The snail Smaragdia bryanae (Neritopsina,Neritidae) is a specialist herbivore of the seagrass Halophila hawaiiana (Alismatidae,Hydrocharitaceae)

Abstract. The endemic Hawaiian gastropod Smaragdia bryanae is a specialized marine herbivore that uses the endemic seagrass Halophila hawaiiana as both food and habitat. These small neritids, their grazing scars, and their egg capsules are found year‐round on seagrass leaves, where they feed on protoplast contents released as the sharp outer‐lateral teeth of the snail's radula puncture leaf epidermal cells; the contents of these cells are likely swept into the mouth by the long, wispy cusps of the marginal teeth. Structural differences from the typical neritid radula include elongated outer‐lateral teeth with two sharply pointed cusps, delicate marginal teeth reduced in both size and number, and a compressed central section. Snails grazed on leaves of H. hawaiiana steadily in laboratory culture, and grew and reproduced on this diet. In laboratory choice experiments, snails did not graze the thalli of any of six macroalgal species growing near seagrass where snails were collected, and strongly preferred occupying seagrass. Seagrass samples from five field sites on Oahu and one on Maui showed from 30% to 94% of leaves damaged, with 11% of the total leaf standing area grazed. Snails are smaller (mean length 2.74±0.32 mm, mean width 2.15±0.17 mm, n=217) than the width of the leaves of H. hawaiiana (mean 3.24±1.26 mm, n=790). The snails associate constantly with their host, despite the scattered distribution, small patch size, and variability of the seagrass resource, demonstrated by a sevenfold range in the leaf area index (mean 1.11±0.61 cm² blade surface cm^?2, n=31) among samples. Damage on grazed leaves (mean 8.21±7.05 mm² per leaf, or 16.5% of leaf surface, n=511) is concentrated in the apical and central epithelia between the midrib and the marginal veins, where snails may access cells with thinner walls and few fibers. Details of the grazing interaction between these extant species in Hawai'i shed light on the ecological specialization of members of the genus Smaragdia to seagrasses over geological time.     

Responses of pre‐dispersal seed predators to sequential flowering of Dipterocarps in Malaysia

Many species of Dipterocarpaceae and other plant families reproduce synchronously at irregular, multi‐year intervals in Southeast Asian forests. These community‐wide general flowering events are thought to facilitate seed survival through satiation of generalist seed predators. During a general flowering event, closely related Shorea species (Dipterocarpaceae) stagger their flowering times by several weeks, which may minimize cross pollination and interspecific competition for pollinators. Generalist, pre‐dispersal seed predators might also track flowering hosts and influence predator satiation. We addressed the question of whether pre‐dispersal seed predation differed between early and late flowering Shorea species by monitoring flowering, fruiting and seed predation intensity over two general flowering events at the Pasoh Research Forest, Malaysia. Pre‐dispersal insect seed predators killed up to 63 percent of developing seeds, with Nanophyes shoreae, a weevil that feeds on immature seeds being the most important predator for all Shorea species. This weevil caused significantly greater pre‐dispersal seed predation in earlier flowering species. Long larval development time precluded oviposition by adults that emerged from the earliest flowering Shorea on the final flowering Shorea. In contrast, larvae of weevils that feed on mature seeds before seed dispersal (Alcidodes spp.), appeared in seeds of all Shorea species almost simultaneously. We conclude that general flowering events have the potential to satiate post‐dispersal seed predators and pre‐dispersal seed predators of mature fruit, but are less effective at satiating pre‐dispersal predators of immature fruit attacking early flowering species.     

Host plant genetic control of associated fungal and insect species in a Populus hybrid cross

Plants employ a diverse set of defense mechanisms to mediate interactions with insects and fungi. These relationships can leave lasting impacts on host plant genome structure such as rapid expansion of gene families through tandem duplication. These genomic signatures provide important clues about the complexities of plant/biotic stress interactions and evolution. We used a pseudo‐backcross hybrid family to identify quantitative trait loci (QTL) controlling associations between Populus trees and several common Populus diseases and insects. Using whole‐genome sequences from each parent, we identified candidate genes that may mediate these interactions. Candidates were partially validated using mass spectrometry to identify corresponding QTL for defensive compounds. We detected significant QTL for two interacting fungal pathogens and three insects. The QTL intervals contained candidate genes potentially involved in physical and chemical mechanisms of host–plant resistance and susceptibility. In particular, we identified adjoining QTLs for a phenolic glycoside and Phyllocolpa sawfly abundance. There was also significant enrichment of recent tandem duplications in the genomic intervals of the native parent, but not the exotic parent. Tandem gene duplication may be an important mechanism for rapid response to biotic stressors, enabling trees with long juvenile periods to reach maturity despite many coevolving biotic stressors.     

