How do herbivorous insects respond to drought stress in trees?

Increased frequency and severity of drought, as a result of climate change, is expected to drive critical changes in plant–insect interactions that may elevate rates of tree mortality. The mechanisms that link water stress in plants to insect performance are not well understood. Here, we build on previous reviews and develop a framework that incorporates the severity and longevity of drought and captures the plant physiological adjustments that follow moderate and severe drought. Using this framework, we investigate in greater depth how insect performance responds to increasing drought severity for: (i) different feeding guilds; (ii) flush feeders and senescence feeders; (iii) specialist and generalist insect herbivores; and (iv) temperate versus tropical forest communities. We outline how intermittent and moderate drought can result in increases of carbon‐based and nitrogen‐based chemical defences, whereas long and severe drought events can result in decreases in plant secondary defence compounds. We predict that different herbivore feeding guilds will show different but predictable responses to drought events, with most feeding guilds being negatively affected by water stress, with the exception of wood borers and bark beetles during severe drought and sap‐sucking insects and leaf miners during moderate and intermittent drought. Time of feeding and host specificity are important considerations. Some insects, regardless of feeding guild, prefer to feed on younger tissues from leaf flush, whereas others are adapted to feed on senescing tissues of severely stressed trees. We argue that moderate water stress could benefit specialist insect herbivores, while generalists might prefer severe drought conditions. Current evidence suggests that insect outbreaks are shorter and more spatially restricted in tropical than in temperate forests. We suggest that future research on the impact of drought on insect communities should include (i) assessing how drought‐induced changes in various plant traits, such as secondary compound concentrations and leaf water potential, affect herbivores; (ii) food web implications for other insects and those that feed on them; and (iii) interactions between the effects on insects of increasing drought and other forms of environmental change including rising temperatures and CO₂ levels. There is a need for larger, temperate and tropical forest‐scale drought experiments to look at herbivorous insect responses and their role in tree death.     

The abundance of invertebrate herbivores in relation to the availability of nitrogen in stressed food plants

Summary It has previously been postulated that when plants are stressed by certain changes in patterns of weather they become a better source of food for invertebrate herbivores because this stress causes an increase in the amount of nitrogen available in their tissues for young herbivores feeding on them. And this may cause outbreaks of such phytophagous invertebrates.Evidence is now presented that a similar physiological mechanism appears to operate when a wide variety of apparently unrelated environmental factors impinge on plants or parts of plants in such a way as to perturb their metabolism. A broken branch, lightning strike, fire, nutrient deficiencies or an otherwise adverse site; all may have this effect. With the advent of modern man the available agencies increase and diversify to include pesticides, irradiation and air pollutants.One common metabolic response by plants to all such agents impinging on them seems to be equivalent to that found in senescing plant tissues — the breakdown and mobilization of nitrogen in soluble form away from the senescing/stressed tissues. Young herbivores which chance to feed on such stressed/senescing tissues have a greater and more readily available supply of nitrogen in their food than they would have had if feeding on unstressed plants. As a result many more of them survive, and there is an increase in abundance of their kind. Such increases may be quite localised and short-lived or more widespread and persistent, depending on the extent and duration of the stress experienced by the plants. And in the face of this improved nutrition and survival of the very young, predators and parasites seem to have only a minor influence on subsequent changes in abundance of their herbivorous prey.Another effect of increased mobilization of nitrogen in stressed plants is an increase in the quantity of the seed that they set. This has led to the conclusion that increased abundance of some species of birds at such times is due to a greater supply of seeds as winter food for recent fledglings. But it may be that the increased abundance is due to the synchronous increase in phytophagous insects providing a richer source of protein food for laying hens and growing nestlings.     

