Foliar Herbivory Research Articles

Nectar chemistry modulates the impact of an invasive plant on native pollinators

Summary Invasive species are considered a main driver of pollinator declines, yet the direct effects of invasive alien plants on pollinators are poorly understood. Abundant, invasive plant species can provide a copious nectar resource for native pollinators. However, the nectar of some plants contains secondary compounds, usually associated with defence against herbivores. The impacts of these compounds on pollinators are often unknown. We compared how consumption of grayanotoxin I and III, natural secondary compounds in the nectar of invasive Rhododendron ponticum L., affected three native bee species: a honeybee, (Apis mellifera L.), a solitary mining bee (Andrena carantonica, Pérez) and a bumblebee, (Bombus terrestris, L.). Survival of the solitary bee and the bumblebee species was not affected by either grayanotoxin, but honeybees were ∼20× more likely to die when fed solutions containing grayanotoxin I. Furthermore, solitary bees were deterred from feeding and exhibited malaise behaviours indicative of sublethal toxicity in response to consumption of grayanotoxin I. In contrast, grayanotoxins did not affect bumblebee survival or behaviour, even when bees were subjected to multiple stressors (parasite infection or food stress). Our experiments suggest that while R. ponticum provides abundant floral nectar, it is only available as a food resource to pollinators that tolerate grayanotoxins. Pollinators whose health is negatively affected by grayanotoxins may experience negative impacts from R. ponticum invasion directly (if they consume R. ponticum nectar) or indirectly (if native floral resources are replaced by R. ponticum). Our study makes a novel comparison of the effects of a natural nectar secondary compound on three pollinator species and clearly demonstrates drastic variation in the responses of different key pollinator taxa to a nectar toxin. Our findings are thus in congruence with literature demonstrating the varying effects of invasive plant chemistry on native foliar herbivores, and our work demonstrates that nectar chemistry should be taken into account when determining the impacts of plant invasion for native pollinators.

Quantifying direct vs. indirect effects of nectar robbers on male and female components of plant fitness

Summary Plants interact simultaneously with both mutualists and antagonists. While webs of plant–animal interactions in natural systems can be highly complex, most interactions can be simplified into those that are either direct (mediated through pairwise interactions) or indirect (mediated through third‐party species). Mechanistic studies of the direct and indirect pathways by which foliar herbivores affect plants have been well explored; however, mechanistic explorations of how floral herbivores, such as nectar robbers, affect total plant fitness via direct vs. indirect pathways have received less attention. The goal of this study was to assess the importance of direct vs. pollinator‐mediated indirect effects of a floral antagonist on female and male components of plant fitness. We focused on the hummingbird‐pollinated plant scarlet gilia, Ipomopsis aggregata, which is nectar‐robbed by the bumblebee Bombus occidentalis. Prior studies have found evidence for pollinator‐mediated indirect effects of robbing on female and male components of I. aggregata fitness, but the mechanisms by which these indirect effects occur, and experimental evidence supporting or refuting direct effects of robbing, have been lacking. We found no evidence for direct effects of robbing on plant fitness. Robbers did not act as pollinators of I. aggregata nor did they directly affect seed production by making nectar‐robber holes or removing nectar in hand‐pollinated flowers. Moreover, robbing had no direct effect on pollen production per flower or the ability of pollen from robbed flowers to sire seeds in hand pollinations. However, nectar robbing had indirect effects on plant reproduction mediated through per‐visit pollinator effectiveness at depositing pollen in robbed vs. unrobbed flowers. A simple model of a plant‐robber‐pollinator system suggested that robbing effects in general may occur through more indirect mechanisms when nectar removal by robbers is high relative to nectar replenishment, and that compensation for robbing is then more profitable through the production of additional flowers. Synthesis. Our results highlight the importance of indirect effects in mediating the fitness consequences of species interactions.