Fine‐scale frequency differentiation along a herbivory gradient in the trichome dimorphism of a wild Arabidopsis

Geographic variation is commonly observed in plant resistance traits, where plant species might experience different selection pressure across a heterogeneous landscape. Arabidopsis halleri subsp. gemmifera is dimorphic for trichome production, generating two morphs, trichome‐producing (hairy) and trichomeless (glabrous) plants. Trichomes of A. halleri are known to confer resistance against the white butterfly, cabbage sawfly, and brassica leaf beetle, but not against flea beetles. We combined leaf damage, microclimate, and microsatellite loci data of 26 A. halleri populations in central Japan, to explore factors responsible for fine‐scale geographic variation in the morph frequency. We found that hairy plants were less damaged than glabrous plants within populations, but the among‐site variation was the most significant source of variation in the individual‐level damage. Fixation index () of a putative trichome locus exhibited a significant divergence along population‐level damage with an exception of an outlier population, inferring the local adaptation to herbivory. Notably, this outlier was a population wherein our previous study reported a balancing role of the brassica leaf beetle Phaedon brassicae on the morph frequency. This differentiation of the trichome locus was unrelated to neutral genetic differentiation (evaluated by of microsatellite loci) and meteorological factors (including temperature and solar radiation). The present findings, combined with those of our previous work, provide suggestive evidence that herbivore‐driven divergence and occasional outbreak of a specific herbivore have jointly contributed to the ecogeographic pattern in the frequency of two morphs.     

Fusarium oxysporum mediates systems metabolic reprogramming of chickpea roots as revealed by a combination of proteomics and metabolomics

Molecular changes elicited by plants in response to fungal attack and how this affects plant–pathogen interaction, including susceptibility or resistance, remain elusive. We studied the dynamics in root metabolism during compatible and incompatible interactions between chickpea and Fusarium oxysporum f. sp. ciceri (Foc), using quantitative label‐free proteomics and NMR‐based metabolomics. Results demonstrated differential expression of proteins and metabolites upon Foc inoculations in the resistant plants compared with the susceptible ones. Additionally, expression analysis of candidate genes supported the proteomic and metabolic variations in the chickpea roots upon Foc inoculation. In particular, we found that the resistant plants revealed significant increase in the carbon and nitrogen metabolism; generation of reactive oxygen species (ROS), lignification and phytoalexins. The levels of some of the pathogenesis‐related proteins were significantly higher upon Foc inoculation in the resistant plant. Interestingly, results also exhibited the crucial role of altered Yang cycle, which contributed in different methylation reactions and unfolded protein response in the chickpea roots against Foc. Overall, the observed modulations in the metabolic flux as outcome of several orchestrated molecular events are determinant of plant's role in chickpea–Foc interactions.     

Shrub facilitation drives tree establishment in a semiarid fog‐dependent ecosystem

Questions

The exceptional occurrence of tall rain forest patches on foggy coastal mountaintops, surrounded by extensive xerophytic shrublands, suggests an important role of plant–plant interactions in the origin and persistence of these patches in semi‐arid Chile. We asked whether facilitation by shrubs can explain the growth and survival of rain forest tree species, and whether shrub effects depend on the identity of the shrub species itself, the drought tolerance of the tree species and the position of shrubs in regard to wind direction.

Location

Open area–shrubland–forest matrix, Fray Jorge Forest National Park, Chile.

Methods

We recorded survival after 12 years of a ~3600 tree saplings plantation (originally ~30‐cm tall individuals) of Aextoxicon punctatum, Myrceugenia correifolia and Drimys winteri placed outside forests, beneath the shrub Baccharis vernalis, and in open (shrub‐free) areas. We assessed the effects of neighbouring shrubs and soil humidity on survival and growth along a gradient related to the direction of fog movement.

