding fundamental features of pest invasions including pest population demographics and adaptation of pests to novel and/or changing environments. Here, we review current applications of molecular genetics and genomics in the study of invertebrate pest invasions and outbreaks, and we highlight shortcomings from the current body of research. We then discuss recent conceptual and methodological advances in the areas of molecular genetics/genomics and data analysis, and we highlight how these advances will further our understanding of the demographic, ecological, and evolutionary features of invertebrate pest invasions. We are now well equipped to use molecular data to understand invertebrate dispersal and adaptation, and this knowledge has valuable applications in agriculture at a time when these are critically required.

Plant acclimation to elevated CO2 affects important plant functional traits, and concomitantly reduces plant colonization rates by an herbivorous insect.

Plants growing under elevated CO2 concentration may acclimatize to this environmental change by modification of chemical, physiological, and/or morphological traits. As a consequence, not only plant functioning but also plant-insect interactions might be altered, with important consequences particularly for agricultural systems. Whereas most studies have focused on the plant acclimation effects of elevated CO2 with regard to crop growth and productivity, acclimation effects on the behavioral response of insects associated with these plants have been largely neglected. In this study, we used a model system comprised of Brussels sprout Brassica oleraceae var. gemmifera and a specialized herbivorous insect, the cabbage aphid Brevicoryne brassicae, to test for the effects of various periods of exposure to an elevated (2× ambient) CO2 concentration on key plant functional traits and on host plant location behavior by the insect, assessed as plant colonization rates. Elevated CO2 had no measurable effect on colonization rates or total plant volatile emissions after a 2-week exposure, but it led to 15 and 26 % reductions in plant colonization rates after 6- and 10-week exposures, respectively. This reduction in plant colonization was associated with significant decreases in leaf stomatal conductance and plant volatile emission. Terpene emission, in particular, exhibited a great reduction after the 10-week exposure to elevated CO2. Our results provide empirical evidence that plants might acclimatize to a future increase in CO2, and that these acclimation responses might affect host plant choice and colonization behavior by herbivorous insects, which might be advantageous from the plant's perspective.

Too low to kill: concentration of the secondary metabolite ranunculin in buttercup pollen does not affect bee larval survival.

Growing evidence suggests that the freely accessible pollen of some plants is chemically protected against pollen-feeding flower visitors. For example, a diet of pollen from buttercup plants (Ranunculus) recently was shown to have a deleterious effect on developing larvae of several bee species not specialized on Ranunculus. Numerous Ranunculus species contain ranunculin, the glucosyl hydrate form of the highly reactive and toxic lactone protoanemonin, that causes the toxicity of these plants. We tested whether the presence of ranunculin is responsible for the lethal effects of R. acris pollen on the larvae of two bee species that are not Ranunculus specialists. To investigate the effect on bee larval development, we added ranunculin to the pollen provisions of the Campanula specialist bee Chelostoma rapunculi and the Asteraceae specialist bee Heriades truncorum, and allowed the larvae to feed on these provisions. We quantified ranunculin in pollen of R. acris and in brood cell provisions collected by the Ranunculus specialist bee Chelostoma florisomne. We demonstrated that although ranunculin was lethal to both tested bee species in high concentrations, the concentration in the pollen of R. acris was at least fourfold lower than that tolerated by the larvae of C. rapunculi and H. truncorum in the feeding experiments. Ranunculin concentration in the brood cells of C. florisomne was on average even twentyfold lower than that in Ranunculus pollen, suggesting that a mechanism different from ranunculin intoxication accounts for the larval mortality reported for bees not specialized on Ranunculus pollen.

Detection of IgM and IgG against hepatitis E virus in serum and meat juice samples from pigs at slaughter in Bavaria, Germany.

