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The Plant Vigor Hypothesis (PVH) predicts that herbivorous insects will preferentially feed on large plants with more vigorous growth, or on more vigorous parts of the plant. However, we still need to understand how these predictions work throughout the ontogenic stages of plants and with other groups of natural plant enemies. We tested the PVH using gall-inducing insects, free-living herbivorous insects, and pathogenic fungi associated with Hymenaea courbaril seedlings of the same age cohort (six months) from seeds germinated in greenhouses. We classified the plants into three different sizes: small, medium, and large seedlings, and then measured above and belowground biomass, plant height, leaf area, and leaf area consumed by free-living herbivores. We also measured nutrient partitioning among roots, stems, and leaves and estimated hypersensitive responses. We found no differences in leaf area consumed by free-living herbivorous insects between plant size groups. However, we observed that larger seedlings were more vigorous and had greater nutritional value and, thus, represented adequate niches for colonization by pathogens and gall-inducing insects, confirming the PVH for both groups. The hypersensitive response was greater in medium and large seedlings, proving to be an effective defense mechanism against gall-inducing insects. Therefore, we highlight that nutritional quality, plant vigor and hypersensitivity response are factors that influence the incidence of pathogens and galling insects on H. courbaril seedlings. [ABSTRACT FROM AUTHOR]

Seasonal windows of opportunity represent intervals of time within a year during which organisms have improved prospects of achieving life history aims such as growth or reproduction, and may be commonly structured by temporal variation in abiotic factors, bottom-up factors, and top-down factors. Although seasonal windows of opportunity are likely to be common, few studies have examined the factors that structure seasonal windows of opportunity in time. Here, we experimentally manipulated host-plant age in two milkweed species (Asclepias fascicularis and Asclepias speciosa) in order to investigate the role of plant-species-specific and plant-age-varying traits on the survival and growth of monarch caterpillars (Danaus plexippus). We show that the two plant species showed diverging trajectories of defense traits with increasing age. These species-specific and age-varying host-plant traits significantly affected the growth and survival of monarch caterpillars through both resource quality- and quantity-based constraints. The effects of plant age on monarch developmental success were comparable to and sometimes larger than those of plant-species identity. We conclude that species-specific and age-varying plant traits are likely to be important factors with the potential to structure seasonal windows of opportunity for monarch development, and examine the implications of these findings for both broader patterns in the ontogeny of plant defense traits and the specific ecology of milkweed-monarch interactions in a changing world.

Comparative studies with taxonomically and geographically paired alien species that exhibit different degrees of success in their invasions may help to identify the factors that determine invasiveness. Examples of such species in Europe include the noninvasive Impatiens balfourii and invasive I. glandulifera. We tested whether the low invasiveness of I. balfourii in Europe may be explained by strong pressure from local enemies. Earlier studies of these two species provided support for their hybridization. We tested this phenomenon as the potential occurrence of I. glandulifera × I. balfourii hybrids might promote the evolution of the invasiveness of I. balfourii. Both species were germinated from seeds collected in 2015 on the Swiss-Italian border in Insubria and utilized in three experiments: (1) a common garden enemy release test (leaf damage or pest pressure), (2) a test of the pressure exerted by a generalist enemy and (3) hybridization test. In the first test, the effect of enemies was assessed by the level of leaf damage and the number of pests. In the second test, a food choice experiment with a generalist herbivore (Cepaea snails) was performed. In the hybridization test, the plants were placed in a climatic chamber for self-pollination and hand cross-pollination. Analyses of enemy release and Cepaea snail preference revealed that I. balfourii experienced higher enemy pressure than I. glandulifera; however, this was not reflected in the performance of the plants. Although I. glandulifera was larger, I. balfourii had greater fecundity. Thus, the invasion success of I. glandulifera could not be unambiguously attributed to its greater degree of release from enemies compared with the noninvasive I. balfourii. Additionally, we did not obtain any evidence of hybridization between the two species. Thus, we obtained no support for the hypothesis that the evolution of the invasiveness of I. balfourii could be enhanced through hybridization with I. glandulifera. [ABSTRACT FROM AUTHOR]