Echinacea angustifolia and its specialist ant‐tended aphid: a multi‐year study of manipulated and naturally‐occurring aphid infestation

1. The impact of herbivores on plant fitness depends on multiple ecological mechanisms, including interactions between herbivore guilds.2. This study assessed the effects of a specialist aphid (Aphis echinaceae) on performance and foliar herbivore damage of a long‐lived perennial plant (Echinacea angustifolia) native to the North American tallgrass prairie. A 2‐year field experiment manipulating aphid infestation on 100 plants was compared with concurrent and past observations of unmanipulated plants in the same outdoor experimental plot. Because ants co‐occur with aphids, the experiment tested the combined effects of aphids and ants.3. Neither manipulated nor naturally‐occurring aphid infestations led to measurable declines in plant performance. Results for foliar herbivore damage differed between experimental and observational studies: the occurrence of foliar herbivore damage decreased with aphid infestation in the first year of the experiment and increased with aphid infestation over 5 years in unmanipulated plants.4. While the experimental results concur with other experiments of ant–hemipteran–herbivore relationships, the observational results suggest that ant–aphid interactions do not naturally play a major role in determining patterns of foliar herbivory in this system. This result demonstrates the value of using field observations to interpret the relevance of experimental results.

Insect herbivory, plant-host specialization and tissue partitioning on mid-Mesozoic broadleaved conifers of Northeastern China

Insect-mediated damage was examined on 756 specimens of three broadleaved conifers Podozamites, Lindleycladus and Liaoningocladus, originating from five mid-Mesozoic localities in Northeastern China. These localities are the Late Triassic Yangcaogou Fm. (T3, ca. 205Ma), the latest Middle Jurassic Jiulongshan Fm. (J2, 165Ma), and the mid Early Cretaceous Yixian Fm. (K1, 125Ma). Plant hosts from these three time intervals harbor five functional feeding groups (FFGs) of herbivores and 23 distinctive damage types (DTs), categorized using the widely applied DT system. The DTs were classified into the five FFGs of external foliage feeding (6 DTs), piercing and sucking (5 DTs), oviposition (3 DTs), galling (8 DTs) and leaf mining (1 DT). Damage-type richness and abundance was established for each FFG, encompassing from 10 to 16 DTs for each of the three time intervals examined. For this 80 million-year-long interval, foliar herbivory on broadleaved conifers was transformed from early predominance of external foliage feeding (T3), later replaced by an emphasis on piercing and sucking (J2), followed by bimodal expansion of endophytic interactions from oviposition and leaf mining (K1). This trajectory of herbivore succession indicates that, from T3 to K1, plant–insect associations were transformed from earlier reliance on a greater number of exophytic modes of herbivory to a later, increased variety in endophytic consumption. The transformation also was demonstrated by finer-grained partitioning of food resources and specialization on particular host-plant tissue types. This subdivision of tissue types likely promoted greater dietary saturation of tissue space by functional feeding groups. Possible explanations for these shifts in herbivory include ecological causes, long-term environmental changes, or both. Ecological factors, such as (i), evolution of a more differentiated plant-host spectrum available for consumption; (ii), long-term changes in plant physiognomy and deployment of antiherbivore defenses; (iii), change in herbivore partitioning of plant-host tissues; and (iv), emergence of the parasitoid guild for efficient regulation of insect herbivores. Long-term environmental variables may be linked to these shifts in insect herbivory style.

Thiamethoxam Seed Treatments Have No Impact on Pest Numbers or Yield in Cultivated Sunflowers.

The use of neonicotinoid seed treatments is a nearly ubiquitous practice in sunflower (Helianthus annuus) pest management. Sunflowers have a speciose pest complex, but also harbor a diverse and abundant community of beneficial, nontarget organisms which may be negatively affected by pest management practices. Here, we investigate how the foliar and subterranean arthropod pest communities in sunflower fields were affected by a thiamethoxam seed treatment over three site years (two years on one farm, and another year at an additional field in the second year). Thiamethoxam and its metabolite clothianidin in leaf tissue were quantified throughout the growing season, and yield differences between treatments were measured. Across site years, foliar herbivores and key pests of sunflowers were unaffected by the seed treatment. Likewise, subterranean herbivores were unaffected. Thiamethoxam was measurable in leaf tissue through the R1 plant stage, while its metabolite clothianidin was detected throughout flowering (R6). No difference in sunflower yield was observed between treatments across site years. This research suggests that neonicotinoid seed treatments in sunflowers do not always provide economic benefits to farmers in the form of pest reductions or yield improvements. Future research should focus on sunflower integrated pest management strategies that limit nontarget effects of agrochemicals, while providing greater economic returns to farmers.

