Fruit traits of vertebrate-dispersed alien plants: smaller seeds and more pulp sugar than indigenous species

Vertebrates play a major role in dispersing seeds of fleshy-fruited alien plants. However, we know little of how the traits of alien fleshy fruits compare with indigenous fleshy fruits, and how these differences might contribute to invasion success. In this study, we characterised up to 38 fruit morphology, pulp nutrient and phenology traits of an assemblage of 34 vertebrate-dispersed alien species in south-eastern Queensland, Australia. Most alien fruits were small (81% < 15 mm in mean width), and had watery fruit pulps that were high in sugars and low in nitrogen and lipids. When compared to indigenous species, alien fruits had significantly smaller seeds. Further, alien fruit pulps contained more sugar and more variable (and probably greater) nitrogen per pulp wet weight, and species tended to have longer fruiting seasons than indigenous species. Our analyses suggest that fruit traits could be important in determining invasiveness and could be used to improve pre- and post-border weed risk assessment.     

Diets of fruit-eating birds: what are the causes of interspecific differences?

Differences in fruit choice among the bird species of a Spanish shrubland were related to the size of fruits and to the lipid content of pulp. Lipid-rich fruits were selected by the bird species with slower food passage rates through the digestive tract. These bird species also fed frequently on insects and seeds. Bird species with faster food passage rates fed less on insects and seeds, and ate mainly fruits with pulp poor in lipids (rich in sugars and water). Studies of digestion in birds indicate that lipids require slower food passage rates for efficient digestion and intestinal transport than simple sugars. The available evidence indicates that the European bird species that show stronger preferences for lipid-rich fruits are no better as seed dispersers, from the point of view of the plants, than species choosing lipidpoor fruits. Thus, the degree of frugivory of birds, their fruit choice patterns and their effects on seed dispersal do not seem to be related to each other in the ways expected by the early models of the evolution of fleshy fruits.     

The role of fruit traits of bird-dispersed plants in invasiveness and weed risk assessment

Aim  Birds play a major role in the dispersal of seeds of many fleshy-fruited invasive plants. The fruits that birds choose to consume are influenced by fruit traits. However, little is known of how the traits of invasive plant fruits contribute to invasiveness or to their use by frugivores. We aim to gain a greater understanding of these relationships to improve invasive plant management.
Location  South-east Queensland, Australia.
Methods  We measure a variety of fruit morphology, pulp nutrient and phenology traits of a suite of bird-dispersed alien plants. Frugivore richness of these aliens was derived from the literature. Using regressions and multivariate methods, we investigate relationships between fruit traits, frugivore richness and invasiveness.
Results  Plant invasiveness was negatively correlated to fruit size, and all highly invasive species had quite similar fruit morphology [smaller fruits, seeds of intermediate size and few (< 10) seeds per fruit]. Lower pulp water was the only pulp nutrient trait associated with invasiveness. There were strong positive relationships between the diversity of bird frugivores and plant invasiveness, and in the diversity of bird frugivores in the study region and another part of the plants' alien range.
Main conclusions  Our results suggest that weed risk assessments (WRA) and predictions of invasive success for bird-dispersed plants can be improved. Scoring criteria for WRA regarding fruit size would need to be system-specific, depending on the fruit-processing capabilities of local frugivores. Frugivore richness could be quantified in the plant's natural range, its invasive range elsewhere, or predictions made based on functionally similar fruits.     

Characteristics and abundance of vertebrate-dispersed fruits in temperate wet sclerophyll forest in southeastern Australia

The diversity, abundance and display characteristics of plants producing fleshy fruits were recorded in wet sclerophyll forest near Melbourne, Victoria. Fruit characteristics and nutrient content were recorded for 14 species of fruits occurring in the forest and these characteristics compared with similar studies in north temperate and tropical regions. Plants producing fleshy fruits were confined to understorey layers and comprised 13.5% of the native vascular flora. Most species displayed fruits conspicuously by ripening brightly coloured fruits synchronously in large displays. Annual fruit production at three sites ranged from 6.98 to 37.01 kg/ha. Fruit mass, seed number and size were variable but the proportion of pulp per fruit remained remarkably consistent between species. Most fruit pulp contained high quantities of soluble carbohydrates and little lipid or protein. Fruits from wet sclerophyll forest did not differ from fruits from other geographic areas in eight characteristics measured. Although taxonomically distinct, the species in wet sclerophyll forest were not significantly different from other temperate fruits, suggesting that fruit characteristics are under the influence of factors other than phylogeny.     

