Foliage‐age mixing within balsam fir increases the fitness of a generalist caterpillar

1. Manipulative field studies were carried out to evaluate the foliage age preference–performance relationship for an extreme generalist herbivore, the whitemarked tussock moth (Orygia leucostigma Smith) (Lepidoptera: Lymantriidae), within balsam fir [Abies balsamea (L.) Mill]. 2. Field surveys indicated that early instar caterpillars fed almost exclusively on young (i.e. current‐year) foliage, whereas late instars caterpillars fed on both young and mature (i.e. 1‐ and 2‐year‐old) foliage. 3. Survival of early instar caterpillars was highest in treatments where current‐year and/or 1‐year old foliage were available, but decreased significantly on older foliage. In contrast, late instar caterpillars had the highest survival when allowed to feed on all age classes of foliage, whereas potential fecundity was highest for late instars that fed on young foliage. 4. Overall, caterpillars had 32–65% higher fitness when able to feed on all rather than just one age class of foliage. 5. These results support both the ‘complementary diet' hypothesis, which states that dietary mixing of different‐aged foliage can increase nutrient uptake and/or dilute harmful secondary plant chemicals, and the ‘ontogeny’ hypothesis, which attributes changes in diet to changes in the nutritional needs and/or tolerance to plant defences of juvenile insects as they develop.     

Role of dominant tree species on diversity of herbivorous insect community in temperate forests

Oak (Quercus spp.) and hornbeam (Carpinus spp.) are one of dominant tree species in East Asian temperate broad leaf deciduous forests and many insect species, including more than 65% of Lepidoptera species, feed on these trees. We sampled lepidopteran caterpillars from two 0.1 ha plots in a temperate forest to investigate the role of dominant trees (oaks and hornbeams) in herbivore community. In total, we identified 738 caterpillars from 223 Lepidopteran species on 34 tree species. Most caterpillar species were from species-rich families such as Geometridae 25% (56 spp.), Noctuidae, and Tortricidae. After excluding dominant trees, plant-herbivore network analyses showed increased network specialization and nestedness and decreased generality and vulnerability. These results suggest that oaks and hornbeams support a large Lepidopteran herbivore community, and co-occurring plant species support diverse but specific herbivores. Geographical distribution and plant community are closely related to diversity of the herbivore community. Future work is needed to investigate the likelihood that specialist herbivores become relatively more abundant in the forest as oaks are succeeded by hornbeams.     

Experimental evidence of host race formation in Mitoura butterflies (Lepidoptera: Lycaenidae)

A population of herbivorous insects that shifts to a novel host can experience selection pressures that result in adaptation to the new resource. Host race formation, considered an early stage of the speciation process, may result. The current study investigates host shifts and variation in traits potentially involved in the evolution of reproductive isolation among populations of the juniper hairstreak butterfly, Mitoura gryneus. Mitoura are closely associated with their host trees (Cupressaceae) and exhibit host plant fidelity: in addition to larval development and oviposition, host trees support male leks and mating. Female oviposition preference for the natal host, and differential fitness of larvae when reared on natal versus alternate hosts, are indications that specialization and local adaptation to the natal host plant are occurring. Populations with single host plant associations (Juniperus ashei, J. pinchotii and J. virginiana) as well as populations with multiple hosts (both J. ashei and J. pinchotii) were examined. Concordance between female preference and larval performance was found for J. ashei‐associated populations. Population‐level variation in the patterns of female preference and larval performance, both within and among host associations, may reflect differences in the timing and direction of colonization of hosts. For a single nominal species that otherwise exhibits no morphological or phenological differences, the experimental assessment of specialization and host fidelity in M. gryneus provides strong support for the hypothesis of ongoing host race formation in these butterflies.     

Aboveground phytochemical responses to belowground herbivory in poplar trees and the consequence for leaf herbivore preference

Belowground (BG) herbivory can influence aboveground (AG) herbivore performance and food preference via changes in plant chemistry. Most evidence for this phenomenon derives from studies in herbaceous plants but studies in woody plants are scarce. Here we investigated whether and how BG herbivory on black poplar (Populus nigra) trees by Melolontha melolontha larvae influences the feeding preference of Lymantria dispar (gypsy moth) caterpillars. In a food choice assay, caterpillars preferred to feed on leaves from trees that had experienced attack by BG herbivores. Therefore, we investigated the effect of BG herbivory on the phytochemical composition of P. nigra trees alone and in combination with AG feeding by L. dispar caterpillars. BG herbivory did not increase systemic AG tree defences like volatile organic compounds, protease inhibitors and salicinoids. Jasmonates and salicylic acid were also not induced by BG herbivory in leaves but abscisic acid concentrations drastically increased together with proline and few other amino acids. Leaf coating experiments with amino acids suggest that proline might be responsible for the caterpillar feeding preference via presumptive phagostimulatory properties. This study shows that BG herbivory in poplar can modify the feeding preference of AG herbivores via phytochemical changes as a consequence of root‐to‐shoot signaling.     