Regulated deficit irrigation, which is a common practice to cope with water scarcity, can impact plant-aphid interactions, and possibly lead to a reduction in the use of pesticides. To test the possible effect of water restriction on the apple tree-rosy apple aphid ( Malus domestica-Dysaphis plantaginea ) system, we performed a factorial experiment with two levels of water supply and two genotypes on artificially infested trees. Plant growth and aphid abundance were characterised during the entire infestation period at two scales of analysis: the apical shoot scale and the tree scale, and additional measures were performed to evaluate plant water status. Aphid abundance increased with plant vegetative growth at both scales of analysis, which is consistent with the Plant Vigor Hypothesis (i.e. with the fact that most of the phloem feeders prefer fast growing plants). At the tree scale, aphid abundance was lower on trees that underwent water restriction, but at the shoot scale, aphid abundance responded differently to water restriction depending on the tree genotype. Water restriction modified the relationship between aphid abundance and growth, thus indicating that host suitability for aphids was affected by different plant variables susceptible to water stress, among which growth. The different response patterns at the two scales of analysis highlight the importance of scale choice in the study of plant-insect interactions. [ABSTRACT FROM AUTHOR]

The few existing analyses of broad-scale variation in plant–herbivore interactions are largely exploratory and not based on hypotheses on the effect of varying environmental conditions on the patterns of herbivore damage. However, there are two established hypotheses that relate herbivore damage and environmental stress mostly tested at the within-population level: the Plant Stress Hypothesis (PSH) and the Plant Vigor Hypothesis (PVH). To examine if support for the PSH holds true at broader scales, we tested it at the micro- (among individuals), meso- (between habitats) and macroscale (between geographical regions, Iberia and Macaronesia). We measured plant stress and leaf damage by different natural enemies (chewers, miners and pathogens) in 12 populations across the range of the relict tree Prunus lusitanica . Results regarding the proportion of leaves attacked by any enemy showed no support for the PSH at either micro or mesoscale, observing instead higher damage in less stressed individuals within populations and in high-quality habitats. However, the latter was true exclusively within the Macaronesian region, highlighting the important effect of the type of abiotic stress on plant susceptibility to herbivory. At the macroscale, alternatively, results showed support for the PSH: we observed a higher proportion of leaves attacked in Macaronesia than in Iberia, where plants unexpectedly exhibited greater stress symptoms. Among-scale inconsistencies may respond to the effect of distinct climatic factors governing both plant and herbivore dynamics at large scales, particularly in species with distributional ranges across contrasting climatic regions such as P. lusitanica . Finally, different plant enemies showed different patterns of damage, corroborating the need to study them separately to achieve consistent generalizations of these hypotheses. [ABSTRACT FROM AUTHOR]

Pattern of attack of a galling insect reveals an unexpected preference-performance linkage on medium-sized resources. The Plant Vigor Hypothesis (PVH) predicts oviposition preference and higher offspring performance on longer and fast-growing shoots, and although several studies have tested its predictions, long-term studies concerning the patterns of host selection by galling species are still lacking. The PVH was tested in this study using Bauhinia brevipes (Fabaceae) as the host of a leaf gall midge, Asphondylia microcapillata (Diptera, Cecidomyiidae) during three consecutive years. Shoots were collected from the same 80 plants between 2001 and 2003 and shoot length, number of healthy and galled leaves, gall number, and mortality factors were recorded. Nearly 600 galls were found on the 5,800 shoots collected. Medium-sized shoots supported from 46 to 70% of all galls, with greater gall survival rate in 2002 and 2003. A decrease in parasitism rate coupled with an increase in gall predation lead to a constant similar gall survivorship rate in all years (x = 22.7%). Although gall abundance varied among years (122 in 2001, 114 in 2002 and 359 in 2003) preference for longer shoots was not observed because the percentage of galled shoots and galled leaves were higher on medium shoot length classes in all years. The observed distribution of gall abundance and galled shoots were always greater than the expected distribution on medium shoot length classes. These findings do not support the PVH, and show that A. microcapillata can maximize the female preference and larval performance on medium-sized shoots of B. brevipes.A Hipótese do Vigor de Plantas (HVP) prevê uma oviposição preferencial e alta performance da prole em ramos longos e de crescimento rápido da planta hospedeira. Embora diversos estudos tenham testado suas predições, estudos de longa duração focados no padrão de seleção de planta hospedeira por insetos galhadores ainda são escassos. A HVP foi testada neste estudo usando Bauhinia brevipes (Fabaceae) como planta hospedeira de uma galha foliar induzida por Asphondylia microcapillata (Diptera: Cecidomyiidae) por três anos consecutivos. Ramos foram coletados das mesmas 80 plantas entre 2001 e 2003 e o tamanho do ramo, número de ramos galhados e sadios, número de galhas e fatores de mortalidade foram registrados. Aproximadamente 600 galhas de A. microcapillata foram coletadas em 5.800 ramos de B. brevipes. Ramos médios suportaram entre 46% e 70% do total de galhas, e a taxa de sobrevivência das galhas foi maior em 2002 e 2003. Uma diminuição da taxa de parasitismo associada com um aumento da taxa de predação de galhas resultou em uma sobrevivência similar em todos os anos (x = 22,7%). A abundância de galhas variou entre os anos (122 em 2001, 114 em 2002 e 359 e 2003). A preferência por ramos longos não foi observada porque a porcentagem de ramos galhados e folhas galhadas foram maiores em ramos de classes de tamanho intermediário por dois anos. A distribuição observada da abundância de galhas e ramos galhados foi sempre maior do que a distribuição esperada em classes de ramos médios. Estes resultados não suportam a HVP, e mostram que A. microcapillata pode maximizar a preferência da fêmea e a performance larval em ramos de tamanho médio de B. brevipes.