Herbivory affects male and female reproductive success differently in dioecious Silene latifolia

AbstractIn dioecious plants, reproductive efforts of male and female plants can be differentially affected by herbivory due to sex‐specific allocation and re‐modulation of resources. Here, we investigate the effects of foliar herbivory by the polyphagous species Spodoptera littoralis (Boisduval) (Lepidoptera: Noctuidae) on male and female reproductive traits of Silene latifolia (Poir.) (Caryophyllaceae). We set up experimental populations of male and female control and Spodoptera‐infested plants and estimated flower morphology and phenology, and plant reproductive success. Herbivory did not affect floral traits involved in visual signalling to pollinators (flower size). Female reproductive success did not differ between Spodoptera‐infested and control plants in terms of seed set and seed germination. On the other hand, Spodoptera‐infested male plants produced fewer flowers than male control plants and maintained fewer flowers during flowering peaks. By experimental reduction in male flower number, we showed that male flower number is directly correlated with male reproductive success (pollen export to female flowers). This strongly suggests that male reproductive success is reduced after herbivory, whereas female reproductive success remains unaffected. The fact that females are more tolerant against herbivory is in accordance with sexual selection theory, which predicts that females, in contrast to males, are not selected for maximum investment in reproductive traits such as floral signalling, and should thus be more flexible in the allocation of resources.

Geographic variation in the protective effects of ants and trichomes in a Neotropical ant–plant

Many ant–plants have pubescent leaves, but the extent to which leaf trichomes enhance the defensive capabilities of ant–plants is poorly known. Here, we examined the influence of foliar trichomes and ants on herbivory in the Neotropical ant–plant Tococa guianensis. We performed a feeding preference test and a field experiment in which we manipulated the presence of ants and trichomes in three distinct populations of T. guianensis; two associated with obligate mutualistic ants and one associated with opportunistic ants. We found that both mutualistic ants and leaf trichomes act as mechanisms of defense against herbivores in T. guianensis. However, the relative importance of each of these two mechanisms was context dependent. The magnitude of the effect of trichomes (as measured by the effect size) was relatively low and varied little among plant populations. The effect of ants on foliar herbivory was up to seven times stronger than the effect of the trichomes. Nevertheless, because the magnitude of ant effects was spatially variable—depending on whether mutualistic ants were present or not, and probably also on the local abundance of herbivores—the relative contribution of ants and trichomes for plant defense was also spatially variable. These findings indicate that mechanical defenses (leaf trichomes) alone cannot replace the biotic defenses of T. guianensis. However, trichomes can play a comparatively more important role in plant defense at times or places where mutualistic ants are not present.

Herbivory Increases Fruit Set in Silene latifolia: A Consequence of Induced Pollinator-Attracting Floral Volatiles?

Although the effect of herbivory on plant reproduction has been investigated in some detail, little is known about how herbivores affect floral signalling. Here, we investigated the effect of foliar herbivory by the African Cotton Leafworm (Spodoptera littoralis) on floral signalling and fruit set in the White Campion (Silene latifolia). We found no effects of herbivory on floral traits involved in visual signalling (flower number, corolla diameter, calyx length, petal length) or in amount of nectar produced. However, Spodoptera-infested plants emitted higher amounts of the two floral volatiles, (Z)-3-hexenyl acetate and β-ocimene, than control plants. Open pollinated, infested plants also were found to produce more fruits than control plants, but only with nocturnal pollinators. Experimental addition of the two induced floral volatiles to non-infested Silene flowers also led to the production of more fruits with nocturnal pollination. This suggests that higher fruit production in herbivore-infested plants was caused by increased nocturnal pollinator attraction, mediated by the induced floral emission of these two volatiles. Our results show that the effects of herbivory on plant reproductive success are not necessarily detrimental, as plants can compensate herbivory with increased investment in pollinator attraction.