An altitudinal comparison of caterpillar (Lepidoptera) assemblages on Ficus trees in Papua New Guinea

Aim This analysis of caterpillar (Lepidoptera) beta‐diversity between tropical lowlands and highlands attempts to separate the effects of between‐site (1) turnover of herbivore species on particular host plants, (2) changes in host use by herbivores, and (3) turnover of plant species on changes in herbivore assemblages. Location Two rain forest areas 130 km and 1700 altitudinal metres apart were studied in Papua New Guinea: one in the lowlands (100 m a.s.l.) on the northern coast of the island and one in the central New Guinean cordillera at 1800 m a.s.l. Methods The analysis is based on caterpillar feeding records obtained by quantitative sampling and rearing of caterpillars from four Ficus species studied in the mountains and 21 Ficus species and 62 plant species from other genera and families studied in the lowlands, including three Ficus species studied in both areas. Results Only 17% of species feeding on Ficus in the highlands also occurred in the lowlands. These species represented 1–46% of individuals in caterpillar assemblages on particular Ficus hosts. Widespread species included both Ficus specialists and generalists feeding on numerous plant families. Some of the Ficus specialists changed their preferred host species with altitude. High species turnover was not explained by changes in the species composition of host plants with altitude as lowland and montane assemblages feeding on the same Ficus species showed high turnover. Despite the rarity of widespread caterpillars, the lowland and montane Ficus assemblages were remarkably similar in their dominance structure, species richness, host specificity, generic composition and familial composition. Main conclusions Ficus‐feeding Lepidoptera assemblages between tropical lowlands and highlands are characterized by substantial species turnover not explained by altitudinal changes in the composition of the vegetation. Further, species‐rich plant genera can support caterpillar assemblages with relatively low beta‐diversity compared with species‐poor genera as caterpillars can switch their host preferences from one congeneric host species to another along an altitudinal gradient. Closely related plant species can thus represent a broad, continuously distributed resource along such gradients.     

Induced plant responses to multiple damagers: differential effects on an herbivore and its parasitoid

Herbivore-induced plants responses can affect the preference and performance of herbivores and their natural enemies. These responses may vary depending on the identity and number of herbivore species feeding on the plant so that when herbivores from different guilds feed on plants, the interactions between plants, herbivores, and natural enemies may be disrupted. Tomato plants were damaged either by the caterpillar Spodoptera exigua, or the aphid Macrosiphum euphorbiae, or damaged by both herbivores, or undamaged controls. We measured the preference and performance of S. exigua and its parasitoid Cotesia marginiventris, and activity of proteinase inhibitors (PI) as an indicator of induced resistance. Compared to undamaged plants, caterpillar damage reduced the number of eggs laid by S. exigua adults, reduced growth, consumption, and survival of larval S. exigua and C. marginiventris, and increased activity of PIs 43%; but did not increase attraction of C. marginiventris. While pupal mass of S. exigua was not affected, the pupal mass of C. marginiventris decreased on caterpillar-damaged plants compared to controls. In contrast, plants damaged by aphids were preferred for oviposition by S. exigua, and had increased larval consumption and survival, compared to controls. Aphid feeding did not affect the preference or performance of C. marginiventris, or PI activity, compared to controls. While oviposition was deterred on caterpillar-damaged plants, plants damaged by both herbivores received the same amount of oviposition as controls. The attraction of C. marginiventris to plants damaged by caterpillars and aphids was increased compared to controls. However, plants damaged by both herbivores had similar PI activity, larval growth and survival of S. exigua and C. marginiventris, as plants singly damaged by caterpillars. Overall, the preference component for both the herbivore and parasitoid was more strongly affected by damage due to multiple herbivores than the performance component.     