Host‐plant genotype mediates supply and demand of animal food in an omnivorous insect

1. Omnivorous predators can protect plants from herbivores, but may also consume plant material themselves. Omnivores and their purely herbivorous prey have previously been thought to respond similarly to host‐plant quality. However, different responses of omnivores and herbivores to their shared host plants may influence the fitness, trophic identity, and population dynamics of the omnivores. 2. The aim of the present study was to show that an omnivorous heteropteran (Anthocoris nemorum L.) and two strictly herbivorous prey species respond differently to different genotypes of their shared host plant, Salix. Some plant genotypes were sub‐optimal for the omnivore, although suitable for the herbivores, and vice versa. 3. The contrasting patterns of plant suitability for the omnivore and the herbivores highlight an interaction between plant genotype and omnivores' access to animal food. Plant genotypes that were sub‐optimal for the omnivore when herbivores were experimentally excluded became the best host plants when herbivores were present, as in the latter situation additional prey became available. By contrast, the quality of plant genotypes that were intrinsically suitable for omnivores, did not improve when herbivores were present as these plant genotypes were intrinsically sub‐optimal for herbivores, thus providing omnivores with almost no additional animal food. 4. The differential responses of omnivores and their prey to the same host‐plant genotypes should allow omnivores to colonise sub‐optimal host plants in their capacity as predators, and to colonise more suitable host plants in their capacity as herbivores. It may thus be difficult for Salix to escape herbivory entirely, as it will rarely be unsuitable for both omnivores and pure herbivores at the same time.     

Drought stress drives intraspecific choice of food plants by Atta leaf‐cutting ants

Leaf‐cutting ants (LCA) are polyphagous and dominant herbivores throughout the Neotropics that carefully select plant individuals or plant parts to feed their symbiotic fungus. Although many species‐specific leaf traits have been identified as criteria for the choice of food plants, the factors driving intraspecific herbivory patterns in LCA are less well studied. Herein, we evaluate whether or not drought‐stressed native plants are a preferred food source using free‐living colonies of two leaf‐cutting ants, Atta sexdens L. (Hymenoptera: Formicidae: Attini), in combination with five plant species, Ocotea glomerata Nees (Lauraceae), Lecythis lurida S. A. Mori (Lecythidaceae), Miconia prasina DC (Melastomataceae), Tovomita brevistaminea Engl. (Clusiaceae), and Tapirira guianensis Aubl. (Anacardiaceae), and Atta cephalotes L., in combination with two plant species, O. glomerata and Licania tomentosa Benth. (Chrysobalanaceae). In dual‐choice bioassays, ants removed about three times more leaf area from drought‐stressed plants compared to control plants. Both leaf‐cutting ant species consistently preferred drought‐stressed plants for all species tested, except T. guianensis. The mean acceptability index – expressing the preference for one of two options on a scale of 0 to 1 – of drought‐stressed plants ranged from 0.65 to 0.86 across plant species, and the preference did not differ significantly among the tested plant species. Our results suggest that selection of drought‐stressed individuals is a general feature of food plant choice by leaf‐cutting ants irrespective of ant or plant species. As human‐modified forest assemblages across the Neotropics are increasingly prone to drought stress, the documented preference of Atta for drought‐stressed plants may have tangible ecological implications.     

Effects of native and exotic range‐expanding plant species on taxonomic and functional composition of nematodes in the soil food web

Due to climate warming, many plant species shift ranges towards higher latitudes. Plants can disperse faster than most soil biota, however, little is known about how range‐expanding plants in the new range will establish interactions with the resident soil food web. In this paper we examine how the soil nematode community from the new range responds to range‐expanding plant species compared to related natives. We focused on nematodes, because they are important components in various trophic levels of the soil food web, some feeding on plant roots, others on microbes or on invertebrates. We expected that range expanding plant species have fewer root‐feeding nematodes, as predicted by enemy release hypothesis. We therefore expected that range expanders affect the taxonomic and functional composition of the nematode community, but that these effects would diminish with increasing trophic position of nematodes in the soil food web. We exposed six range expanders (including three intercontinental exotics) and nine related native plant species to soil from the invaded range and show that range expanders on average had fewer root‐feeding nematodes per unit root biomass than related natives. The range expanders showed resistance against rather than tolerance for root‐feeding nematodes from the new range. On the other hand, the overall taxonomic and functional nematode community composition was influenced by plant species rather than by plant origin. The plant identity effects declined with trophic position of nematodes in the soil food web, as plant feeders were influenced more than other feeding guilds. We conclude that range‐expanding plant species can have fewer root‐feeding nematodes per unit root biomass than related natives, but that the taxonomic and functional nematode community composition is determined more by plant identity than by plant origin. Plant species identity effects decreased with trophic position of nematodes in the soil food web.     

