Consequences For Plant Fitness Research Articles

The interplay between defaunation and phylogenetic diversity affects leaf damage by natural enemies in tropical plants

Abstract Natural enemies play an important role in controlling plant population growth and vegetation dynamics. Tropical rainforests host the greatest diversity of herbivores, from large mammalian ungulates to microscopic pathogens, generating and maintaining plant diversity. By feeding on the same resources, large mammalian herbivores may interfere with plant consumption and leaf damage by important enemy guilds such as invertebrate herbivores and pathogens, triggering indirect trophic cascades. However, the impact of local extinctions of large herbivores on plant–enemy interactions is relatively unknown. We experimentally tested the effects of defaunation of large mammalian herbivores (e.g. peccaries, tapirs and brocket deer; hereafter, large herbivores) on the leaf damage of 3350 understorey plants in tropical rainforests of Brazil. We examined leaf damage in 10,050 leaves from 333 morphospecies by assigning the area consumed or damaged by five guilds of insect herbivores and leaf pathogens within 86 paired open‐closed plots and investigating the joint effects of defaunation and plant phylogenetic diversity. Plants released from large herbivores had 9% less leaf damage; this difference was due to the lower leaf pathogens incidence (29%) rather than insect herbivory. Evolutionary distinctness was positively correlated with leaf damage in a similar way in all treatments, suggesting additive effects of defaunation and phylogenetic diversity. Total and pathogenic leaf damage (but not insect damage) decreased with plant richness across treatments, and large herbivores exclusion resulted in increased plant species richness. This suggests that large herbivores exclusion leads to a dilution of total and pathogens' leaf damage by increasing plant species richness. Our results suggest that indirect effects of large herbivores decrease the dilution potential of plant communities against pathogens and rather reinforce their top‐down impact on vegetation, demonstrating a previously overlooked cascading effect of large herbivore extinction on forest ecosystems. Synthesis: The extinction of large mammalian herbivores can lead to a decrease in pathogen‐driven leaf damage, a previously unknown indirect effect in forest ecosystems, which might have consequences for plant fitness and ultimately for plant diversity. Large herbivores and plant pathogens might have synergistic effects in regulating the diversity of plant communities in some of the most diverse ecosystems on Earth.

Insect herbivory but not plant pathogen infection drive floral volatile‐mediated indirect effects on pollinators and plant fitness in Brassica rapa

Abstract Plant enemies can indirectly affect pollinators by modifying plant traits, but simultaneous tests of herbivore and pathogen effects are lacking, and the role of floral volatiles has seldom been assessed. In this study, we tested for indirect effects of insect herbivores and pathogens on pollinator attraction via altered floral volatile emissions, and its consequences for plant fitness in Brassica rapa. Plants in the field were exposed to either no damage or damage by caterpillars (Mamestra brassicae), aphids (Brevicoryne brassicae), a leaf fungus (Sclerotinia sclerotiorum), or a bacterium (Xanthomonas campestris pv. campestris). We then recorded pollinator visits and measured floral traits (flower number, volatiles) and plant fitness‐correlates. We additionally performed a greenhouse experiment with artificial floral emitters to test for effects of target volatiles on pollinator attraction. In the field experiments, relative to controls, plants subjected to herbivory by the aphid B. brassicae (but not those exposed to the other enemies) exhibited a marked reduction in the emission of two volatile organic compounds (nonanal and 2‐butyl‐1‐octanol), experienced lower pollinator visits and produced seeds of lower quality in terms of seed biomass and germination rate, while flower output itself was not affected. Consistently, artificial emitters with reduced amounts of these volatile organic compounds were less attractive to pollinators under greenhouse conditions. Synthesis. These results provide strong evidence for volatile‐mediated indirect interactions between plant enemies and pollinators ultimately impacting plant fitness, and further point at enemy and compound specificity in such effects.