The role of leaf defensive traits in oaks on the preference and performance of a polyphagous herbivore,Orgyia vetusta

1. Leaves possess traits that mediate the preference and performance of herbivores. Most evidence for the importance of leaf traits as defences against herbivory comes from studies of few model plant species. 2. In a phylogenetically explicit comparison, I explain the differences in preference and performance of tussock moth (Orgyia vetusta Boisduval) larvae on leaves of 27 oak (Quercus) species using nine putative leaf defences. 3. The preference for an oak species correlated positively with the survival of caterpillars. The correlation between preference and performance did not differ between oak species native to the range of tussock moth versus those from outside the herbivore's range. 4. The first principal component of leaf traits predicted survival of caterpillars on oak leaves but only marginally predicted their preference between oak species. A multiple regression model showed that evergreenness, toughness, and condensed tannin content were the best predictors of caterpillar survival, and leaf toughness was the best predictor of host preference. 5. Generalist caterpillars may accurately assess the value of novel food sources. Moreover, many leaf traits that have been found to affect herbivory within a plant species can also be used to predict the fitness of a generalist herbivore between species.     

Associative learning of host presence in non‐host environments influences parasitoid foraging

1. Parasitoids are known to utilise learning of herbivore‐induced plant volatiles (HIPVs) when foraging for their herbivorous host. In natural situations these hosts share food plants with other, non‐suitable herbivores (non‐hosts). Simultaneous infestation of plants by hosts and non‐hosts has been found to result in induction of HIPVs that differ from host‐infested plants. Each non‐host herbivore may have different effects on HIPVs when sharing the food plant with hosts, and thus parasitoids may learn that plants with a specific non‐host herbivore also contain the host. 2. This study investigated the adaptive nature of learning by a foraging parasitoid that had acquired oviposition experience on a plant infested with both hosts and different non‐hosts in the laboratory and in semi‐field experiments. 3. In two‐choice preference tests, the parasitoid Cotesia glomerata shifted its preference towards HIPVs of a plant–host–non‐host complex previously associated with an oviposition experience. It could, indeed, learn that the presence of its host is associated with HIPVs induced by simultaneous feeding of its host Pieris brassicae and either the non‐host caterpillar Mamestra brassicae or the non‐host aphid Myzus persicae. However, the learned preference found in the laboratory did not translate into parasitisation preferences for hosts accompanying non‐host caterpillars or aphids in a semi‐field situation. 4. This paper discusses the importance of learning in parasitoid foraging, and debates why observed learned preferences for HIPVs in the laboratory may cancel out under some field experimental conditions.     

Plant genotype and induced responses affect resistance to herbivores on evening primrose (Oenothera biennis)

Abstract. 1. Although both genotype and induced responses affect a plant's resistance to herbivores, little is known about their relative and interactive effects. This study examined how plant genotype of a native plant (Oenothera biennis) and induced plant responses to herbivory affect resistance to, and interactions among, several herbivores. 2. In a field experiment, genetic and environmental variation among habitats led to variation in the amount of early season damage and plant quality. The pattern of variation in early season infestation by spittlebugs (Philaenus spumarius, a piercing–sucking herbivore) negatively correlated with oviposition preference by a later feeding specialist weevil (Tyloderma foveolatum, a leaf‐chewer). 3. To determine if plant genotype and induced responses to herbivory might be responsible for these field patterns, we performed no‐choice and choice bioassays using four genotypes of O. biennis that varied in resistance. Plants were induced by either spittlebugs or weevils and assays measured the responses of the same specialist weevil as well as a generalist caterpillar (Spodoptera exigua). 4. Resistance to adult weevils was largely unaffected by plant genotype, while they experienced induced resistance following damage by conspecific weevils in no‐choice assays. Caterpillars were more strongly affected by plant genotype than induced responses in both no‐choice and choice assays, but they also fed less and experienced higher mortality on plants previously damaged by weevils. In contrast to the pattern suggested by the field experiment, spittlebugs did consistently induce resistance against either weevils or caterpillars in the bioassay experiment. 5. These results support recent findings that show herbivore species can compete via induced plant responses. Additionally, a quantitative review of the literature demonstrates that plant genotype tends to be more important than interspecific competition among herbivores (plant‐mediated or otherwise) in affecting herbivore preference and performance.     