Community‐wide effects of below‐ground rhizobia on above‐ground arthropods

1. Plants take nutrients for their growth and reproduction from not only soil but also symbiotic microbes in the rhizosphere, and therefore below‐ground microbes may indirectly influence the above‐ground arthropod community through changes in the quality and quantity of plants. 2. Rhizobia are root‐nodulating bacteria that provide NH₄⁺ to legume plants. We examined bottom‐up effects of rhizobia on the community properties of the arthropods on host plants, using a root‐nodulating soybean strain (R+) and a non‐nodulating strain (R?) in a common garden. 3. R+ plants grew larger and produced a greater number of leaves than R? plants. We observed 28 species of herbivores and three taxonomic groups of predators on R+ and R? plants. The herbivorous species were classified into sap feeders (12 species) and chewers (16 species). 4. The species richness of overall herbivores, sap feeders, and chewers on R+ plants was greater than that on R? plants. Rhizobia positively affected the abundance of chewers. 5. The community composition of herbivores was significantly different between R? and R+ plants, although species diversity and evenness did not differ. 6. Rhizobia‐induced bottom‐up effects were transmitted to the third trophic level. The abundance, taxonomic richness, and diversity of the predators on R+ plants were greater but evenness was lower than those on R? plants. The community composition of predators was not affected by rhizobia. 7. These results indicate that the below‐ground microbes initiated bottom‐up effects on above‐ground herbivores and predators through trophic levels.     

Predicting novel trophic interactions in a non‐native world

Humans are altering the global distributional ranges of plants, while their co‐evolved herbivores are frequently left behind. Native herbivores often colonise non‐native plants, potentially reducing invasion success or causing economic loss to introduced agricultural crops. We developed a predictive model to forecast novel interactions and verified it with a data set containing hundreds of observed novel plant–insect interactions. Using a food network of 900 native European butterfly and moth species and 1944 native plants, we built an herbivore host‐use model. By extrapolating host use from the native herbivore–plant food network, we accurately forecasted the observed novel use of 459 non‐native plant species by native herbivores. Patterns that governed herbivore host breadth on co‐evolved native plants were equally important in determining non‐native hosts. Our results make the forecasting of novel herbivore communities feasible in order to better understand the fate and impact of introduced plants.     

Increased cytokinin levels in transgenic PSAG12–IPT tobacco plants have large direct and indirect effects on leaf senescence, photosynthesis and N partitioning

We studied the impact of delayed leaf senescence on the functioning of plants growing under conditions of nitrogen remobilization. Interactions between cytokinin metabolism, Rubisco and protein levels, photosynthesis and plant nitrogen partitioning were studied in transgenic tobacco (Nicotiana tabacum L.) plants showing delayed leaf senescence through a novel type of enhanced cytokinin syn‐thesis, i.e. targeted to senescing leaves and negatively auto‐regulated (P_SAG12–IPT), thus preventing developmental abnormalities. Plants were grown with growth‐limiting nitrogen supply. Compared to the wild‐type, endogenous levels of free zeatin (Z)‐ and Z riboside (ZR)‐type cytokinins were increased up to 15‐fold (total ZR up to 100‐fold) in senescing leaves, and twofold in younger leaves of P_SAG12–IPT. In these plants, the senescence‐associated declines in N, protein and Rubisco levels and photosynthesis rates were delayed. Senescing leaves accumulated more (¹⁵N‐labelled) N than younger leaves, associated with reduced shoot N accumulation (–60%) and a partially inverted canopy N profile in P_SAG12–IPT plants. While root N accumulation was not affected, N translocation to non‐senescing leaves was progressively reduced. We discuss potential consequences of these modified sink–source relations, associated with delayed leaf senescence, for plant productivity and the efficiency of utilization of light and minerals.     