The effect of species role and trait-matching on plant fitness in a plant-hummingbird interaction network

In animal-pollinated plants, fitness is affected by pollinator attraction and plant-pollinator trait matching. Plant-pollinator interactions are often characterized using metrics derived from mutualistic networks, such as centrality and interaction selectivity. This approach often focuses on the species level, thus neglecting significant among-individual variation in attracting foraging pollinators and associated fitness consequences for plants. Here, we tested if centrality and interaction selectivity were related to fitness at the species and the individual level for 14 hummingbird-pollinated plant species in the Atlantic Forest. In addition, we tested if plant-pollinator trait-matching influenced plant fitness. At the species-level, we did not find a relationship between plant fitness and centrality nor interaction selectivity; we found instead that plant fitness increased with higher trait-matching between bill and corolla lengths. At the individual-level we found that individuals with higher centrality values produced less seeds, likely due to lower quality in pollen deposition. Moreover, individuals with a lower interaction selectivity also produced more seeds, probably because of higher conspecific pollen flow. Our results show that individual position and interaction selectivity within pollination networks may have important implications for pollination effectiveness and hence plant fitness.

Insect exuviae as soil amendment affect flower reflectance and increase flower production and plant volatile emission.

Soil composition and herbivory are two environmental factors that can affect plant traits including flower traits, thus potentially affecting plant-pollinator interactions. Importantly, soil composition and herbivory may interact in these effects, with consequences for plant fitness. We assessed the main effects of aboveground insect herbivory and soil amendment with exuviae of three different insect species on visual and olfactory traits of Brassica nigra plants, including interactive effects. We combined various methodological approaches including gas chromatography/mass spectrometry, spectroscopy and machine learning to evaluate changes in flower morphology, colour and the emission of volatile organic compounds (VOCs). Soil amended with insect exuviae increased the total number of flowers per plant and VOC emission, whereas herbivory reduced petal area and VOC emission. Soil amendment and herbivory interacted in their effect on the floral reflectance spectrum of the base part of petals and the emission of 10 VOCs. These findings demonstrate the effects of insect exuviae as soil amendment on plant traits involved in reproduction, with a potential for enhanced reproductive success by increasing the strength of signals attracting pollinators and by mitigating the negative effects of herbivory.

Flight-Fecundity Trade-offs: A Possible Mechanistic Link in Plant-Herbivore-Pollinator Systems.

Plant–herbivore and plant–pollinator interactions are both well-studied, but largely independent of each other. It has become increasingly recognized, however, that pollination and herbivory interact extensively in nature, with consequences for plant fitness. Here, we explore the idea that trade-offs in investment in insect flight and reproduction may be a mechanistic link between pollination and herbivory. We first provide a general background on trade-offs between flight and fecundity in insects. We then focus on Lepidoptera; larvae are generally herbivores while most adults are pollinators, making them ideal to study these links. Increased allocation of resources to flight, we argue, potentially increases a Lepidopteran insect pollinator’s efficiency, resulting in higher plant fitness. In contrast, allocation of resources to reproduction in the same insect species reduces plant fitness, because it leads to an increase in herbivore population size. We examine the sequence of resource pools available to herbivorous Lepidopteran larvae (maternally provided nutrients to the eggs, as well as leaf tissue), and to adults (nectar and nuptial gifts provided by the males to the females), which potentially are pollinators. Last, we discuss how subsequent acquisition and allocation of resources from these pools may alter flight–fecundity trade-offs, with concomitant effects both on pollinator performance and the performance of larval herbivores in the next generation. Allocation decisions at different times during ontogeny translate into costs of herbivory and/or benefits of pollination for plants, mechanistically linking herbivory and pollination.

Pollinator-mediated facilitation alleviates pollen limitation in a plant-hummingbird network.

Facilitation and competition among plants sharing pollinators have contrasting consequences for plant fitness. However, it is unclear whether pollinator-mediated facilitation and competition may affect pollen limitation (potential contribution of pollination to fitness) in pollination networks. Here, we investigated how pollinator sharing affects pollen limitation in a tropical hummingbird-pollinated community marked by facilitation. We employed indices describing how much a plant species potentially affects the pollination of other co-flowering species through shared pollinators (acting degree) and is affected by other co-flowering species (target degree) within the plant-hummingbird network. Since facilitation often increases pollination quantity but not necessarily quality, we expected both indices to be associated with reductions in pollen limitation estimates that depend on pollination quantity (fruit set and seed number) rather than estimates more strictly related to quality (seed weight and germination). We found that both indices were associated with reductions in pollen limitation only for seed weight and germination. Thus, facilitation occurred via qualitative estimates of pollen limitation. Our results suggest that facilitation may enhance plant fitness estimates even if quantitative components of plant fecundity are already saturated. Overall, we showed that pollinator-mediated indirect effects in a multispecies context are important drivers of plant fitness estimates with consequences for coexistence in diverse communities.