Seed dispersal anachronisms: rethinking the fruits extinct megafauna ate

Background

Some neotropical, fleshy-fruited plants have fruits structurally similar to paleotropical fruits dispersed by megafauna (mammals >10³ kg), yet these dispersers were extinct in South America 10–15 Kyr BP. Anachronic dispersal systems are best explained by interactions with extinct animals and show impaired dispersal resulting in altered seed dispersal dynamics.

Methodology/Principal Findings

We introduce an operational definition of megafaunal fruits and perform a comparative analysis of 103 Neotropical fruit species fitting this dispersal mode. We define two megafaunal fruit types based on previous analyses of elephant fruits: fruits 4–10 cm in diameter with up to five large seeds, and fruits >10 cm diameter with numerous small seeds. Megafaunal fruits are well represented in unrelated families such as Sapotaceae, Fabaceae, Solanaceae, Apocynaceae, Malvaceae, Caryocaraceae, and Arecaceae and combine an overbuilt design (large fruit mass and size) with either a single or few (<3 seeds) extremely large seeds or many small seeds (usually >100 seeds). Within-family and within-genus contrasts between megafaunal and non-megafaunal groups of species indicate a marked difference in fruit diameter and fruit mass but less so for individual seed mass, with a significant trend for megafaunal fruits to have larger seeds and seediness.

Conclusions/Significance

Megafaunal fruits allow plants to circumvent the trade-off between seed size and dispersal by relying on frugivores able to disperse enormous seed loads over long-distances. Present-day seed dispersal by scatter-hoarding rodents, introduced livestock, runoff, flooding, gravity, and human-mediated dispersal allowed survival of megafauna-dependent fruit species after extinction of the major seed dispersers. Megafauna extinction had several potential consequences, such as a scale shift reducing the seed dispersal distances, increasingly clumped spatial patterns, reduced geographic ranges and limited genetic variation and increased among-population structuring. These effects could be extended to other plant species dispersed by large vertebrates in present-day, defaunated communities.     

Relationships between bird-dispersed plants and avian fruit consumers with different feeding strategies in Japan

We examined the interactions between bird-dispersed plants and fruit-consuming birds with various feeding strategies, by reviewing the plant species consumed by 14 bird species in Japan with four feeding types: gulpers (five species), grinders (four species), crushers (four species), and peckers (one species). Our literature review provided information on the plant species consumed by the birds in Japan and the morphological traits of the fruits: fruit volume, seed mass and number, pulp type (fleshy, dry, or arillate), and plant height (tall, medium, or small). Using these data, we examined the diversity of plant species consumed by each bird and the fruit morphological traits that affected fruit selection. The five gulpers consumed fruits from the largest number of plants, followed by the four grinders, the four crushers, and the one pecker. The gulpers and grinders consumed a wider variety of fruits than were consumed by the crushers and the pecker. Logistic regression analysis revealed that some crushers and the pecker preferred plants with dry or arillate pulp around the seeds. Our results suggest that a frugivorous bird’s feeding strategies, and particularly its fruit-handling behaviors and the fruit parts it ingests, influence the diversity of plants it consumes. The crushers and the pecker, which feed exclusively on seeds, require more effort and time to consume this type of food, and this might cause a strong preference for specific fruit traits and thus, consumption of a lower diversity of plant species.     

Positive relationship between fruit removal by animals and seedling recruitment in a tropical forest

Fleshy-fruited plants rely on animal frugivores to disperse their seeds, and seed removal by frugivores may leave an imprint on seedling recruitment. However, to what extent plant–frugivore interactions are related to seedling recruitment has rarely been quantified at the community level, especially in species-rich tropical forests. In this study, we tested the effect of different plant traits on fruit removal by frugivores and tested the relative importance of fruit removal, plant traits and abiotic factors for seedling recruitment. We quantified plant–frugivore interactions of 22 fleshy-fruited plant species consumed by 56 diurnal frugivore species, and counted the number of seedlings that emerged along an elevational gradient in the Colombian Andes. We measured a set of plant traits (i.e., crop size; fruit size; seed load and mass; fruit nutritional contents), estimated the density of adult plants and recorded relevant abiotic factors (light, temperature and humidity). We found that fruit removal by frugivores was positively associated with crop size, but negatively associated with fruit length and unrelated to seed load and fruit nutritional content. Seedling densities were positively related to the density of adult plants, seed mass and fruit removal by animals. We found no relationship between abiotic factors and seedling recruitment. Our results indicate that fruit abundance and morphology are important determinants of fruit removal and that fruit removal is positively associated with seedling recruitment accounting for effects of species abundance and plant traits. We conclude that plant traits shape fruit removal and seedling recruitment at the community level, while these two crucial processes of forest regeneration are directly linked by seed dispersal of animals.     