The availabilities of water and nitrogen resources are among the main abiotic factors modifying plant physiology and growth. Consequently, via bottom-up processes, they also can have an impact on the performance of herbivorous insects. However, the extent to which the modification of plant growth is responsible for such an impact remains unclear., We conducted a factorial experiment quantifying Myzus persicae aphid abundance and the vegetative growth of Prunus persica peach tree shoots under contrasting levels of nitrogen and water supplies. We used a hierarchical analysis of multiple regression models to determine whether the consequences of the availability of nitrogen and water on aphid abundance could be the result of a modification of plant growth., Maximum aphid abundance was achieved under nitrogen and water comfort conditions. The best model explaining variance in aphid abundance took into account vegetative growth and water supply, as well as their interaction., The results of the present study suggest that a higher nitrogen supply increases aphid abundance by fostering plant growth. Additionally, the positive response of aphid abundance to vegetative growth is lower in the case of water restriction because, under such conditions, aphids cannot take full advantage of tree vigour. Such a result provides new insights into aphid control in agriculture, as well as on the possible effects of climate change. [ABSTRACT FROM AUTHOR]

The effects of bottom-up forces on the community structure of tropical insect gallers and within species variation in gall morphotype assemblages are not well understood., We tested the roles of increased nutritional quality and density of host plants with respect to structuring the galling insect communities and gall morphotype assemblages on Neoboutonia macrocalyx trees in Uganda., Plant nutritional quality and resource concentration were manipulated with four levels of fertilization and the group size of trees, respectively. After applying these treatments in May 2011, gallers established naturally on experimental replicates. Five months later, we sampled gallers and recorded their numbers and the different morphotypes. We analyzed the effects of treatments on the structures of galler communities and gall morphotype assemblages., We recorded 5237 individuals, representing four galler species. One species, Cecidomyiid leaf galler, was represented by three morphologically distinct galls. Fertilization, host tree density and their interactions significantly changed the structures of galler species communities and gall morphotype assemblages., The results of the present study demonstrate the important role of bottom-up factors in structuring galler communities and the gall morphotype assemblages. These changes are likely caused by differential responses of different galler species and gall morphs to plant quality or quantity changes. [ABSTRACT FROM AUTHOR]

Gall insects select vigorously growing plants and plant parts when initiating gall formation. Vigor is associated with rapid growth rate, and in turn, rapid growth confers competitiveness. Are there conditions under which the cost of vigor, in the form of increased susceptibility to attack, outweighs the benefit of competitive success? I present a simulation model to explore the interaction between susceptibility and competition on the selective advantage of increased growth rate. Assuming size-symmetric competition, the model shows that in general, vigor is favored (benefit > cost) at low to intermediate gall loads. At very high plant densities, however, plants with high gall loads may lose standing in the competitive size hierarchy from which they cannot recover. The details of this result, however, change somewhat when competition is size-asymmetric, that is, when a larger focal plant suppresses smaller neighbors, but smaller neighbors cannot exert a reciprocal effect on the focal. At low densities, the pattern of selection on growth rate is qualitatively similar to the size-symmetric case. At higher plant densities, however, fast-growing genotypes can suppress slow ones so much during the preattack phase that even at the highest gall loads they maintain their standing in the competitive hierarchy. Thus, heavy gall insect attack on vigorous plants can impose selection against high intrinsic growth rates under strong symmetric competition, but not strong asymmetric competition. While life history traits can evolve as a correlated response to selection on defensive traits that reduce susceptibility, this model reveals that susceptibility can evolve as a correlated response to selection on basic life history traits. [ABSTRACT FROM AUTHOR]