Pollinator-Mediated Indirect Effects on Plant Fecundity Revealed by Network Indices.

AbstractIndirect effects arise when one species influences how another species interacts with a third. Pollinator-mediated indirect effects are widespread in many plant communities and are often not restricted to plant species pairs. An analytical framework does not exist yet that allows for the evaluation of indirect effects through shared pollinators in a community context as well as their consequences for plant fitness. We used network indices describing pollinator sharing to assess the extent to which plant species affect and are affected by others in a pollination network from a species-rich dune community. For 23 plant species, we explore how these indices relate to plant fecundity (seeds/flower) over two years. We further linked plant traits and indices to uncover functional aspects of pollinator-mediated indirect interactions. Species frequently visited by shared pollinators showed higher fecundity and exhibited traits that increase pollinator attraction and generalization. Conversely, species whose shared pollinators frequently visited other plants had lower fecundity and more specialized traits. Thus, pollinator sharing benefited some species while others suffered reproductive disadvantages, consistent with competition. The framework developed here uses network tools to advance our understanding of how pollinator-mediated indirect interactions influence a species' relative reproductive success at the community level.

Recipient and donor characteristics govern the hierarchical structure of heterospecific pollen competition networks

Abstract Pollinator sharing can have negative consequences for plant fitness via competition for visits as well as with the arrival of heterospecific pollen. Plant traits and relatedness of donor and recipient species have been suggested to drive the observed variation in plant fitness effects of both processes, but how they shape the structure of interspecific pollen competition networks has been overlooked at the community level. To understand the importance of reproductive traits and relatedness to the impact of heterospecific pollen, we conducted a controlled glasshouse experiment with an artificial co‐flowering community. We performed 2,200 hand pollination crosses by experimentally transferring conspecific pollen alone or with 50% and 100% foreign pollen among 10 species belonging to three different plant families. Relative to conspecific pollen alone, there was a significant reduction in seed set with 50% heterospecific pollen for 67% of the crosses. This effect is driven largely by recipient traits and the interaction between recipient and donor traits under specific circumstances of trait matching. In general, species with shorter styles, smaller stigmas and lower pollen:ovule ratios were more impacted by heterospecific pollen. These traits and their differences among species led to a hierarchical (or transitive) structure of pollen competition with clear winners and losers. However, phylogenetic distance among recipient and donor species did not explain the effects. Synthesis. Our study shows that specific traits and trait combinations between donor and recipient species are important in determining seed production outcomes with heterospecific pollen receipt. Moreover, the differences in traits between species lead to a competitive structure with clear ‘winners’ or ‘losers’ species. The results of this study highlight the importance of specific traits in understanding the position of the competitive hierarchy of the species and the mechanisms underlying heterospecific pollen impacts upon plant reproductive success.

Setting Up an Arabidopsis LED Culture Module that Simulates Plant Neighbor Proximity.

Competition for light between neighboring plants has important consequences for plant fitness and crop productivity. Studies on the molecular mechanisms of plant responses to neighbor proximity have been largely based on the model species Arabidopsis thaliana grown under controlled light environments. These controlled conditions commonly use fluorescent tubes for the main light source for photosynthesis and filtered light form incandescent bulbs to adjust the ratio of red (R) to far-red (FR) radiation. However, both of these types of bulbs are being discontinued and replaced by more efficient sources based on light emitting diodes (LEDs). For that reason, there is a need to evaluate alternative light sources, which can phenocopy the physiological and molecular results obtained with traditional lighting systems. Here we evaluate a custom-made LED culture module that can be used to effectively evaluate shade-avoidance responses, yielding results that, in Arabidopsis, are comparable to those obtained using traditional lighting systems.