Water and fish select for fleshy fruits in tropical wetland forests

Adjacent floodplain and upland tropical forests experience the same temperature and precipitation regimes, but differ substantially in plant species composition and biotic interactions because of extensive flooding. We hypothesize that flooded forests filter fruiting traits linked to seed dispersal by water and fishes, such that selection by water and fish led to (1) trees that synchronize the timing of fruiting with annual floods, and (2) the evolution of fleshy tissues on fruits to improve buoyancy and increase fruit consumption rates by fish. To test this hypothesis, we compared plant communities in seasonally flooded forests and adjacent upland forest in terms of fruiting phenology, the frequency of trees bearing fleshy fruit, and the role of fleshy tissues in buoyancy and seed viability. Beta‐diversity in this system is high, with significant differences in species composition across habitats. As predicted, the production of ripe fleshy fruits was significantly greater in flooded than upland forests during the flood season. Furthermore, we found that trees with fleshy fruit were significantly more abundant in flooded forests even though species richness of fleshy fruit‐bearing trees was proportionally similar in flooded and upland forests. Additionally, fleshy pulp increased buoyancy. Likewise, time afloat decreased for denser fruit and those with high seed to pulp ratios. In concert, these results suggest that fleshy fruits in Neotropical floodplain forests facilitated hydrochory and ichthyochory. Once established, water and fish became important agents of selection on fruiting traits.     

The influence of pulp lipids on fruit preference by birds

Use patterns of lipid-rich, fleshy fruits were examined for frugivorous birds. Approximately 20% of fleshy-fruited species within plant communities bear fruit with >10% dry weight lipids. These fruits are fed upon by the most heavily frugivorous bird species in those communities studied and are the focus of highly specialized relationships among birds and fruits. In aviary tests some temperate frugivores show preference for higher lipid native fruits and prefer higher-lipid artificial fruits with as little as 3% wet weight difference in lipid content. The importance of lipids in fruit preference by birds, and the possible effects of differing digestive physiologies in birds are discussed.     

Can Functional Traits Predict Ecological Interactions? A Case Study Using Rain forest Frugivores and Plants in Australia

In rain forest, the large numbers of species of fleshy-fruited plants and frugivorous animals result in a large number of potential fruit–frugivore interactions, which are challenging to survey in the field. Yet, knowledge of these relationships is needed to predict consequences of changes in the frugivore assemblage for seed dispersal. In the absence of comprehensive dietary information, it may be possible to delineate between frugivores that disperse different plants using ‘functional traits,’ or morphological and behavioral attributes of frugivores that interact with differences in salient characteristics of plant species. Here we use data on the consumption of 244 Australian rain forest plant species by 38 bird species to test for associations between patterns of frugivory and birds': (1) degree of frugivory, (2) gape width, and (3) seed treatment (seed crushing or seed dispersing). Degree of frugivory and gape width explain 74 percent of the variation in the sizes of fruits consumed by frugivorous birds. Among birds that consume a substantial dietary proportion of fruit, birds with wider gapes consume larger fruits. In contrast, this relationship was not shown by birds for which fruit is only a minor dietary component. Degree of frugivory and gape width, together with seed treatment, also strongly predict the overall taxonomic composition and diversity of plants consumed by bird species. Functional classifications of frugivore species may prove useful in developing a predictive understanding of fruit–frugivore interactions in other rain forest regions where detailed dietary information is not available for most frugivores.     