How does the host‐specialized aphid deal with food deficiency?

Aphis gossypii Glover shows obvious host specialization, with cucurbit‐ and cotton‐specialized biotypes or host races in many regions. Because its annual natal host crops senesce earlier the cucurbit‐specialized biotype may suffer food deficiency. The method this biotype uses to overcome this challenge is still poorly understood. In order to understand the potential of the cucurbit‐specialized biotype aphids in host shift and usage, the performance of this biotype on cotton (Gossypium hirsutum), a common but poor quality host plant, was explored in this study. The cucurbit‐specialized aphids could establish populations on cotton only when these plants had at least nine leaves, and subsequent populations developed rather slowly. The presence of whitefly populations on cotton improved the success rate of cucurbit‐specialized aphids. The cucurbit‐specialized aphids were mainly distributed on the older leaves of cotton, with only a few settling on the upper leaves. The cucurbit‐specialized aphids reared on cotton for 40, 54 and 61 days still maintained strong preference for their natal host plant, cucumber (Cucumis sativus), rather than cotton, and their net reproductive rates and intrinsic rates of natural increase were dramatically lower when they were transferred onto new six‐leaf cotton plants or detached leaves. Therefore, we concluded that the cucurbit‐specialized aphids have the potential to utilize mature or whitefly‐stressed cotton plants, but that this feeding experience on cotton did not alter their specialization for cucurbits. Some cotton plants could act as a temporary host for the cucurbit‐specialized aphids to overcome food deficiency arising from senescing cucurbits.     

Influence of corn on stink bugs (Heteroptera: Pentatomidae) in subsequent crops

In southeastern United States farmscapes, corn, Zea mays L., is often closely associated with peanut, (Arachis hypogaea L.), cotton, (Gossypium hirsutum L.), or both. The objective of this 3-yr on-farm study was to examine the influence of corn on stink bugs (Heteroptera: Pentatomidae), Nezara viridula (L.), and Euschistus servus (Say), in subsequent crops in these farmscapes. Adults of both stink bug species entered corn first, and seasonal occurrence of stink bug eggs, nymphs, and adults indicated that corn was a suitable host plant for adult survival and nymphal development to adults. Stink bug females generally oviposited on cotton or peanut near the interface, or common boundary, of the farmscape before senescence of corn, availability of a new food, or both. Adult stink bugs dispersed from crop to crop at the interface of a farmscape in response to senescence of corn, availability of new food, or both. In corn-cotton farmscapes, adult stink bugs dispersed from senescing corn into cotton to feed on bolls (fruit). In corn-peanut farmscapes, adult stink bugs dispersed from senescing corn into peanut, which apparently played a role in nymphal development in these farmscapes. In the corn-cotton-peanut farmscape, stink bug nymphs and adults dispersed from peanut into cotton in response to newly available food, not senescence of peanut. Stink bug dispersal into cotton resulted in severe boll damage. In conclusion, N. viridula and E. servus are generalist feeders that exhibit edge-mediated dispersal from corn into subsequent adjacent crops in corn-cotton, corn-peanut, and corn-peanut-cotton farmscapes to take advantage of suitable resources available in time and space for oviposition, nymphal development, and adult survival. Management strategies for crops in this region need to be designed to break the cycle of stink bug production, dispersal, and expansion by exploiting their edge-mediated movement and host plant preferences.     

Protein carbonylation during natural leaf senescence in winter wheat,as probed by fluorescein‐5‐thiosemicarbazide