Pollination and fruit infestation under artificial light at night:light colour matters

Rapid human population growth and associated urbanization lead to increased artificial illumination of the environment. By changing the natural light–dark cycle, artificial lighting can affect the functioning of natural ecosystems. Many plants rely on insects in order to reproduce but these insects are known to be disturbed by artificial light. Therefore, plant–insect interactions may be affected when exposed to artificial illumination. These effects can potentially be reduced by using different light spectra than white light. We studied the effect of artificial lighting on plant–insect interactions in the Silene latifolia–Hadena bicruris system using a field set-up with four different light treatments: red, green, white and a dark control. We compared the proportion of fertilized flowers and fertilized ovules as well as the infestation of fruits by Hadena bicruris, a pollinating seed predator. We found no difference in the proportion of fertilized flowers among the treatments. The proportion of fruits infested by H. bicruris was however significantly higher under green and white light and a significantly lower proportion of fertilized ovules was found under green light. We show that artificial light with different colours impacts plant–insect interactions differently, with direct consequences for plant fitness.

Annual and perennial high-Andes species have a contrasting freezing-resistance mechanism to cope with summer frosts

In high-mountain habitats, summer frosts have negative consequences for plant fitness, therefore high-mountain plants have developed mechanisms of avoidance and tolerance to cope with freezing temperatures. Various hypotheses have been proposed to explain the prevalence of one freezing-resistance mechanism over another, focusing on thermal conditions without a consensus. We hypothesize that the prevalence of a freezing-resistance mechanism depends on the life-history strategy of the species, and is probably the consequence of a trade-off between growth/reproduction and the cost of the mechanism. Specifically, short-lived annual species should be freezing avoidant, whereas perennial long-lived species should be freezing tolerant. We used thermal analysis to determine the mechanism of freezing resistance of leaves and flowers for 10 annual and 14 perennial herb species from an alpine ecosystem in the Central Chilean Andes. We found that 70% of the annual species, their flowers and leaves were freezing avoidant, indicating that avoidance was their predominant freezing-resistance mechanism. In the case of perennial species, both mechanisms were almost equally represented in flowers and leaves. Overall, our results showed that a species freezing-resistance mechanism depends on its life-history strategy, and that leaves and flowers of single species exhibit the same freezing resistance mechanism, suggesting a common whole plant strategy. Further, freezing resistance strategies were not found to be mutually exclusive. In some cases, a specific combination of phenological, structural, and functional strategies may determine how freezing resistant vegetative and reproductive organs are to freezing during the growing season.

Does the life-history strategy determine the freezing resistance of flowers and leaves of alpine herbaceous species?

In high-mountain habitats, summer frost events can have negative consequences for plant fitness. Despite this, most studies have evaluated the consequences of frosts for vegetative structures of perennial plants, and neither for leaves nor for flowers of annual plants. We hypothesize that the degree of freezing resistance of flowers and leaves of a species depends on its life-history strategy (LHS), and is probably the consequence of a trade-off between growth/reproduction and the cost of the freezing resistance. Specifically, flowers and leaves of short-lived annual species should be less freezing resistant than those of perennial plant species. We compared the freezing resistance of flowers and leaves of 10 annual and 12 perennial plant species from the Andes of central Chile using the electrolyte leakage method. Temperature damage for 50% tissue (LT50) of annual species was − 9.6 °C in flowers and − 11.9 °C in leaves. In perennial species, LT50 was similar in flowers (− 12.3 °C) and leaves (− 12.5 °C). Despite that, these differences were not significant (except the flowers of annual species), we found remarkable differences between LHS when freezing resistance was analyzed species by species. Like this, 58% and 83% of perennial species resist temperatures ≤ − 10 °C in their flowers and leaves, respectively, compared with only 30% and 40% of annual species. Additionally, in most of the species, the freezing resistance of leaves was greater than that of flowers, with this proportion being greater in annual (58%) than in perennial species (43%). Thus, we concluded that the degree of freezing resistance depends on the LHS, such that annual species, which are less freezing resistant than perennial species, have an infrequent occurrence and a distribution restricted to low elevation in high-mountain habitats.

Plant ontogeny determines strength and associated plant fitness consequences of plant-mediated interactions between herbivores and flower visitors.