Results

B. vernalis had a clear facilitative effect on tree establishment and survival since, after ~12 years, saplings only survived beneath the shrub canopy. Long‐term survival strongly depended on tree species identity, drought tolerance and position along the soil moisture gradient, with higher survival of A. punctatum (>35%) and M. correifolia (>14%) at sites on wind‐ and fog‐exposed shrubland areas. Sites occupied by the shrub Aristeguietia salvia were unsuitable for trees, presumably due to drier conditions than under B. vernalis.

Conclusions

Interactions between shrubs and fog‐dependent tree species in dry areas revealed a strong, long‐lasting facilitation effect on planted tree's survival and growth. Shrubs acted as benefactors, providing sites suitable for tree growth. Sapling mortality in the shrubland interior was caused by lower soil moisture, the consequence of lower fog loads in the air and thus insufficient facilitation. While B. vernalis was a key ecosystem engineer (nurse) and intercepted fog water that dripped to trees planted underneath, drier sites with A. salvia were unsuitable for trees. Consequently, nurse effects related to water input are strongly site and species specific, with facilitation by shrubs providing a plausible explanation for the initiation of forest patches in this semi‐arid landscape.     

Plant–plant interactions as a mechanism structuring plant diversity in a Mediterranean semi‐arid ecosystem

Plant–plant interactions are among the fundamental processes that shape structure and functioning of arid and semi‐arid plant communities. Despite the large amount of studies that have assessed the relationship between plant–plant interactions (i.e., facilitation and competition) and diversity, often researchers forget a third kind of interaction, known as allelopathy. We examined the effect of plant–plant interactions of three dominant species: the perennial grass Lygeum spartum, the allelopathic dwarf shrub Artemisia herba‐alba, and the nurse shrub Salsola vermiculata, on plant diversity and species composition in a semi‐arid ecosystem in NE Spain. Specifically, we quantified the interaction outcome (IO) based on species co‐occurrence, we analyzed diversity by calculation of the individual species–area relationship (ISAR), and compositional changes by calculation of the Chao‐Jaccard similarity index. We found that S. vermiculata had more positive IO values than L. spartum, and A. herba‐alba had values between them. Lygeum spartum and A. herba‐alba acted as diversity repellers, whereas S. vermiculata acted as a diversity accumulator. As aridity increased, A. herba‐alba transitioned from diversity repeller to neutral and S. vermiculata transitioned from neutral to diversity accumulator, while L. spartum remained as diversity repeller. Artemisia herba‐alba had more perennial grass species in its local neighborhood than expected by the null model, suggesting some tolerance of this group to its “chemical neighbor”. Consequently, species that coexist with A. herba‐alba were very similar among different A. herba‐alba individuals. Our findings highlight the role of the nurse shrub S. vermiculata as ecosystem engineer, creating and maintaining patches of diversity, as well as the complex mechanism that an allelopathic plant may have on diversity and species assemblage. Further research is needed to determine the relative importance of allelopathy and competition in the overall interference of allelopathic plants.     

El Niño,Host Plant Growth,and Migratory Butterfly Abundance in a Changing Climate

In the wet forests of Panama, El Niño typically brings a more prolonged and severe dry season. Interestingly, many trees and lianas that comprise the wet forests increase their productivity as a response to El Niño. Here, we quantify the abundance of migrating Marpesia chiron butterflies over 17 yr and the production of new leaves of their hostplants over 9 yr to test the generality of the El Niño migration syndrome, i.e., whether increased abundance of migrating insects and productivity of their food plants are associated with El Niño and La Niña events. We find that the quantity of M. chiron migrating across the Panama Canal was directly proportional to the sea surface temperature (SST) anomaly of the Pacific Ocean, which characterizes El Niño and La Niña events. We also find that production of new leaves by its larval host trees, namely Brosimum alicastrum, Artocarpus altilis, and Ficus citrifolia, was directly proportional to the SST anomaly, with greater leaf flushing occurring during the period of the annual butterfly migration that followed an El Niño event. Combining these and our previously published results for the migratory butterfly Aphrissa statira and its host lianas, we conclude that dry season rainfall and photosynthetically active radiation can serve as primary drivers of larval food production and insect population outbreaks in Neotropical wet forests, with drier years resulting in enhanced plant productivity and herbivore abundance. Insect populations should closely track changes in both frequency and amplitude of the El Niño Southern Oscillation with climate change.     