Leaf senescence is characterised by a massive degradation of proteins in order to recycle nitrogen to other parts of the plant, such as younger leaves or developing grain/seed. Protein degradation during leaf senescence is a highly regulated process and it is suggested that proteins to be degraded are marked by an oxidative modification (carbonylation) that makes them more susceptible to proteolysis. However, there is as yet no evidence of an increase in protein carbonylation level during natural leaf senescence. The aim of our study was thus to monitor protein carbonylation level during the process of natural senescence in the flag leaf of field‐grown winter wheat plants. For this purpose, we adapted a fluorescence‐based method using fluorescein‐5‐thiosemicarbazide (FTC) as a probe for detecting protein carbonyl derivatives. As used for the first time on plant material, this method allowed the detection of both quantitative and qualitative modifications in protein carbonyl levels during the last stages of wheat flag leaf development. The method described herein represents a convenient, sensitive and reproducible alternative to the commonly used 2,4‐dinitrophenylhydrazine (DNPH)‐based method. In addition, our analysis revealed changes in protein carbonylation level during leaf development that were associated with qualitative changes in protein abundance and carbonylation profiles. In the senescing flag leaf, protein carbonylation increased concomitantly with a stimulation of endoproteolytic activity and a decrease in protein content, which supports the suggested relationship between protein oxidation and proteolysis during natural leaf senescence.     

Elongated hindguts in desert‐living dung beetles (Scarabaeidae: Scarabaeinae) feeding on dry dung pellets or plant litter

Most adult dung beetles (Scarabaeidae: Scarabaeinae) feed on fresh, wet dung of larger herbivorous or omnivorous mammals. As refractory plant fragments are selected out before ingestion, the food is presumed easily digestible. However, members of the desert‐living scarabaeine genus Pachysoma (probably evolved from an ancestor closely related to the wet‐dung feeding genus Scarabaeus) select dry dung pellets and/or plant litter. Thus, they ingest a much higher proportion of structural plant material, which nevertheless appears to be digested rather efficiently. This study investigates morphological modifications of the gut for this digestion in adults of eight Pachysoma species, both pellet and litter feeders. To ascertain hypothesized ancestral conditions, four fresh‐dung feeding Scarabaeus species were also examined. The latter have the usual dung beetle gut consisting of a long, simple midgut, followed by an equally simple, much shorter hindgut of the same width. Lengths of midguts (M) and hindguts (H) divided by body length (B) for comparison between species of different size are: 4.9–6.3 (M/B) and 0.7–0.8 (H/B), which is normal for dung feeders. In Pachysoma, lengths are 6.3–6.5 (M/B) and 1.0–1.4 (H/B) in pellet feeders, and 4.4–5.0 (M/B) and 2.0–2.5 (H/B) for litter feeders. Hindguts are still morphologically undifferentiated and of midgut width, but clearly longer, particularly in litter feeders. Presumably, plant fragments in the food are digested, at least partly, in the hindgut. If so, the morphological adaptation is unusual: simple elongation rather than the expansion of part of the hindgut, as found in several other plant‐ or detritus‐feeding scarabaeids. J. Morphol. 2013. © 2013 Wiley Periodicals, Inc.     

The form of direct interspecific competition modifies secondary extinction patterns in multi‐trophic food webs

Forcibly removing species from ecosystems has important consequences for the remaining assemblage, leading to changes in community structure, ecosystem functioning and secondary (cascading) extinctions. One key question that has arisen from single‐ and multi‐trophic ecosystem models is whether the secondary extinctions that occur within competitive communities (guilds) are also important in multi‐trophic ecosystems? The loss of consumer–resource links obviously causes secondary extinction of specialist consumers (topological extinctions), but the importance of secondary extinctions in multi‐trophic food webs driven by direct competitive exclusion remains unknown. Here I disentangle the effects of extinctions driven by basal competitive exclusion from those caused by trophic interactions in a multi‐trophic ecosystem (basal producers, intermediate and top consumers). I compared food webs where basal species either show diffuse (all species compete with each other identically: no within guild extinctions following primary extinction) or asymmetric competition (unequal interspecific competition: within guild extinctions are possible). Basal competitive exclusion drives extra extinction cascades across all trophic levels, with the effect amplified in larger ecosystems, though varying connectance has little impact on results. Secondary extinction patterns based on the relative abundance of the species lost in the primary extinction differ qualitatively between diffuse and asymmetric competition. Removing asymmetric basal species with low (high) abundance triggers fewer (more) secondary extinctions throughout the whole food web than removing diffuse basal species. Rare asymmetric competitors experience less pressure from consumers compared to rare diffuse competitors. Simulations revealed that diffuse basal species are never involved in extinction cascades, regardless of the trophic level of a primary extinction, while asymmetric competitors were. This work highlights important qualitative differences in extinction patterns that arise when different assumptions are made about the form of direct competition in multi‐trophic food webs.     