Plants show ontogenetic variation in growth–defence strategies to maximize reproductive output within a community context. Most work on plant ontogenetic variation in growth–defence trade‐offs has focussed on interactions with antagonistic insect herbivores. Plants respond to herbivore attack with phenotypic changes. Despite the knowledge that plant responses to herbivory affect plant mutualistic interactions with pollinators required for reproduction, indirect interactions between herbivores and pollinators have not been included in the evaluation of how ontogenetic growth–defence trajectories affect plant fitness.In a common garden experiment with the annual Brassica nigra, we investigated whether exposure to various herbivore species on different plant ontogenetic stages (vegetative, bud or flowering stage) affects plant flowering traits, interactions with flower visitors and results in fitness consequences for the plant.Effects of herbivory on flowering plant traits and interactions with flower visitors depended on plant ontogeny. Plant exposure in the vegetative stage to the caterpillar Pieris brassicae and aphid Brevicoryne brassicae led to reduced flowering time and flower production, and resulted in reduced pollinator attraction, pollen beetle colonization, total seed production and seed weight. When plants had buds, infestation by most herbivore species tested reduced flower production and pollen beetle colonization. Pollinator attraction was either increased or reduced. Plants infested in the flowering stage with P. brassicae or Lipaphis erysimi flowered longer, while infestation by any of the herbivore species tested increased the number of flower visits by pollinators.Our results show that the outcome of herbivore–flower visitor interactions in B. nigra is specific for the combination of herbivore species and plant ontogenetic stage. Consequences of herbivory for flowering traits and reproductive output were strongest when plants were attacked early in life. Such differences in selection pressures imposed by herbivores to specific plant ontogenetic stages may drive the evolution of distinct ontogenetic trajectories in growth–defence–reproduction strategies and include indirect interactions between herbivores and flower visitors. Synthesis. Plant ontogeny can define the direct and indirect consequences of herbivory. Our study shows that the ontogenetic stage of plant individuals determined the effects of herbivory on plant flowering traits, interactions with flower visitors and plant fitness.

Quint Rusman.

Quint Rusman.

Dryness, wetness and temporary flooding reduce floral resources of plant communities with adverse consequences for pollinator attraction

Abstract Climate change alters precipitation regimes world‐wide and is regarded as a major threat for pollinators and pollination services. Yet, not much is known on how wetter as well as drier conditions affect food resources for pollinators and pollinator attraction in a multi‐species community context. It is unclear how community shifts under changed hydrological conditions might affect pollinators. This mesocosm study addresses existing research gaps by investigating effects of drought, wetness and temporary flooding on floral resources and pollinator attraction within a plant community, initially including nine insect‐pollinated and six wind‐pollinated species. Floral resources were assessed over three growing seasons. I monitored community descriptors (percentage of herbs and herbaceous species richness, presence of flowering insect‐pollinated individuals), floral traits (flower height, size and weight, floral sugar content of one key species) and proxies for plant fitness (pollinator attraction, seed and flower weight of one key species). Wetness and especially dryness decreased the species richness and biomass proportion of insect‐pollinated herbaceous species and flowering herbaceous plant individuals occurred less frequently. Permanent wetness decreased the floral sugar content of the key species Trifolium pratense. Wet, temporary flooded and especially dry communities were visited by pollinators less often. Active pollinators spent less time within wet and flooded communities, while not at all visiting dry communities. Seed and flower weight of the obligate xenogamous T. pratense in flooded and dry communities decreased, indicating negative consequences for plant fitness caused by a lack of pollination. Synthesis. While dryness had negative effects for floral resources both in terms of community descriptors and floral traits, negative effects of wetness and temporary flooding were mostly caused by a decrease of insect‐pollinated herbaceous species. The study thus indicates that shifts in plant community composition are decisive for a predictive understanding of plant–pollinator interactions under environmental change, but these have been neglected in past research. Changing precipitation patterns will adversely affect floral resources and pollinator attraction in agriculturally used temperate grassland, which might have widespread negative consequences for pollination services and food security in the coming decades.