Neofusicoccum parvum and Diaporthe foeniculina associated with twig and shoot blight and branch canker of citrus in Greece

In a survey performed in Chania and Aetoloacarnania, Greece in years 2013–2014, fungal isolates causing twig and shoot blight and branch canker of citrus trees were morphologically characterized and identified by multiple gene sequence analysis. By sequencing the ITS‐5.8S rRNA, the elongation factor 1‐α (EF1‐α), the β‐tubulin and the RNA polymerase II subunit (Rpb2) genes, the isolates examined were associated with Diaporthe foeniculina (six isolates) and Neofusicoccum parvum (one isolate). All six D. foeniculina isolates showed slow colony growth rates (7.4 ± 3.2 mm/day), while the N. parvum isolate exhibited fast growth (41.6 mm/day). Koch's criteria were met after re‐isolation of D. foeniculina isolates from all inoculated Citrus spp. and N. parvum from inoculated C. reticulata “Ortanique” and after having developed symptoms similar to those detected on shoots and branches collected from citrus fields. Based on lesion length on detached C. medica “Lia Kritis” shoots, N. parvum caused long necrotic lesions (58 mm in length) in comparison with a length of 12–21 mm lesions caused by D. foeniculina isolates. Pathogenicity trials on nine Citrus spp., which had been inoculated with D. foeniculina and N. parvum, revealed different levels of susceptibility, indicating a host‐dependent infection effect, with Poncirus trifoliate × C. paradisi (“Citrumelo Swingle”) being the most resistant citrus genotype. Lack of host specificity suggests that their pathogen–host association could be attributed to ecological rather to co‐evolutionary factors. This work represents the first report, accompanied with pathogenicity tests, on botryosphaeriaceous and diaporthaceous pathogens associated with twig and shoot blight and branch canker of citrus in Greece.     

Interactions between a pollinating seed predator and its host plant: the role of environmental context within a population

Plant–insect interactions often are important for plant reproduction, but the outcome of these interactions may vary with environmental context. Pollinating seed predators have positive and negative effects on host plant reproduction, and the interaction outcome is predicted to vary with density or abundance of the partners. We studied the interaction between Silene stellata, an herbaceous perennial, and Hadena ectypa, its specialized pollinating seed predator. Silene stellata is only facultatively dependent upon H. ectypa for pollination because other nocturnal moth co‐pollinators are equally effective at pollen transfer. We hypothesized that for plants without conspecific neighbors, H. ectypa would have higher visitation rates compared to co‐pollinators, and the plants would experience lower levels of H. ectypa pollen deposition. We predicted similar oviposition throughout the study site but greater H. ectypa predation in the area without conspecific neighbors compared to plants embedded in a naturally high density area. We found that H. ectypa had consistently higher visitation than moth co‐pollinators in all host plant contexts. However, H. ectypa pollinator importance declined in areas with low conspecific density because of reduced pollen deposition, resulting in lower seed set. Conversely, oviposition was similar across the study site independent of host plant density. Greater likelihood of very high fruit predation combined with lower pollination by H. ectypa resulted in reduced S. stellata female reproductive success in areas with low conspecific density. Our results demonstrate local context dependency of the outcomes of pollinating seed predator interactions with conspecific host plant density within a population.     

Lack of evolution in a leaf beetle that lives on two contrasting host plants

The interactions between plant‐eating insects and their hosts have shaped both the insects and the plants, driving evolution of plant defenses and insect specialization. The leaf beetle Trirhabda eriodictyonis (Chrysomelidae) lives on two shrubs with differing defenses: Eriodictyon crassifolium has hairy leaves, whereas E. trichocalyx has resinous leaves. We tested whether these beetles have differentiated onto the two host plants, and if not, whether the beetles prefer the better host plant and prefer mates who are from that host plant. In feeding tests, adult beetles strongly preferred eating E. trichocalyx regardless of which host they came from. In addition, females laid more eggs if they ate E. trichocalyx than E. crassifolium. So, E. trichocalyx is generally the better host. However, beetle mate preference was not in line with food choice. Males did not prefer to mate with females from E. trichocalyx. Females from E. crassifolium did prefer males from E. trichocalyx over males from E. crassifolium, but did not lay more eggs as a result of these matings. We conclude that the beetle populations we studied have not differentiated based on their host plants and may not have even adapted to the better host. Although to humans these host plant defenses differ dramatically, signs that they have caused evolution in the beetles are lacking. The case of T. eriodictyonis stands counter to many other studies that have seen the differentiation of ecotypes and/or adaptive coordination of an herbivore's life cycle based on host plant differences.     