Although several hypotheses aim to explain insect herbivory on plants, the relative importance of plant traits, environment, and organizational scale (i.e., individual or community) to herbivory damage level is not well understood. We used an approach based on a local scale, divided into individual and community levels, to test if plant traits, soil characteristics, and plant density explain leaf damage. We sampled 983 individuals in 49 plots distributed over dense and open savanna formations in Emas National Park. In order to explain plant damage, we performed a multi-model inference analysis of four plant traits associated with plant damage, five soil characteristics, and plant density. We did not find any support to plant vigor or plant stress hypotheses at individual plant level. However, the resource concentration hypothesis and plant stress hypothesis explained leaf damage at the community level. We found that increased availability of calcium (Ca) in soils reduced plant damage at the community level. Because soil Ca concentration is a major constraint to plant development in the Brazilian savanna, we postulated that its increasing availability permits plants to invest more in defense strategies. We demonstrate that plant density, Ca soil concentration, and leaf size can be used to predict the plant damage suffered by woody species in savannas at community level. [ABSTRACT FROM AUTHOR]

Gall-site selection by the aphid Kaltenbachiella japonica was evaluated in relation to leaf position in a shoot, and gall positions within a leaf. First-instar fundatrices induce closed galls on the midribs of host leaves, and several galls were often induced on one leaf. Leaves with many galls were often withered before emergence of sexuparae from the galls. Within a leaf, gall volume was positively correlated with the sum of lateral-vein length in the leaf segment at which the gall was induced. The observed pattern in gall volume among the leaf segments corresponded with that in the lateral-vein length. These results show that a foundatrix selects the most vigorous position within a leaf to produce more offspring. Although distal leaves grew faster than did basal leaves, gall density was highest on leaves at the middle order when a shoot has more than seven leaves. Optimal gall-site selection seems to be constrained by the asynchrony in timing between the hatching of fundatrices and leaf growth within a shoot. These results suggest that the observed gall distribution is affected by both the distribution of suitable galling sites within a leaf and the synchrony with leaf phenology of the host plant. [ABSTRACT FROM AUTHOR]

The Plant Vigor Hypothesis (PVH) proposes that natural selection on female oviposition choice results from higher fitness of larvae on more vigorous and larger plant modules. For six consecutive years we tested the PVH predictions by investigating the effect of shoot size of Bauhinia brevipes (Fabaceae) on the oviposition preference and offspring survival of the gall-midge Schizomyia macrocapillata (Diptera: Cecidomyiidae). Additionally, we analyzed the effects of bottom-up and top-down mortality forces on the system. The hypersensitive reaction (bottomup effect) accounted for more than 90% larval mortality of S. macrocapillata, making available few galls to be found and killed by natural enemies (top-down effect). Smaller shoots were always more abundant while longer shoots were rare. Nevertheless, the percent number of galls induced by S. macrocapillata was up to 10-fold greater on the largest shoots, corroborating the preference prediction of the PVH. Schizomyia macrocapillata should use over-exploit larger shoots to maximize the preference for, and consequently increase the performance on these shoots. Our results partially support the performance prediction of the PVH: (1) the observed survival was higher than expected on longer shoots, and (2) the ratio of survival per shoot was positively related with shoot length only in 2 years. Thus, we found a link between female preference and larval performance on large-sized shoots, at least in some years. The gall-midge attack pattern in this study might be an evolved response to maximize the female preference and increased larval performance on longer shoots of the host plant. [ABSTRACT FROM AUTHOR]