Response of Brassica oleracea to temporal variation in attack by two herbivores affects preference and performance of a third herbivore

1. Plants are frequently under attack by multiple insect herbivores, which may interact indirectly through herbivore‐induced changes in the plant's phenotype. The identity, order, and timing of herbivore arrivals may influence the outcome of interactions between two herbivores. How these aspects affect, in turn, subsequently arriving herbivores that feed on double herbivore‐induced plants has not been widely investigated. 2. This study tested whether the order and timing of arrival of two inducing herbivores from different feeding guilds affected the preference and performance of a subsequently arriving third herbivore, caterpillars of Mamestra brassicae L. (Lepidoptera: Noctuidae). Aphids [Brevicoryne brassicae L. (Hemiptera: Aphididae)] and caterpillars [Plutella xylostella L. (Lepidoptera: Yponomeutidae)] were introduced onto wild Brassica oleracea L. (Brassicaceae) plants in different sequences and with different arrival times. The effects of these plant treatments on M. brassicae caterpillars were assessed in pairwise preference tests and no‐choice performance tests. 3. The caterpillars of M. brassicae preferred to feed from undamaged plants rather than double herbivore‐induced plants. Compared with undamaged plants, they preferred plant material on which aphids had arrived first followed by caterpillars, whereas they avoided plant material with the reverse order of herbivore arrival. Performance of the caterpillars increased with increasing arrival time between herbivore infestations in double herbivore‐induced plants. Although M. brassicae grew faster on plants induced by aphids than on those induced by caterpillars alone, its performance was not affected by the order of previous herbivore arrival. 4. These results imply that the timing of colonisation by multiple herbivores determines the outcome of plant‐mediated herbivore–herbivore interactions.     

Larval experience induces adult aversion to rearing host plants: a novel behaviour contrary to Hopkins' host selection principle

1. Hopkins' host selection principle (HSP) states that insects should prefer foliage from their rearing host plant over that of an alternative host. 2. The current study tested whether eastern spruce budworm, Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae), that were laid and developed on, respectively, resistant and susceptible white spruce Picea glauca (Moench) Voss, showed differences in their feeding and oviposition preferences for these two hosts. 3. The data revealed that previous experience of spruce budworm on a host tree type does not influence the host acceptance and feeding behaviour of later larval stages. However, adult budworm reared on resistant white spruce needles preferentially selected susceptible white spruce needles as the host for their progeny, whereas those reared on susceptible needles showed no preference. 4. To the authors' knowledge, this is the first report of an insect showing an oviposition preference for the non‐rearing host plant. This would tend to increase mixing between insects from susceptible and resistant trees. The present results thus argue against Hopkins' HSP and suggest that learned aversion to resistant foliage experienced by larvae is carried into the adult stage.     

Elevated volatile concentrations in high‐nutrient plants: do insect herbivores pay a high price for good food?

1. A tritrophic perspective is fundamental for understanding the drivers of insect–plant interactions. While host plant traits can directly affect insect herbivore performance by either inhibiting or altering the nutritional benefits of consumption, they can also have an indirect effect on herbivores by influencing rates of predation or parasitism. 2. Enhancing soil nutrients available to trees of the genus Eucalyptus consistently modifies plant traits, typically improving the nutritional quality of the foliage for insect herbivores. We hypothesised that resulting increases in volatile essential oils could have an indirect negative effect on eucalypt‐feeding herbivores by providing their natural enemies with stronger host/prey location cues. 3. Eucalyptus tereticornis Smith seedlings were grown under low‐ and high‐nutrient conditions and the consequences for the release of volatile cues from damaged plants were examined. The influence of 1,8‐cineole (the major volatile terpene in many Eucalyptus species) on rates of predation on model caterpillars in the field was then examined. 4. It was found that the emission of cineole increased significantly after damage (artificial or herbivore), but continued only when damage was sustained by herbivore feeding. Importantly, more cineole was emitted from high‐ than low‐nutrient seedlings given an equivalent amount of damage. In the field, predation was significantly greater on model caterpillars baited with cineole than on unbaited models. 5. These findings are consistent with the hypothesis that any performance benefits insect herbivores derive from feeding on high‐nutrient eucalypt foliage could be at least partially offset by an increased risk of predation or parasitism via increased emission of attractive volatiles.     