The effects of the presence of Bt-transgenic oilseed rape in wild mustard populations on the rhizosphere nematode and microbial communities

The adventitious presence of transgenic crops in wild plant populations is of ecological and regulatory concern. In this context, their effects on non-target, below-ground organisms are not well understood. Here, we introduced, at various frequencies, Bt-transgenic oilseed rape (OSR, Brassica napus) into wild mustard (Brassica juncea) populations in the presence and absence of the target herbivore (Plutella xylostella). The impacts on soil nematode and microbial communities were assessed in this system. There were no significant changes on the number of nematode genera and abundance in proportions of OSR with mustard. Nonetheless, the Shannon–Wiener and Pielou evenness index was lowest in plant stands containing 50% of Bt-transgenic OSR. Among treatments, there was no significant variation for culturable soil microbes. There was a positive association between foliar herbivory and the abundance of plant parasitic (PP) and cp-3 nematodes, whereas there was no association between herbivory and soil microbial populations. There was no direct effects of the presence of Bt-transgenic OSR in wild mustard populations on the rhizosphere nematode and microbial communities, whereas its indirect effects via aboveground herbivory might be important to consider for biosafety assessments.

Plant herbivory responses through changes in leaf quality have no effect on subsequent leaf-litter decomposition in a neotropical rain forest tree community.

It is commonly accepted that plant responses to foliar herbivory (e.g. plant defenses) can influence subsequent leaf-litter decomposability in soil. While several studies have assessed the herbivory-decomposability relationship among different plant species, experimental tests at the intra-specific level are rare, although critical for a mechanistic understanding of how herbivores affect decomposition and its consequences at the ecosystem scale. Using 17 tree species from the Yasuní National Park, Ecuadorian Amazonia, and applying three different herbivore damage treatments, we experimentally tested whether the plant intra-specific responses to herbivory, through changes in leaf quality, affect subsequent leaf-litter decomposition in soil. We found no effects of herbivore damage on the subsequent decomposition of leaf litter within any of the species tested. Our results suggest that leaf traits affecting herbivory are different from those influencing decomposition. Herbivore damage showed much higher intra-specific than inter-specific variability, while we observed the opposite for decomposition. Our findings support the idea that interactions between consumers and their resources are controlled by different factors for the green and the brown food-webs in tropical forests, where herbivory may not necessarily generate any direct positive or negative feedbacks for nutrient cycling.

Foliar anti-herbivore defenses in Qualea multiflora Mart. (Vochysiaceae): Changing strategy according to leaf development

Knowledge about plant defense systems against herbivory is crucial to understanding trophic relationships in terrestrial ecosystems. Distinct defenses may act either alone or in combination during foliar sprouting. In our study, we explored the effect of the three most conspicuous defenses found in the leaves of Qualea multiflora (Vochysiaceae) during foliar sprouting, which are: (1) the presence of trichomes, (2) foliar toughness and (3) activity of extrafloral nectaries (EFNs). In addition, we quantified foliar herbivory at different leaf stages. We determined that the three defenses evaluated vary in effectiveness during leaf development. The number of trichomes was higher during early leaf development, toughness at the end, and the activity of EFNs (biotic defense) during the middle period. Foliar herbivory was similar at different leaf ages. The results indicate that Q. multiflora synchronizes its foliar defenses in order to optimize performance in anti-herbivory protection over time.