Hepatitis E virus (HEV) is an emerging foodborne pathogen with domestic and wild pigs (and likely other species such as deer or rabbits) recognized as reservoir. Pathogenesis in pigs usually leads to an asymptomatic course of disease. Since there is no enzyme-linked immunosorbent assay (ELISA) kit for the detection of anti-HEV antibodies in pigs commercially available, the objective of this study was to assess the seroprevalence in fattening pigs at slaughter and at herd level using a newly developed ELISA based on genotype (GT) 1 and GT 3 in Bavaria, Germany. Based on 516 serum and 198 meat juice samples collected from different herds at four different Bavarian slaughterhouses, the overall seroprevalence of anti-HEV IgG in serum and meat juice samples was 68.6% and 67.6%, respectively. Analyzing the serum for the presence of anti-HEV IgM, 36/516 (7%) were positive for anti-HEV IgM. At herd level, most of the herds were seropositive for anti-HEV antibodies. The present study shows that HEV is widespread among the Bavarian pig population and that some pigs might test positive for anti-HEV IgM even at the age of slaughter. Also, meat juice serves as an equivalent matrix to serum to test for anti-HEV antibodies in pigs.

Associational resistance and associational susceptibility: specialist herbivores show contrasting responses to tree stand diversification.

Heterospecific neighbors may reduce damage to a focal plant by lowering specialist herbivore loads (associational resistance hypothesis), or enhance damage by increasing generalist herbivore loads (associational susceptibility hypothesis). We tested the associational effects of tree diversity on herbivory patterns of the tropical focal tree Tabebuia rosea in an experimental plantation setup, which contained tree monocultures and mixed stands. We found higher herbivore damage to T. rosea at higher tree diversity, indicating that T. rosea did not benefit from associational resistance but rather experienced associational susceptibility. The specific consideration of the two dominant insect herbivore species of T. rosea, the specialist chrysomelid Walterianella inscripta and the specialist pyralid Eulepte gastralis, facilitated understanding of the detected damage patterns. Tree diversity exerted opposite effects on tree infestation by the two herbivores. These findings point to resource concentration effects for the chrysomelid beetle (favored by tree monoculture) and to resource dilution effects for the pyralid caterpillar (favored by tree mixture) as underlying mechanisms of herbivore distribution. A strong contribution of the pyralid to overall damage patterns in diversified stands suggests that associational susceptibility may not necessarily be related to higher abundances of generalist herbivores but may also result from specialized herbivores affected by resource dilution effects. Thus, the identity and biology of herbivore species has to be taken into account when attempting to predict damage patterns in forest ecosystems.

Seroprevalence study in forestry workers from eastern Germany using novel genotype 3- and rat hepatitis E virus-specific immunoglobulin G ELISAs.

Hepatitis E virus (HEV) is the causative agent of an acute self-limiting hepatitis in humans. In industrialized countries, autochthonous cases are linked to zoonotic transmission from domestic pigs, wild boar and red deer. The main route of human infection presumably is consumption of contaminated meat. Farmers, slaughterers and veterinarians are expected to be risk groups as they work close to potentially infected animals. In this study, we tested four Escherichia coli-expressed segments of the capsid protein (CP) of a German wild boar-derived HEV genotype 3 strain for their diagnostic value in an indirect immunoglobulin G (IgG) ELISA. In an initial validation experiment, a carboxy-terminal CP segment spanning amino acid (aa) residues 326-608 outperformed the other segments harbouring aa residues 112-608, 326-660 and 112-335. Based on this segment, an indirect ELISA for detection of anti-HEV IgG antibodies in human sera was established and validated using a commercial line immunoassay as reference assay. A total of 563 sera from forestry workers of all forestry offices of Brandenburg, eastern Germany and 301 sera of blood donors from eastern Germany were surveyed using these assays. The commercial test revealed seroprevalence rates of 11% for blood donors and 18% for forestry workers. These rates are in line with data obtained by the in-house test (12 and 21%). Hence, the in-house test performed strikingly similar to the commercial test (sensitivity 0.9318, specificity 0.9542). An initial screening of forestry worker and blood donor sera with a corresponding CP segment of the recently discovered Norway rat-associated HEV revealed several strong positive sera exclusively in the forestry worker panel. Future investigations have to prove the performance of this novel IgG ELISA in large-scale seroepidemiological studies. In addition, the observed elevated seroprevalence in a forestry worker group has to be confirmed by studies on groups of forestry workers from other regions. The epidemiological role of ratHEV in human disease should be assessed in a large-scale study of risk and non-risk groups.