Seed size selection by olive baboons

Seed size is an important plant fitness trait that can influence several steps between fruiting and the establishment of a plant’s offspring. Seed size varies considerably within many plant species, yet the relevance of the trait for intra-specific fruit choice by primates has received little attention. Primates may select certain seed sizes within a species for a number of reasons, e.g. to decrease indigestible seed load or increase pulp intake per fruit. Olive baboons (Papio anubis, Cercopithecidae) are known to select seed size in unripe and mature pods of Parkia biglobosa (Mimosaceae) differentially, so that pods with small seeds, and an intermediate seed number, contribute most to dispersal by baboons. We tested whether olive baboons likewise select for smaller ripe seeds within each of nine additional fruit species whose fruit pulp baboons commonly consume, and for larger seeds in one species in which baboons feed on the seeds. Species differed in fruit type and seed number per fruit. For five of these species, baboons dispersed seeds that were significantly smaller than seeds extracted manually from randomly collected fresh fruits. In contrast, for three species, baboons swallowed seeds that were significantly longer and/or wider than seeds from fresh fruits. In two species, sizes of ingested seeds and seeds from fresh fruits did not differ significantly. Baboons frequently spat out seeds of Drypetes floribunda (Euphorbiaceae) but not those of other plant species having seeds of equal size. Oral processing of D. floribunda seeds depended on seed size: seeds that were spat out were significantly larger and swallowed seeds smaller, than seeds from randomly collected fresh fruits. We argue that seed size selection in baboons is influenced, among other traits, by the amount of pulp rewarded per fruit relative to seed load, which is likely to vary with fruit and seed shape.     

Factors affecting seed rain beneath fleshy-fruited plants

We investigated factors affecting seed rain beneath nine fleshy-fruited fruiting plant species growing in a 1-ha plot of planted Pinus thunbergii in central Japan. We tested whether the numbers of seeds and seed species dropped by birds beneath fruiting plants were correlated with the number of fruits removed by birds from the plants. Most of fruiting plant species with high fruit removal had significantly high seed rain. Both the numbers of seeds and seed species dropped were significantly, positively correlated with the number of fruits removed across for all fruiting plant species. Therefore, fruit removal predicted the difference among heterospecific fruiting plants in seed rain. We also tested whether the number of fruits removed from fruiting plants by birds was related with fruit crop size, fruit size, and height of the plants, and the numbers of fruits and fruit species of neighboring plants near the plants. Most of fruiting plant species with high fruit crop size had significantly high fruit removal. The number of fruits removed was significantly, positively correlated with both the fruit crop size and the number of neighboring fruits across the nine fruiting plant species. However, the effect of the neighboring fruit density on fruit removal was lower remarkably than that of fruit crop size. Therefore, fruit crop size best predicted the differences among heterospecific fruiting plants in fruit removal. We suggest that fruiting plant species with high fruit crop size and high fruit removal contribute to intensive seed rain beneath them. This revised version was published online in June 2006 with corrections to the Cover Date.     

Interaction between juniper Juniperus communis L. and its fruit pest insects: Pest abundance, fruit characteristics and seed viability

The relationships between the fruit features of Juniperus communis and the presence of fruit pests were studied in Sierra Nevada, SE Spain. The abundance of two insect species — a pulp-sucking scale and a seed-predator wasp — was surveyed with respect both to fruit characteristics and to viability of seeds contained therein. Seed-predator pressure was not significantly related to any fruit characteristics; however, pulp suckers tended to be more abundant in plants with low pulp: seed ratios and high fruit-water content. In addition, fruits with high levels of pulp-sucker attack tended to have higher water content. A multi-factor ANOVA, considering the identity of the plant and the attack of the different pests as factors, showed that plant identity accounts for most of the variation in fruit characteristics. The viability of seeds tended to be lower in plants strongly attacked by both pests. Fruits attacked by seed predators showed significantly lower proportions of viable and unviable seeds than did unattacked fruits. Seed viability was also lower in those fruits heavily attacked by pulp suckers, but this pattern is strongly mediated by plant identity. Pest activity proved to be clearly associated with a direct decrease in juniper reproductive capacity. This loss involved a reduction of the viable-seed number, mainly related to the seed predator, as well as a reduction of fruit attractiveness to frugivorous dispersers, related to the pulp sucker.     