Abrupt changes in invertebrate herbivory on woody plants at the forest–tundra ecotone

Invertebrate herbivores, insects in particular, play important functional roles in terrestrial ecosystems. At background (non-outbreak) densities, they consume 5–15 % of woody plant foliage in forests across the planet. At the same time, almost nothing is known about the levels of insect herbivory in Arctic tundra. To test the hypothesis that the amount of plant biomass lost to insects in tundra is substantially less than in subarctic forests, in 2013, we explored foliar herbivory in woody plants at three sites in the Arctic tundra and four sites in the subarctic forests of European Russia. A vast majority of foliar damage was imposed by externally feeding defoliators. In forests, defoliators damaged three times more leaves and consumed eight times more leaf area than in the tundra. No miners were found in the tundra, and gallers affected five times less leaf area in the tundra compared with forests. An abrupt decrease in loss of woody plant foliage to insects between subarctic forests and tundra (from 4.34 to 0.56 %) supports the existence of a latitudinal gradient in herbivory in terrestrial ecosystems. More studies are needed to predict how tundra plants, which have been historically exposed to low levels of insect herbivory, will cope with the increased levels of damage that are expected to occur due to climate-driven range expansion and increased abundances of plant-feeding insects.

Anthocyanin production in the hyperaccumulator plant Noccaea caerulescens in response to herbivory and zinc stress

In the current study, potential involvement of anthocyanins in the defence of the Zn hyperaccumulator plant, Noccaea caerulescens, was investigated in two independent glasshouse experiments. Plants were grown in compost amended with a range of Zn concentrations. In the first experiment, foliar herbivory was simulated by clipping the shoots with scissors, whilst in second experiment, plants were subjected to natural herbivory by thrips (Frankliniella occidentalis). In both experiments, foliar Zn concentrations were approximately twice as high as that applied to the soil, indicating accumulation occurred. In the clipping experiment, foliar anthocyanin concentrations decreased by 22–45 % with increasing soil Zn amendments and clipping increased anthocyanin concentrations by 17–24 % relative to unclipped plants. In a thrips feeding experiment, Zn treatment had little effect on anthocyanin content although there was a trend towards increased anthocyanin production in plants grown at the lower concentrations of Zn. Anthocyanin concentration increased by 49–52 % in N. caerulescens leaves subjected to thrip attack compared to that of thrip-free shoots. Thus, anthocyanins increased as a result of foliar damage (manual clipping and natural herbivory) and generally decreased with enhanced Zn concentrations. Foliar glucosinolate concentrations were higher in ungrazed leaves than in grazed leaves of the same thrip-infested plants, whilst anthocyanin production was higher in the grazed leaves. The possibility of an interaction between glucosinolate and anthocyanin production resulting from herbivory is discussed.

Genetic variation in plant below-ground response to elevated CO2 and two herbivore species

It is unclear how changing atmospheric conditions, including rising carbon dioxide concentration, influence interactions between above and below-ground systems and if intraspecific variation exists in this response. We assessed interactive effects of atmospheric CO2 concentration, above-ground herbivory, and plant genotype on root traits and mycorrhizal associations. Plants from five families of Asclepias syriaca, a perennial forb, were grown under ambient and elevated atmospheric CO2 concentrations. Foliar herbivory by either lepidopteran caterpillars or phloem-feeding aphids was imposed. Mycorrhizal colonization, below-ground biomass, root biomass, and secondary defensive chemistry in roots were quantified. We observed substantial genetic variation among A. syriaca families in their mycorrhizal colonization levels in response to elevated CO2 and herbivory treatments. Elevated CO2 treatment increased root biomass in all genetic families, whereas foliar herbivory tended to decrease root biomass. Root cardenolide concentration and composition varied greatly among plant families, and elevated CO2 treatment increased root cardenolides in two of the five plant families. Moreover, herbivores differentially affected the composition of cardenolides expressed below ground. Increased atmospheric CO2 has the potential to influence interactions among plants, herbivores and mycorrhizal fungi and intraspecific variation suggests that such interactions can evolve.

Variation in the Outcomes of an Ant-Plant System: Fire and Leaf Fungus Infection Reduce Benefits to Plants with Extrafloral Nectaries