Tracking changes in life‐history traits related to unnecessary virulence in a plant‐parasitic nematode

Evaluating trade‐offs in life‐history traits of plant pathogens is essential to understand the evolution and epidemiology of diseases. In particular, virulence costs when the corresponding host resistance gene is lacking play a major role in the adaptive biology of pathogens and contribute to the maintenance of their genetic diversity. Here, we investigated whether life‐history traits directly linked to the establishment of plant–nematode interactions, that is, ability to locate and move toward the roots of the host plant, and to invade roots and develop into mature females, are affected in Meloidogyne incognita lines virulent against the tomato Mi‐1.2 resistance gene. Virulent and avirulent near‐isogenic lines only differing in their capacity to reproduce or not on resistant tomatoes were compared in single inoculation or pairwise competition experiments. Data highlighted (1) a global lack of trade‐off in traits associated with unnecessary virulence with respect to the nematode ability to successfully infest plant roots and (2) variability in these traits when the genetic background of the nematode is considered irrespective of its (a)virulence status. These data suggest that the variation detected here is independent from the adaptation of M. incognita to host resistance, but rather reflects some genetic polymorphism in this asexual organism.     

Persistence of flower visitors and pollination services of a generalist tree in modified forests

Flower‐visiting insects provide essential pollination services to many plant species. It is thus of critical importance to understand the effects of anthropogenic landscape modification on these animals. Particularly at the landscape scale, we still lack information on how flower visitors are affected by different intensities of human disturbance. In this study, we chose six representative types of forest modification across a heterogeneous South African landscape. At 36 study sites we observed insect visitation to Celtis africana flowers in two consecutive years. This generalist tree species has small unspecialized flowers which we found to be pollinated by a diverse array of insects as well as by wind. Visitation rates to flowers of C. africana differed significantly among the six forest types and between two study years. Visitation rates were enhanced in modified forests, facilitated by a high abundance of feral honeybees (Apis mellifera). Fruit set in C. africana showed significant positive associations with insect visitation and with the diversity of flower visitors, but was only weakly predicted by forest type. Our findings imply that even though forest modification can strongly alter flower visitors, pollination services for trees with unspecialized flowers may persist at a landscape scale. We advise conservation managers to maintain modified forest fragments in addition to natural forests as these may contribute to sustain pollination services in human‐modified landscapes.     

Effects of gene flow on phenotype matching between two varieties of Joshua tree (Yucca brevifolia; Agavaceae) and their pollinators

In animal‐pollinated plants, local adaptation to pollinator behaviour or morphology can restrict gene flow among plant populations; but gene flow may also prevent divergent adaptation. Here, we examine possible effects of gene flow on plant–pollinator trait matching in two varieties of Joshua tree (Agavaceae: Yucca brevifolia). The two varieties differ in strikingly in floral morphology, which matches differences in the morphology of their pollinators. However, this codivergence is not present at a smaller scale: within the two varieties of Joshua tree, variation in floral morphology between demes is not correlated with differences in moth morphology. We use population genetic data for Joshua tree and its pollinators to test the hypotheses that gene flow between Joshua tree populations is structured by pollinator specificity, and that gene flow within the divergent plant–pollinator associations ‘swamps’ fine‐scale coadaptation. Our data show that Joshua tree populations are structured by pollinator association, but the two tree varieties are only weakly isolated – meaning that their phenotypic differences are maintained in the face of significant gene flow. Coalescent analysis of gene flow between the two Joshua tree types suggests that it may be shaped by asymmetric pollinator specificity, which has been observed in a narrow zone of sympatry. Finally, we find evidence suggesting that gene flow among Joshua tree sites may shape floral morphology within one plant–pollinator association, but not the other.     