Variation in Below-to Aboveground Systemic Induction of Glucosinolates Mediates Plant Fitness Consequences under Herbivore Attack.

Plants defend themselves against herbivore attack by constitutively producing toxic secondary metabolites, as well as by inducing them in response to herbivore feeding. Induction of secondary metabolites can cross plant tissue boundaries, such as from root to shoot. However, whether the potential for plants to systemically induce secondary metabolites from roots to shoots shows genetic variability, and thus, potentially, is under selection conferring fitness benefits to the plants is an open question. To address this question, we induced 26 maternal plant families of the wild species Cardamine hirsuta belowground (BG) using the wound-mimicking phytohormone jasmonic acid (JA). We measured resistance against a generalist (Spodoptera littoralis) and a specialist (Pieris brassicae) herbivore species, as well as the production of glucosinolates (GSLs) in plants. We showed that BG induction increased AG resistance against the generalist but not against the specialist, and found substantial plant family-level variation for resistance and GSL induction. We further found that the systemic induction of several GSLs tempered the negative effects of herbivory on total seed set production. Using a widespread natural system, we thus confirm that BG to AG induction has a strong genetic component, and can be under positive selection by increasing plant fitness. We suggest that natural variation in systemic induction is in part dictated by allocation trade-offs between constitutive and inducible GSL production, as well as natural variation in AG and BG herbivore attack in nature.

Daily fluctuations in pollination effectiveness explain higher efficiency of native over exotic bees in Lepechinia floribunda (Lamiaceae).

Despite Stebbins' principle of the most efficient pollinator being proposed decades ago, the most important pollinators are still mainly identified using the frequency of visits to flowers. This shortcoming results in a gap between the characterization of the flower visitors of a plant species and a reliable estimation of the plant fitness consequences of the mutualistic interaction. The performance of a mutualistic visitor depends on its abundance, behaviour, effectiveness (pollen removal and deposition per unit time) and efficiency (seed set per unit time) conditioned by the temporal matching between pollinator activity and temporal patterns of maturation of the sexual functions of flowers. Although there have been recent attempts to provide a conceptual and methodological framework to characterize pollinators' performance, few have combined all key elements of visitors and plants to provide an accurate estimation of pollinators' performance under natural conditions. We complement information on the flower biology and mating system of the sub-shrub Lepechinia floribunda (Lamiaceae) to provide a daily quantitative estimation of performance (effectiveness and efficiency) of the more abundant pollinators, i.e. native bumble-bees (Bombus spp.) and leafcutter bees (Megachile sp.), and the exotic honey-bee (Apis mellifera). Unlike honey-bees or leafcutter bees, native bumble-bees matched the daily pattern of nectar production and stigma receptivity, and showed higher effectiveness and efficiency. Despite the overabundance of honey-bees, visits occurred mainly when stigmas were not receptive, thus reducing the honey-bees' overall performance. Bumble-bees appear to be the most important pollinators and potential historical mediators of reproductive trait evolution in L. floribunda. Because the production of seeds by bumble-bees involved fewer pollen grains for plants and less investment in floral display than honey-bees, contemporary and expected changes in pollinator abundance may affect future L. floribunda floral evolution. If bumble-bees were to be further displaced by anthropogenic disturbance or by competition with honey-bees, their lower efficiency will select for a larger floral display increasing reproductive costs. This scenario may also impose selection to reduce dichogamy to match honey-bee foraging activity.

Cross-seasonal legacy effects of arthropod community on plant fitness in perennial plants.