Interactions between species are evolutionary malleable and may suffer changes in small timescales. Environmental disturbances, such as fire, can deeply affect species interactions, but how they influence the outcome of a mutualistic interaction has yet to be studied. In order to test the hypothesis that an environmental disturbance, in this case fire, may produce differences in the outcome of the association of ants with the extrafloral-nectaries-bearing plant Qualea multiflora Mart. (Myrtales: Vochysiaceae), a previous study was replicated, but this time after fire incidence, at the same study site and with the same plant species. Eight ant species visited Q. multiflora, and the most abundant genera were Crematogaster, Cephalotes, and Camponotus. Herbivores were found in branches with and without ants with no statistical difference, but foliar herbivory was always higher in branchs where ants were absent. Leaves were infested by fungi, and fungi spots were higher in branches where ants were present. Compared to the previous study, it was clearly observed that ant benefits to Q. multiflora varied over time. The most common ant species still protected leaves against chewing herbivores, but a new kind of leaf damage appeared, namely fungi spots. Data also support that ants may be acting as vectors of fungi spores on plants, as ant visited branches had higher fungus incidence than non-visited branches. Fire is a major source of disturbance in tropical savannas, and we suggest that it can cause strong variation in the outcomes of interactions between ants and plants with extrafloral nectaries in the Brazilian tropical savanna.

Increase of fast nutrient cycling in grassland microcosms through insect herbivory depends on plant functional composition and species diversity

Nutrient cycling in terrestrial ecosystems is affected by various factors such as plant diversity and insect herbivory. While several studies suggest insect herbivory to depend on plant diversity, their interacting effect on nutrient cycling is unclear.In a greenhouse experiment with grassland microcosms of one to six plant species of two functional groups (grasses and legumes), we tested the influence of plant species richness (diversity) and functional composition on plant community biomass production, insect foliar herbivory, soil microbial biomass, and nutrient concentrations in throughfall. To manipulate herbivory, zero, three or six generalist grasshoppers (Chorthippus parallelus) were added to the plant communities.Increasing plant species richness increased shoot biomass and grasshopper performance, without significantly affecting root biomass or insect herbivory. Plant functional composition affected all of these parameters, e.g. legume communities showed the highest shoot biomass, the lowest grasshopper performance and suffered the least herbivory. Nutrient concentrations (dissolved mineral N, PO4‐P, SO4‐S) and pH in throughfall increased with herbivory. PO4‐P and pH increases were positively affected by plant diversity, especially under high herbivore pressure. Plant functional composition affected several throughfall variables, sometimes fully explaining diversity effects. Increasing plant diversity tended to increase soil microbial biomass, but only under high herbivore pressure. Faeces quantities strongly correlated with changes in pH and PO4‐P; frass may therefore be an important driver of throughfall pH and a main source of PO4‐P released from living plants. Our results indicate that insect herbivory may significantly influence fast nutrient cycling processes in natural communities, particularly so in managed grasslands.

Herbivory and floral signaling: phenotypic plasticity and tradeoffs between reproduction and indirect defense

Plant defense against herbivores may compromise attraction of mutualists, yet information remains limited about the mechanisms underlying such signaling tradeoffs. Here, we investigated the effects of foliar herbivory by two herbivore species on defense compounds, floral signaling, pollinator and parasitoid attraction, and seed production. Herbivory generally reduced the quantity of many floral volatile organic compounds VOCs) in Brassica rapa. By contrast, floral color, flower diameter, and plant height remained unaffected. The decreased amounts of floral volatiles led to reduced attractiveness of flowers to pollinators, but increased the attractiveness of herbivore-infested plants to parasitoids. Plants infested with the native butterfly Pieris brassicae produced more flowers during early flowering, effectively compensating for the lower olfactory attractiveness. Herbivory by the invasive Spodoptera littoralis increased the amounts of glucobrassicanapin, and led to delayed flowering. These plants tended to attract fewer pollinators and to produce fewer seeds. Our study indicates a tradeoff between pollinator attraction and indirect defense (parasitoid attraction), which can be mitigated by reduced floral VOC emission and production of more early flowers. We suggest that this compensatory mechanism is specific to plant-herbivore associations with a coevolutionary history.