Ability of the oriental fruit moth Grapholita molesta (Lepidoptera: Tortricidae) to detoxify juglone, the main secondary metabolite of the non-host plant walnut.

Many plant species produce toxic secondary metabolites that limit attacks by herbivorous insects, and may thereby constrain insect expansion to new hosts. Walnut is a host for the codling moth Cydia pomonella, which efficiently detoxifies the main walnut defensive compound juglone (5-hydroxy-1,4-naphthoquinone). The oriental fruit moth Grapholita molesta, which also belongs to the tribe Grapholitini, does not feed on walnut. We tested the performance of G. molesta, a highly invasive species, on artificial diets containing juglone at levels mimicking those found in walnut over the growing season. Juglone-fed G. molesta survived relatively well to adulthood, but larval and adult body weights were reduced, and larval developmental time was prolonged in a dose-dependent fashion. Chemical analysis of frass from larvae that had been fed a juglone-containing diet suggests that G. molesta reduces juglone to non-toxic 1,4,5-trihydroxynaphthalene in its gut. This unexpected tolerance of G. molesta to high levels of juglone may facilitate expansion of the host range beyond the current rosacean fruit trees used by this invasive pest.

Can pollen headspace volatiles and pollenkitt lipids serve as reliable chemical cues for bee pollinators?

Chemical analysis of putative contact chemical cues for pollinators from pollen of two plant species, Ranunculus bulbosus (Ranunculaceae) and Campanula rapunculoides (Campanulaceae), showed high consistency in the qualitative and quantitative composition of pollenkitt surface lipids in all samples analyzed per species. The pollenkitt lipids of R. bulbosus included an aldehyde, fatty acid amides, saturated and unsaturated hydrocarbons, and secondary alcohols; the lipids of C. rapunculoides consisted of an aldehyde, monoketones, and ß-diketones. In marked contrast, the pollen headspace volatiles showed a wide qualitative and quantitative variability among all samples per species, whereby the variability was more pronounced in R. bulbosus. Hence, the highly species-specific pollenkitt lipids may provide pollinators with more reliable information on pollen identity.

How the oligophage codling moth Cydia pomonella survives on walnut despite its secondary metabolite juglone.

Besides apple, its primary host, the codling moth Cydia pomonella uses walnut as a secondary host. Abundance of toxic naphthoquinones, among which juglone prevails, does not restrain this economically important pest insect from infesting walnut, but processes underlying the suitability of this host were yet unknown. Larvae feeding on an artificial diet supplemented with juglone at naturally occurring concentrations survived to adulthood at a similarly high proportion as those in the juglone-devoid control. However, their development time was prolonged, their weight gain was reduced, and adult sex ratio was distorted. Results from the natural system with walnut and apple fruits were in line with data gained on artificial diet. Remarkably, a twofold increase of the maximal juglone content reported from the walnut husk was lethal to the larvae. Chemical analyses showed that larvae feeding on the artificial diet supplemented with juglone concentrations present in walnut contained 1,4,5-trihydroxynaphthalene and excreted it in their frass, whereas the hemolymph contained neither detectable amounts of juglone nor the product of its reduction. Hence, effective metabolism of juglone in the intestinal system of the larvae underlies their survival on host plants containing this defensive compound.

Within-plant distribution of induced resistance in apple seedlings: rapid acropetal and delayed basipetal responses.