The persistence of ripe fleshy fruits in the presence and absence of frugivores

A trade-off between antimicrobial defences and palatability to dispersers may place limits on fruit persistence in nature. The retention times of ripe fruits on 34 wild plant species under natural conditions (unbagged persistence) and when fruits had been bagged with nylon mesh to exclude frugivores (bagged persistence) were compared in Hong Kong, China (22°N). Bagged persistence is a measure of the effectiveness of fruit defence while unbagged persistence is an inverse measure of attractiveness to vertebrate frugivores. Bagged fruits persisted significantly longer than unbagged fruits in 30 species, with half the species tested persisting for more than 2 months. There was a significant positive relationship between the median persistence times of bagged and unbagged fruits, suggesting that species with a high resistance to microbial infection are also less attractive to frugivores. Both bagged and unbagged fruits persisted significantly longer at lower temperatures. There was a significant positive relationship between bagged persistence time and fibre content of the fruit pulp, but no significant relationships between unbagged persistence and the six fruit traits tested (diameter, pulp as a percentage of fruit fresh weight, and lipid, total soluble carbohydrate, nitrogen and fibre as percentages of pulp dry weight). Mechanical damage significantly decreased the bagged persistence time for half of the species. Although some fruits decayed or dried up while attached to the plant, fruits of 53% of the species remained visually attractive until they fell off.     

EFFECT OF TEMPERATURE AND PHOTOPERIOD ON GROWTH AND DORMANCY OF BETULA PAPYRIFERA

The effects of day/night temperatures and photoperiod on the growth and dormancy of paper birch (Betula papyrifera) were studied in seedlings from different geographic origins. The response of Alaskan plants to temperature and photoperiod was distinctly different from other seed sources. Alaskan plants required very long days to prevent cessation of growth while plants from southern seed sources grew on photoperiods as short as 14 hr. Low night temperature (14 C) antagonized the promotive action of long photoperiods in Alaskan plants but had little effect in other seed sources. High day temperatures offset the inhibitory effect of the cool night to a lesser degree in Alaskan plants than in plants from other locations. Dormancy induced by short photoperiods was antagonized (relieved ?) to a lesser degree by high night temperatures in Alaskan birch than in other seed sources. Betula papyrifera var. humilis from Alaska may be an incipient species since its morphological traits are accompanied by adaptive physiological responses to its environment. These responses are as distinct as its morphological characteristics.     

Host specificity and interaction networks of insects feeding on seeds and fruits in tropical rainforests

In the tropics, antagonistic seed predation networks may have different properties than mutualistic pollination and seed dispersal networks, but the former have been considerably less studied. We tested whether the structure of antagonistic tripartite networks composed of host plants, insects developing within seeds and fruits, and their insect parasitoids could be predicted from plant phylogenetic distance and plant traits. We considered subsets of the networks (‘subnetworks') at three rainforest locations (Panama, Thailand, Papua New Guinea), based on insect families, plant families or plant functional groups. We recorded 3197 interactions and observed a low percentage of realized interactions, especially in Panama, where insect host specificity was higher than in Thailand or New Guinea. Several factors may explain this, including insect faunal composition, incidence of dry fruits, high fruit production and high occurrence of Fabaceae at the Panamanian site. Host specificity was greater among seed-eaters than pulp-eaters and for insects feeding on dry fruits as opposed to insects feeding on fleshy fruits. Plant species richness within plant families did not influence insect host specificity, but site characteristics may be important in this regard. Most subnetworks were extremely specialized, such as those including Tortricidae and Bruchinae in Panama. Plant phylogenetic distance, plant basal area and plant traits (fruit length, number of seeds per fruit) had important effects on several network statistics in regressions weighted by sampling effort. A path analysis revealed a weak direct influence of plant phylogenetic distance on parasitoid richness, indicating limited support for the ‘nasty host hypothesis'. Our study emphasizes the duality between seed dispersal and seed predation networks in the tropics, as key plant species differ and host specificity tends to be low in the former and higher in the latter. This underlines the need to study both types of networks for sound practices of forest regeneration and conservation.     

Nutrients in fruits as determinants of resource tracking by birds

Fruit pulp is an important source of nutrients for many bird species. Fruit‐eating birds use a variety of strategies to cope with changes in the availability of fruits, exhibiting a remarkable ability to track resources. We assessed the role of nutrient availability in the fruiting environment as a factor driving resource tracking by fruit‐eating birds. Fruit consumption by the four most common frugivorous species in a 6‐ha plot in the Southern Yungas montane forest of Argentina was assessed. We determined the content of selected nutrients (soluble carbohydrates, proteins, phenols, ascorbic acid and essential minerals) in 22 fruiting plant species eaten by birds, and measured fruit–frugivore interactions and the availability of nutrients and dry fruit pulp mass over 2 years. There was strong temporal covariation in the availability of the selected nutrients in fruits across the study period. Similarly, the availability of nutrients in the fruiting environment covaried with pulp mass. Fruit consumption by the four commonest bird species and the abundance of most species were positively associated with nutrient availability and dry pulp mass. Nutrient availability was a good predictor of temporal fruit tracking by three of the four commonest frugivores. Despite large differences in particular nutrient concentrations in fruits, overall nutrient (and pulp) quantity in the fruiting environment played a greater role in fruit tracking than did the nutritional quality of individual fruits. While overall nutrient availability (i.e. across fruit) and total pulp mass were important determinants of fruit tracking, we suggest that plant species‐specific differences in fruit nutrient concentration may be important in short‐term foraging decisions involved in fruit choice and nutritional balance of birds.     