Sequential effects of root and foliar herbivory on aboveground and belowground induced plant defense responses and insect performance

Plants are often simultaneously or sequentially attacked by multiple herbivores and changes in host plants induced by one herbivore can influence the performance of other herbivores. We examined how sequential feeding on the plant Plantago lanceolata by the aboveground herbivore Spodoptera exigua and the belowground herbivore Agriotes lineatus influences plant defense and the performance of both insects. Belowground herbivory caused a reduction in the food consumption by the aboveground herbivore independent of whether it was initiated before, at the same time, or after that of the aboveground herbivore. By contrast, aboveground herbivory did not significantly affect belowground herbivore performance, but significantly reduced the performance of later arriving aboveground conspecifics. Interestingly, belowground herbivores negated negative effects of aboveground herbivores on consumption efficiency of their later arriving conspecifics, but only if the belowground herbivores were introduced simultaneously with the early arriving aboveground herbivores. Aboveground-belowground interactions could only partly be explained by induced changes in an important class of defense compounds, iridoid glycosides (IGs). Belowground herbivory caused a reduction in IGs in roots without affecting shoot levels, while aboveground herbivory increased IG levels in roots in the short term (4 days) but only in the shoots in the longer term (17 days). We conclude that the sequence of aboveground and belowground herbivory is important in interactions between aboveground and belowground herbivores and that knowledge on the timing of exposure is essential to predict outcomes of aboveground-belowground interactions.

Continued foliar herbivory by the red spider mite Tetranychus macfarlenei on Plumbago zeylanica severely reduces the levels of a medicinally important metabolite in the roots

Red spider mite, Tetranychus macfarlanei, was identified as a causative parasite for Plumbago zeylanica in India. Present study was undertaken to evaluate the detrimental role of T. macfarlanei on P. zeylanica. The relative change in biomass, stem length, stem diameter, and leaf number was significantly reduced by mite infestation as compared with uninfected plants. A significant reduction of chlorophyll a, chlorophyll b, and total chlorophyll was observed in mite-infested plants. Mite infestation significantly hampered the normal physiological process of the plant, namely, CO2 assimilation, stomatal conductance, electrolyte leakage, cell membrane injury, and carboxylation efficiency. A significant increase in tissue H2O2 and lipid peroxidation coupled with depletion of antioxidant enzymes, namely superoxide dismutase and catalase indicated augmented oxidative stress associated with biotic interaction. Finally, a significant reduction of root-specific secondary metabolite, plumbagin, production highlighted the potential damage of T. macfarlanei in P. zeylanica. The experimental outcome could conclude that the aboveground symptoms due to mite infestation could also exert profound detrimental effect in the belowground tissues.

Drought and Root Herbivory Interact to Alter the Response of Above-Ground Parasitoids to Aphid Infested Plants and Associated Plant Volatile Signals

Multitrophic interactions are likely to be altered by climate change but there is little empirical evidence relating the responses of herbivores and parasitoids to abiotic factors. Here we investigated the effects of drought on an above/below-ground system comprising a generalist and a specialist aphid species (foliar herbivores), their parasitoids, and a dipteran species (root herbivore).We tested the hypotheses that: (1) high levels of drought stress and below-ground herbivory interact to reduce the performance of parasitoids developing in aphids; (2) drought stress and root herbivory change the profile of volatile organic chemicals (VOCs) emitted by the host plant; (3) parasitoids avoid ovipositing in aphids feeding on plants under drought stress and root herbivory. We examined the effect of drought, with and without root herbivory, on the olfactory response of parasitoids (preference), plant volatile emissions, parasitism success (performance), and the effect of drought on root herbivory. Under drought, percentage parasitism of aphids was reduced by about 40–55% compared with well watered plants. There was a significant interaction between drought and root herbivory on the efficacy of the two parasitoid species, drought stress partially reversing the negative effect of root herbivory on percent parasitism. In the absence of drought, root herbivory significantly reduced the performance (e.g. fecundity) of both parasitoid species developing in foliar herbivores. Plant emissions of VOCs were reduced by drought and root herbivores, and in olfactometer experiments parasitoids preferred the odour from well-watered plants compared with other treatments. The present work demonstrates that drought stress can change the outcome of interactions between herbivores feeding above- and below-ground and their parasitoids, mediated by changes in the chemical signals from plants to parasitoids. This provides a new insight into how the structure of terrestrial communities may be affected by drought.