Induction of plant resistance by herbivory is a complex process, which follows a temporal dynamic and varies spatially at the within-plant scale. This study aimed at improving the understanding of the induction process in terms of time scale and within-plant allocation, using apple tree seedlings (Malus × domestica) as plant model. Feeding preferences of a leaf-chewing insect (Spodoptera littoralis) for previously damaged and undamaged plants were assessed for six different time intervals with respect to the herbivore damage treatment and for three leaf positions. In addition, main secondary defense compounds were quantified and linked to herbivore feeding preferences. Significant herbivore preference for undamaged plants (induced resistance) was first observed 3 days after herbivore damage in the most apical leaf. Responses were delayed in the other leaf positions, and induced resistance decreased within 10 days after herbivore damage simultaneously in all tested leaf positions. Chemical analysis revealed higher concentrations of the flavonoid phloridzin in damaged plants as compared to undamaged plants. This indicates that herbivore preference for undamaged apple plants may be linked to phloridzin, which is the main secondary metabolite of apple leaves. The observed time course and distribution of resistance responses within plants contribute to the understanding of induction processes and patterns, and support the optimal defense theory stating young tissue to be prioritized. Moreover, induced resistance responses occurred also basipetally in leaves below the damage site, which suggests that signaling pathways involved in resistance responses are not unidirectional.

No evidence for increased extinction proneness with decreasing effective population size in a parasitoid with complementary sex determination and fertile diploid males.

BACKGROUND: In species with single locus complementary sex determination (sl-CSD), the sex of individuals depends on their genotype at one single locus with multiple alleles. Haploid individuals are always males. Diploid individuals are females when heterozygous, but males when homozygous at the sex-determining locus. Diploid males are typically unviable or effectively sterile, hence imposing a genetic load on populations. Diploid males are produced from matings of partners that share an allele at the sex-determining locus. The lower the allelic diversity at the sex-determining locus, the more diploid males are produced, ultimately impairing the growth of populations and jeopardizing their persistence. The gregarious endoparasitoid wasp Cotesia glomerata is one of only two known species with sl-CSD and fertile diploid males. RESULTS: By manipulating the relatedness of the founders, we established replicated experimental populations of the parasitoid C. glomerata differing in their genetic effective size, and thus in allelic richness at the sex-determining locus and in the expected magnitude of diploid male production. Our long-term survey of population welfare and persistence did not provide evidence for increased proneness to population extinction with decreasing initial genetic effective population size. Most recorded surrogates of fitness nevertheless decayed over time and most experimental populations eventually went extinct, suggesting that the negative effects of inbreeding outweighed any premium from the fertility of diploid males. CONCLUSIONS: The fertility of diploid males may have evolved as an adaptation prompted by the risk of extinction looming over small isolated populations of species with sl-CSD. However, fertility of diploid males does not negate the costs imposed by their production, and although it may temporarily stave off extinction, it is not sufficient to eradicate the negative effects of inbreeding.

Early-season headspace volatiles from apple and their effect on the apple blossom weevil Anthonomus pomorum.

Apple volatiles emitted at early phenological stages are little investigated, although they may influence behavior of early-season pests. The apple blossom weevil Anthonomus pomorum is a herbivore pest of orchards in Europe. It colonizes apple trees in early season and oviposits into developing flower buds, often leading to economic damage. Using in situ radial diffusive sampling and thermal desorption, followed by GC/MS analysis, headspace volatiles from apple twigs with flower buds at three early phenological tree stages were identified and quantified. The volatile blend consisted of 13 compounds for the first, and increased to 15 compounds for the third phenological stage sampled. These blends included benzenoids, terpenes, and derivatives of fatty acids. A recombined synthetic blend served as the odor source in a still-air dual-choice olfactometer bioassay, in which individual male and female weevils were tested. Results from this behavioral test document an attraction of both sexes to odors of their host plant, suggesting that apple volatiles emitted in early season serve as olfactory cues for host location of A. pomorum in the field.

Structured populations of the oriental fruit moth in an agricultural ecosystem.