Invasive plants have different effects on trophic structure of green and brown food webs in terrestrial ecosystems: a meta‐analysis

Although invasive plants are a major source of terrestrial ecosystem degradation worldwide, it remains unclear which trophic levels above the base of the food web are most vulnerable to plant invasions. We performed a meta‐analysis of 38 independent studies from 32 papers to examine how invasive plants alter major groupings of primary and secondary consumers in three globally distributed ecosystems: wetlands, woodlands and grasslands. Within each ecosystem we examined if green (grazing) food webs are more sensitive to plant invasions compared to brown (detrital) food webs. Invasive plants have strong negative effects on primary consumers (detritivores, bacterivores, fungivores, and/or herbivores) in woodlands and wetlands, which become less abundant in both green and brown food webs in woodlands and green webs in wetlands. Plant invasions increased abundances of secondary consumers (predators and/or parasitoids) only in woodland brown food webs and green webs in wetlands. Effects of invasive plants on grazing and detrital food webs clearly differed between ecosystems. Overall, invasive plants had the most pronounced effects on the trophic structure of wetlands and woodlands, but caused no detectable changes to grassland trophic structure.     

Plant-mediated effects of drought stress on host preference and performance of a grass miner

The plant stress hypothesis predicts that environmental stress increases the suitability of plants as food for herbivores, especially for senescence feeders. Yet, performance is enhanced only at moderate stress intensities in several herbivores. Even more paradoxically, a large number of insect species prefer and perform better on vigorously growing plants. In order to test plant stress theory, we conducted a laboratory experiment in which the influence of plant water stress on host preference and the performance of the grass miner Chromatomyia milii was studied. We imposed a gradient of stress intensities, i.e. 25, 50, 75, 150, and 300‐ml weekly‐administered water per grass pot, in order to study the full range of responses of C. milii to water stressed plants. Plant stress intensity was quantified by measuring individual plant mass, foliar water content and the concentration of the photosynthetic pigments chlorophyll a and b. Plant mass had decreased from the 150 and 300‐ml treatments to the lowest water treatment at the end of our experiment, which was mainly a result of a reduction in leaf area and leaf number. Foliar water content was clearly negatively affected by water shortage. Chlorophyll a and b also decreased with water shortage. Finally, the stress intensity measurements showed that plants acclimated to water stress conditions throughout the experiment. Feeding and oviposition preference of C. milii was positively related to water supply. No larvae survived on two lowest water treatments and only 38% survived on the 75‐ml treatment, while more than 80% survived on the 150 and the 300‐ml treatments. Offspring development time was longer on the 75‐ml treatment than on the 150 and the 300‐ml treatments. We also evaluated the mechanisms that could explain the response of C. milii to water stressed plants. Although no relationship between water treatment and foliar amino acid concentration was found, we observed significantly higher foliar protein concentrations in the 25 and the 50‐ml treatments. This supports the hypothesis that abiotic stress causes an increase of nitrogenous compounds in plants. Leaf senescence following self‐pruning, a process by which H. lanatus plants acclimate to drought conditions, was responsible for the dramatically high offspring mortality on the water stressed plants. The shape of the plant stress intensity–herbivore response relationship showed strong variation and depended both on the type of plant stress intensity measure and herbivore response variable involved. Yet, all relationships showed a monotonic increase of herbivore preference and performance with decreasing plant stress intensity. This indicates that C. milii prefers and performs better on vigorously growing plants. We found no support for an increased herbivore performance on moderately or severely stressed plants.     