In perennial plants, interactions with other community members during the vegetative growth phase may influence community assembly during subsequent reproductive years and may influence plant fitness. It is well‐known that plant responses to herbivory affect community assembly within a growing season, but whether plant–herbivore interactions result in legacy effects on community assembly across seasons has received little attention. Moreover, whether plant–herbivore interactions during the vegetative growing season are important in predicting plant fitness directly or indirectly through legacy effects is poorly understood.Here, we tested whether plant–arthropod interactions in the vegetative growing season of perennial wild cabbage plants, Brassica oleracea, result in legacy effects in arthropod community assembly in the subsequent reproductive season and whether legacy effects have plant fitness consequences. We monitored the arthropod community on plants that had been induced with either aphids, caterpillars or no herbivores in a full‐factorial design across 2 years. We quantified the plant traits ‘height’, ‘number of leaves’ and ‘number of flowers’ to understand mechanisms that may mediate legacy effects. We measured seed production in the second year to evaluate plant fitness consequences of legacy effects.Although we did not find community responses to the herbivory treatments, our data show that community composition in the first year leaves a legacy on community composition in a second year: predator community composition co‐varied across years. Structural equation modelling analyses indicated that herbivore communities in the vegetative year correlated with plant performance traits that may have caused a legacy effect on especially predator community assembly in the subsequent reproductive year. Interestingly, the legacy of the herbivore community in the vegetative year predicted plant fitness better than the herbivore community that directly interacted with plants in the reproductive year. Synthesis. Thus, legacy effects of plant–herbivore interactions affect community assembly on perennial plants across growth seasons and these processes may affect plant reproductive success. We argue that plant–herbivore interactions in the vegetative phase as well as in the cross‐seasonal legacy effects caused by plant responses to arthropod herbivory may be important in perennial plant trait evolution such as ontogenetic variation in growth and defence strategies.

Heavy metals in nectar modify behaviors of pollinators and nectar robbers: Consequences for plant fitness

Plants growing in heavy-metal-rich soils can accumulate metals into their nectar. Nectar chemical composition can alter foraging behavior of floral visitors (including pollinators and floral antagonists) and further affect plant reproductive fitness. The role of nectar heavy metals in deterring pollinators (e.g., shortening foraging time) has been recently studied, but their effects on plant reproduction via changes in behaviors of both pollinators and floral antagonists (e.g., nectar robbers) are less understood. We experimentally manipulated four nectar heavy metals (Zn, Cu, Ni, and Pb) in a native ornamental plant, Hosta ensata F. Maekawa, to investigate the effect of nectar metals on plant reproductive success. We also recorded nectar robbing as well as foraging time and visitation rate of pollinators to assess whether nectar metals could alter the behavior of antagonists and mutualists. Although metals in nectar had no significant direct effects on plant reproduction via hand-pollination, we detected their positive indirect effects on components of female fitness mediated by pollinators and nectar robbers. Matching effects on female plant fitness, nectar robbers responded negatively to the presence of metals in nectar, robbing metal-treated flowers less often. Pollinators spent less time foraging on metal-treated flowers, but their visitation rate to metal-treated flowers was significantly higher than to control flowers. Moreover, pollinators removed less nectar from flowers treated with metals. Our results provide the first direct evidence to date that heavy metals in nectar are capable of deterring nectar robbers and modifying pollinator foraging behavior to enhance plant reproductive fitness.

Predator and floral traits change pollinator behaviour, with consequences for plant fitness

1. Flower‐dwelling predators may interfere in plant–pollinator interactions through changes in pollinator behaviour, leading to a reduction in pollination services. Although the context‐dependency of tri‐trophic interactions is often acknowledged, the relative contribution of predator, pollinator and flower traits in shaping pollinator behaviour has been rarely studied simultaneously.2. Pollinators' responses to predation risk were evaluated using artificial spiders on Alstroemeria aurea flowers in order to experimentally test whether predator traits (colour and size) and nectar availability affect their behaviour and plant fitness.3. Regarding the predator's traits, both sizes of artificial spiders used were similarly rejected, but spider colour was detected differently. In particular, red and black artificial spiders were more strongly rejected than the rest. In turn, nectar availability increased the time spent in the inflorescences, yet most pollinators rejected flowers with artificial spiders independently of nectar availability. Furthermore, responses to artificial spider colour and nectar availability varied significantly between dipterans and hymenopterans. Finally, artificial spiders reduced seed set and fruit weight of plants owing to changes in the behaviour of the most efficient pollinators.4. It was shown that poorly studied predator traits such as colour and size may affect pollinator behaviour in different ways among taxa. In addition, feeding necessity in a rich‐resource environment was a weaker selecting force than predation risk. Hence, it is argued here that trade‐offs between predation and flower reward may arise depending on predator detection, relationship between pollinator and predator size, and resource availability in the ecosystem with consequences for plant fitness.