Spatial variation in the demography and population dynamics of a perennial shrub (Atriplex vesicaria) under sheep grazing in semi‐arid Australian rangelands

Atriplex vesicaria Heward ex Benth. (Chenopodiaceae) is a widespread perennial shrub in southern Australia's chenopod rangelands but is sensitive to grazing. A detailed investigation of the demography and population dynamics of A. vesicaria under sheep grazing was conducted over 6.5 years at a range of sites across a typical paddock to assess the long‐term effects of grazing on the species and elucidate the mechanisms of population change under grazing. The effects of rainfall on recruitment and mortality were also examined. Six‐monthly censuses of all A. vesicaria individuals were conducted in permanent grazed and ungrazed plots at sites located across an 1100‐ha paddock. Grazing increased adult shrub mortality close to water and reduced recruitment over a broader area of the paddock, but seedling survival did not appear to be affected by grazing. As a result of these changes, the population declined on grazed plots up to 2200 m from water during the study, but the decline was greatest closer to water. The population was most dynamic at the sites furthest from the water point where it was unaffected by grazing because of the greater recruitment and mortality of young plants, but because these processes balanced out over time, population density was effectively unchanged by the end of the study. Although statistical models indicated that six‐monthly rainfall did not explain temporal variation in recruitment or mortality, rainfall nevertheless has a central role in both processes. In particular, longer periods of favourable rainfall and drought appear to have an important influence on recruitment and mortality, respectively, with heavy grazing during a drought period increasing mortality. Occasional shortages of seed or rains occurring during the warmer months when seed germination is limited possibly explain poor recruitment at sites unaffected by grazing following good rainfall.     

Late‐onset progressive retinal atrophy in the Gordon and Irish Setter breeds is associated with a frameshift mutation in C2orf71

Progressive retinal atrophy (PRA) in dogs is characterised by the degeneration of the photoreceptor cells of the retina, resulting in vision loss and eventually complete blindness. The condition affects more than 100 dog breeds and is known to be genetically heterogeneous between breeds. Around 14 mutations have now been identified that are associated with PRA in around 49 breeds, but for the majority of breeds the mutation(s) responsible have yet to be identified. Using genome‐wide association with 16 Gordon Setter PRA cases and 22 controls, we identified a novel PRA locus, termed rod–cone degeneration 4 (rcd4), on CFA17 (P_raw = 2.22 × 10^?8, P_genome = 2.00 × 10^?5), where a 3.2‐Mb region was homozygous within cases. A frameshift mutation was identified in C2orf71, a gene located within this region. This variant was homozygous in 19 of 21 PRA cases and was at a frequency of approximately 0.37 in the Gordon Setter population. Approximately 10% of cases in our study (2 of 21) are not associated with this C2orf71 mutation, indicating that PRA in this breed is genetically heterogeneous and caused by at least two mutations. This variant is also present in a number of Irish Setter dogs with PRA and has an estimated allele frequency of 0.26 in the breed. The function of C2orf71 remains unknown, but it is important for retinal development and function and has previously been associated with autosomal recessive retinitis pigmentosa in humans.     

Differential susceptibility and suitability of domestic and wild apple species for a florivorous weevil and its parasitoids

Crop plant domestication can change plant resistance to herbivores leading to differences in pest pressure experienced by crop plants and their wild relatives. To compare resistance to herbivores between domesticated and wild fruit trees, we quantified direct resistance and indirect resistance to a pest insect, the florivorous apple blossom weevil Anthonomus pomorum (Coleoptera: Curculionidae), for the cultivated apple Malus domestica and two wild apple species, the European crab apple M. sylvestris and the exotic M. kirghisorum. We measured weevil infestation and performance (weight, sex ratio), and weevil parasitism by parasitoid wasps for different cultivars of M. domestica and for the two wild apple species. To explain weevil and parasitoid responses to different apple species, we quantified tree characteristics including nitrogen content, size of flower buds, bark roughness, tree size, tree phenology and tree position. We found significant differences in susceptibility to weevil infestation between apple species, with lowest infestation (highest apple resistance) in M. domestica and highest infestation in M. kirghisorum. The suitability of apple species also varied significantly: weevils emerging from M. sylvestris were significantly lighter than those from M. kirghisorum. Parasitism of A. pomorum by different parasitoid species was significantly higher in M. sylvestris than in M. domestica. Infestation, weevil weight and parasitism were positively related to tree characteristics: infestation to bud nitrogen content and bark roughness, weevil size to nitrogen content and bud size, and parasitism to tree height and bud density. Our study revealed marked differences between apple species in susceptibility and suitability for the pest herbivore, but also for antagonistic parasitoids. Whereas direct resistance appeared to be higher in cultivated apple, indirect resistance via parasitoids was apparently higher in wild apple trees. Our findings suggest that wild and cultivated apple trees possess different resistance traits that may be combined to optimize resistance in commercial apple cultivars.