Population Dynamics of the Polychaetous Annelids of an Intertidal Habitat in Upper Old Tampa Bay,Florida

Following an extensive red-tide induced mass mortality of the benthic macrofauna of a sandy intertidal habitat, the population dynamics of the polychaete species were studied. Detailed studies of the 12 most abundant species are presented. Data concerning total population levels, reproduction, recruitment, distribution within the intertidal zone, and gut content analyses are integrated in order to explain the observed spatial and temporal patterns of distribution and abundance. Potential competive interactions for food are considered to be the most important factor for explaining the observed ecological patterns. The polychaete species studied are divided into three trophic guilds: an omnivorous guild that feeds predaciously and as non-selective deposit feeders, a surface feeding guild consisting of species usually considered to be selective surface deposit feeders, and a subsurface feeding guild usually considered as non-selective infaunal deposit feeders. Within and between guild interactions are discussed.     

Environmental and plant genetic effects on tri‐trophic interactions

The effects of plant genotype and environmental factors on tri‐trophic interactions have usually been investigated separately, limiting our ability to compare the relative strength of these effects as well as their potential to interactively shape arthropod communities. We studied the interactions among the herb Ruellia nudiflora, a seed predator, and its parasitoids using 14 maternal plant families grown in a common garden. By fertilizing half of the plants of each family and subsequently recording fruit number, seed predator number, and parasitoid number per plant, we sought to compare the strength of plant genetic effects with those of soil fertility, and determine if these factors interactively shape tri‐trophic interactions. Furthermore, we evaluated if these bottom–up factors influenced higher trophic levels through changes in abundance across trophic levels (density‐mediated) or changes in the function of species interactions (trait‐mediated). Plant genetic effects on seed predators and parasitoids were stronger than fertilization effects. Moreover, we did not find plant genetic variation for fertilization effects on fruit, seed predator, or parasitoid abundance, showing that each factor acted independently on plant resources and higher trophic levels. Both bottom–up forces were transmitted via density‐mediated effects where increased fruit number from fertilization and plant genetic effects increased seed predator and parasitoid abundance; however, seed predator attack was density‐dependent, while parasitoid attack was density‐independent. Importantly, there was evidence (marginally significant in one case) that fertilization modified the function of plant‐seed predator and seed predator–parasitoid interactions by increasing the number of seed predators per fruit and decreasing the number of parasitoids per seed predator, respectively. These findings show that plant genetic and soil fertility effects cascaded up this simple food chain, that plant genetic effects were stronger across all trophic levels, and that these effects were transmitted independently and through contrasting mechanisms.     

Aphid‐tending ants on introduced fennel: can resources derived from non‐native plants alter the trophic position of higher‐order consumers?

1. Although plant invasions often reduce insect abundance and diversity, non‐native plants that support phytophagous insects can subsidise higher trophic levels via elevated herbivore abundance. 2. Here ant–aphid interactions on non‐native fennel on Santa Cruz Island, California are examined. Fennel hosts abundant, honeydew‐producing fennel aphids. The patchiness of fennel and the relative lack of honeydew‐producing insects on other plants at our study sites suggest that assimilation of fennel‐derived honeydew would increase the abundance and decrease the trophic position of the omnivorous, aphid‐tending Argentine ant. 3. To assess the strength of the ant–aphid interaction, a comparison of ant abundance on and adjacent to fennel prior to and 3 weeks after experimental aphid removal was performed. Compared with control plants with aphids, ants declined in abundance on and around fennel plants following aphid removal. At the habitat scale, pitfall traps in fennel‐dominated habitats captured more ants than in fennel‐free scrub habitats. 4. To determine if assimilation of aphid‐produced honeydew reduces the ant's trophic position, variation in δ¹⁵N values among ants, plants and other arthropods was analysed. Unexpectedly, δ¹⁵N values for ants in fennel‐dominated habitats were higher than those of arthropod predators from the same sites and also higher than those of ants from fennel‐free habitats. 5. Our results illustrate how introduced plants that support phytophagous insects appear to transfer energy to higher trophic levels via elevated herbivore abundance. Although assimilation of fennel‐derived honeydew did not appear to reduce consumer trophic position, spatial variation in alternative food resources might obscure contributions from honeydew.