A basic question in insect–plant interactions is whether the insects respond to, or regulate plant traits, or a complex mixture of the two. The relative importance of the directions of the influence in insect–plant interactions has therefore been articulated through both the plant vigor hypothesis (PVH) and the resource regulation hypothesis (RRH). This study tested the applicability of these hypotheses in explaining the interactions betweenParthenium hysterophorusL. (Asteraceae) and its stem-galling moth,Epiblema strenuanaWalker (Lepidoptera: Tortricidae). Parthenium plants exposed to galling were sampled at three sites in north Queensland, Australia, over a 2-year period, and the relationship between gall abundance and plant vigor (plant height, biomass, flowers per plant, and branches per plant) was studied. To test the predictions of PVH and RRH, the vigor of parthenium plants protected from galling using insecticides was compared to galled plants and plants that escaped from galling. The vigor of ungalled plants was less than the vigor of galled plants. The higher plant vigor in galled plants was not due to galling, as was evident from insecticide exclusion trials. The insect seemed to preferentially gall the more vigorous plants. These findings support the predictions of the PVH and are contrary to those of RRH. Since gall abundance is linked to plant vigor, galling may have only a limited impact on the vigor of parthenium. This has implications for weed biological control. If the objective of biological control is to regulate the population of a plant by a galling insect, a preference for more vigorous plants by the insect is likely to limit its ability to regulate plant populations. This may explain the paucity of successes against biocontrol of annual weeds using gall insects. [ABSTRACT FROM AUTHOR]

To test the Plant Vigor Hypothesis, we determined female oviposition preference of Phyllocolpa leavitii (Hymenoptera: Tenthredinidae) for shoot lengths on 20 clones of Salix discolor (Salicaceae) and examined larval performance by determining larval survival based on clone, shoot length, and leaf length for each Phyllocolpa gall observed. Sawfly galls were found on significantly longer shoots compared to shoots without galls, and shoots with 2, 3, 4, and 5 galls were successively longer than shoots with fewer galls. Furthermore, a much higher proportion of longer shoots had galls that did shorter shoots. These data demonstrate strong sawfly oviposition preference for long shoots. However, when shoot length was adjusted for numbers of available leaves per shoot, the number of galls per available leaf showed a weaker, but still positive relationship with shoot size. Using a logistic regression on survival of Phyllocolpa larvae, we detected highly significant effects of clone, shoot length, leaf length, and the square of leaf length on survival. Additionally, much higher sawfly survival on the long shoots of one clone caused a significant clone by shoot-length interaction. Survival was positively correlated with shoot length, indicating directional selection favoring oviposition on longer shoots; still, shoot length explained only 11.8% of the variation in survival. The directional component of selection favored oviposition on smaller leaves as survival was inversely related to leaf length; however, disruptive selection for leaf length oviposition preference was also detected, with survival of Phyllocolpa galls lowest on intermediate leaf lengths. This study provided evidence supporting the preference prediction of the Plant Vigor Hypothesis. However, much of the data contradicted the performance prediction of the Plant Vigor Hypothesis, with only a modest amount of evidence supporting the performance prediction. [ABSTRACT FROM AUTHOR]

The plant stress and plant vigor hypotheses are widely used to explain the distribution and abundance of insect herbivores across their host plants. These hypotheses are the subject of contentious debate within the plant herbivore research community, with several studies finding simultaneous support for both hypotheses for the same plant–herbivore interaction. We address the question of how such support is possible using dynamic site‐occupancy models to quantify the attack dynamics of Cryptorhynchus lapathi (poplar‐willow weevil) on Salix sitchensis (Sitka willow), a dioecious shrub colonizing Mount St. Helens, Washington, USA after the 1980 eruption, in relation to host plant stress, vigor, and sex. We also introduce several scaling criteria as a rigorous test of the plant vigor hypothesis and demonstrate why modeling insect detection is important in plant–insect studies. Weevils responded positively to water stress associated with seasonal dry‐downs, and this response was phenologically compartmentalized by larval feeding mode. Weevils preferentially attacked large and/or flowering stems, imposing an ecological cost on willow reproduction via increased stem mortality and susceptibility to future attack. We propose that the dual response to host plant stress and vigor is due to the synchronization between young weevil larval feeding and willow nutrient pulses that are mediated by environmental stress. In turn, this process drives successional dynamics, causing the juvenilization of upland willow plants and possibly delaying establishment of a willow‐dominated upland sere. These results highlight the common, but often overlooked, phenological basis of the plant stress and plant vigor hypotheses, which both focus on how stress changes the quality of plant resources available to immature insects. [ABSTRACT FROM AUTHOR]