Do seed-dispersing birds exert selection on optimal plant trait combinations? Correlated phenotypic selection on the fruit and seed size of hawthorn (<Emphasis Type="Italic">Crataegus monogyna</Emphasis>)

Seed-dispersing birds can be expected to optimize their energy intake through fruit selection and hence exert a positive selection on fruit pulp content, that is to say, favoring big fruits with small seeds. On the other hand they may select both, average fruit size and its variation. We addressed this issue by analyzing the phenotypic selection exerted by the dispersers of Crataegus monogyna on the fruit and seed size of this species. Fruit and seed size were analyzed at two ontogenic plant stages: the initial size of seeds (and their fruits) as future individuals and the mean size of fruits and seeds produced by adult plants. Fruit diameter and its within-individual variation are the actual targets of selection for maternal fruit traits, negatively affecting relative fitness, although total selection acts only on mean fruit size. For individual seeds, size selection is positive and directed at fruit diameter and seed length, while the remaining traits are subject to indirect selection. Birds exerted a correlational selection favoring big fruits with small seeds. Nevertheless the evolutionary consequences of this selection are expected to be limited by several factors. For example, the positive correlation between fruit and seed sizes, the existence of counteracting selective pressures exerted by other mutualistic or antagonist interactions and temporal or spatial changes in the environmental conditions that may alter the selective forces and even the optimum phenotype in each situation.     

Seed mass and the evolution of fleshy fruits in angiosperms

Fleshy fruits, like drupes and berries, have evolved many times through angiosperm history. Two hypotheses suggest that fleshy fruit evolution is related to changes in the seed mass fitness landscape. The reduced dispersal capability following from an increase in seed mass may be counterbalanced by evolution of traits mediating seed dispersal by animals, such as fleshy fruits. Alternatively, increasing availability and capabilities of frugivores promote evolution of fleshy fruits and allow an increase in seed size. Both these hypotheses predict an association between evolution of fleshy fruits and increasing seed size. We investigated patterns of fruit and seed evolution by contrasting seed mass between fleshy and non‐fleshy fruited sister clades. We found a consistent association between possession of fleshy fruits and heavier seeds. The direction of fruit type change did not alter this pattern; seed mass was higher in clades where fleshy fruits evolved and lower in clades where non‐fleshy fruits evolved, as compared to their sister clades. These patterns are congruent with the predictions from the two hypotheses, but other evidence is needed to distinguish between them. We emphasize the need to integrate studies of seed disperser effectiveness, seed morphology, and plant recruitment success to better understand the frugivores’ role in fleshy fruit evolution.     

Lagged effects of sawfly leaf herbivory on reproductive organs in cherry trees: Overcompensation in flower production reduces quality of fruits and seeds

Herbivores reduce plant productivity by removing part of the assimilation surface. Also, they can alter plant traits that affect plant–pollinator interactions and reproductive success. The objective of this study was to evaluate the impact of defoliation by sawfly (Caliroa cerasi) larvae on fruit production and quality in three cultivars of sweet cherry (Prunus avium). We hypothesized that the fruit production and quality is reduced as a consequence of changes in the allocation of resources within the plant in response to partial leaf removal during the previous year. Number of flowers per branch meter was higher in infested trees than in non-infested trees, while the number of fruits per branch meter was similar due to fruit abortion in all cultivars. Fruit quality was significantly affected by herbivory in different traits depending on cultivar. Infested Lapins and Van trees had significantly lower soluble solid content than non-infested trees. Titratable acidity was higher and ripening index was lower in infested Bing and Lapins trees than non-infested trees. Infested Van trees also exhibited a significant decrease in equatorial diameter and fresh fruit weight as well as pulp fresh weight and dry weight of seed compared to non-infested trees. Overall our study highlights that the direct impact of herbivores at leaf level has lagged effects on productivity in terms of fruit and seed quality in the year following the leaf damage.