Intercontinental trade has led to multiple introductions of invasive pest species at a global scale. Molecular analyses of the structure of populations support the understanding of ecological strategies and evolutionary patterns that promote successful biological invasions. The oriental fruit moth, Grapholita (= Cydia) molesta, is a cosmopolitan and economically destructive pest of stone and pome fruits, expanding its distribution range concomitantly with global climate warming. We used ten newly developed polymorphic microsatellite markers to examine the genetic structure of G. molesta populations in an agricultural ecosystem in the Emilia-Romagna region of northern Italy. Larvae collected in eight sampling sites were assigned to a mosaic of five populations with significant intra-regional structure. Inferred measures of gene flow within populations implicated both active dispersal, and passive dispersal associated with accidental anthropogenic displacements. Small effective population sizes, coupled with high inbreeding levels, highlighted the effect of orchard management practices on the observed patterns of genetic variation within the sampling sites. Isolation by distance did not appear to play a major role at the spatial scale considered. Our results provide new insights into the population genetics and dynamics of an invasive pest species at a regional scale.

Relationship between behavior and physiology in an invasive pest species: oviposition site selection and temperature-dependent development of the oriental fruit moth (Lepidoptera: Tortricidae).

Oviposition site selection is crucial for the reproductive success of a herbivore insect species with relatively sedentary larvae. The optimal oviposition theory, i.e., the preference-performance hypothesis, has thus far mainly been tested with a focus on nutritional quality of the host. This study investigates whether female oriental fruit moth Grapholita (Cydia) molesta choose a microhabitat for oviposition characterized by a temperature range within which their offspring perform best. Thermal preferences of females during oviposition were assessed in a circular temperature gradient arena. Offspring performance and survival were assessed under different constant temperature conditions. Females preferred oviposition sites of approximately 30 degrees C over lower and higher temperatures. At this temperature, egg, larval, and pupal development was significantly faster than at 22 and 25 degrees C, and larval development was also faster than at 33 degrees C. At 30 degrees C and at the lower temperatures tested, survival of eggs and larvae was significantly higher than at 33 degrees C, whereas development was precluded at 35 degrees C. Furthermore, female pupal weight attained at 30 and 33 degrees C exceeded that reached at the lower temperatures tested. Considering the potentially reduced predation risk caused by the shorter developmental time of eggs and larvae, the laboratory data suggest that this species maximizes its fitness by selecting a thermally optimal environment for its offspring, supporting the optimal oviposition theory. Conversely, it is known that the codling moth (C. pomonella) lacks a mechanism to avoid temperatures lethal to progeny development, which may reflect the differences in geographic ranges of these tortricids.

Long foraging distances impose high costs on offspring production in solitary bees.

1. Solitary bees are central place foragers returning to their nests several times a day with pollen and nectar to provision their brood cells. They are especially susceptible to landscape changes that lead to an increased spatial separation of suitable nesting sites and flower rich host plant stands. While knowledge of bee foraging ranges is currently growing, quantitative data on the costs of foraging flights are very scarce, although such data are crucial to understand bee population dynamics. 2. In this study, the impact of increased foraging distance on the duration of foraging bouts and on the number of brood cells provisioned per time unit was experimentally quantified in the two pollen specialist solitary bee species Hoplitis adunca and Chelostoma rapunculi. Females nesting at different sites foraged under the same environmental conditions on a single large and movable flowering host plant patch in an otherwise host plant free landscape. 3. The number of brood cells provisioned per time unit by H. adunca was found to decrease by 23%, 31% and 26% with an increase in the foraging distance by 150, 200 and 300 m, respectively. The number of brood cells provisioned by C. rapunculi decreased by 46% and 36% with an increase in the foraging distance by 500 and 600 m, respectively. 4. Contrary to expectation, a widely scattered arrangement of host plants did not result in longer mean duration of a foraging bout in H. adunca compared to a highly aggregated arrangement, which might be due to a reduced flight directionality combined with a high rate of revisitation of already depleted flowers in the aggregated plant arrangement or by a stronger competition and disturbance by other flower visitors. 5. The results of this study clearly indicate that a close neighbourhood of suitable nesting and foraging habitats is crucial for population persistence and thus conservation of endangered solitary bee species.