Nuclear polyhedrosis virus as a biological control agent for Malacosoma americanum (Lepidoptera: Lasiocampidae)

In addition to damaging trees, the eastern tent caterpillar is implicated in early fetal loss and late‐term abortion in horses. In a field study, we evaluated the potential biological control of the caterpillar using eastern tent caterpillar nuclear polyhedrosis virus (ETNPV), a naturally occurring virus that is nearly species‐specific. Egg masses were hatched and second instar larvae were fed virus‐inoculated foliage to propagate the virus in vivo. Then, a viral pesticide was formulated at concentrations of 10⁴, 10⁶ and 10⁸ polyhedral inclusion bodies per ml. The pesticide was applied to foliage on which second, third and fourth instar caterpillars were feeding. When the majority of surviving larvae reached the sixth instar, colonies were collected and the surviving caterpillars counted. Mean numbers of surviving caterpillars per treatment were compared via 95% bootstrap confidence intervals. The data indicate second instar caterpillars were highly susceptible to the virus, but only at the highest concentration tested. Third instar caterpillars were also somewhat susceptible to high virus concentrations, while fourth instar caterpillars were fairly resistant. Our data provide the strongest evidence to date that ETNPV can be propagated, harvested and refined for formulation as a biological control agent for eastern tent caterpillar. Its use on this insect may be merited in circumstances where landowners and managers need to protect trees and horses.     

Cascading effects of host size and host plant species on parasitoid resource allocation

1. The bottom‐up factors that determine parasitoid host use are an important area of research in insect ecology. Host size is likely to be a primary cue for foraging parasitoids due to its potential influence on offspring development time, the risk of multiparasitism, and host immunocompetence. Host size is mediated in part by host‐plant traits that influence herbivore growth and potentially affect a herbivore's quality as a host for parasitoids. 2. Here, we tested how caterpillar host size and host plant species influence adult fly parasitoid size and whether host size influences wasp parasitoid sex allocation. We measured the hind tibia lengths and determined the sex of wasp and fly parasitoids reared from 11 common host species of polyphagous caterpillars (Limacodidae) that were in turn reared on foliage of seven different host plant species. 3. We also tested how host caterpillar species, host caterpillar size, and host and parasitoid phenology affect how the parasitoid community partitions host resources. We found evidence that parasitoids primarily partition their shared hosts based on size, but not by host species or phenology. One index of specialisation (d′) supports our observation that these parasitoids are quite generalised within the Limacodidae. In general, wasps were reared from caterpillars collected in early instars, while flies were reared from caterpillars collected in late instars. Furthermore, for at least one species of solitary wasp, host size influenced sex allocation of offspring by ovipositing females. 4. Host‐plant quality indirectly affected the size attained by a tachinid fly parasitoid through its direct effects on the size and performance of the caterpillar host. The host plants that resulted in the highest caterpillar host performance in the absence of enemies also yielded the largest parasitoid flies, which suggests that host plant quality can cascade up to influence the third trophic level.     

Plant quality and predation risk mediated by plant ontogeny: consequences for herbivores and plants

Bottom‐up and top‐down impacts on herbivores can be influenced by plant productivity, structural complexity, vigor and size. Although these traits are likely to vary with plant development, the influence of plant ontogeny on the relative importance of plant quality (i.e. bottom‐up forces) and predation risk (i.e. top‐down forces) has been the focus of little previous investigation. We evaluated the role of plant ontogeny for the relative importance of bottom‐up and top‐down forces on insect herbivore abundance, species richness, and species diversity attacking the tropical tree Casearia nitida. We also quantified the cascading effects on herbivory, growth and reproduction of this plant species. Plant quality traits (nitrogen and phenolic compounds) were assessed in saplings and reproductive trees. Bottom‐up forces were manipulated by fertilizing plants from both ontogenetic stages. Top‐down forces were manipulated by excluding insectivorous birds from saplings and reproductive trees. Plant ontogeny influenced foliage quality in terms of total phenolics, which were in greater concentration in reproductive trees than in saplings; however, it did not influence bottom‐up forces as modified by fertilization. Bird exclusion increased herbivore density with the same magnitude on both stages. Ontogeny influenced species diversity, which was greater in reproductive trees than in saplings, and also influenced treatment impacts on species richness and diversity. Although top‐down forces increased herbivory equally on plants of each ontogenetic stage, the two stages showed different overcompensation responses to increased damage: caged saplings produced greater leaf biomass than non‐caged saplings, whereas caged trees increased in height proportionally more than non‐caged trees. In sum, plant ontogeny influenced the impact of bird predation on herbivore density, species richness, and species diversity, and the growth variables affected by increased damage in caged plants. We suggest that plant ontogeny can contribute to some extent to the influence of plant quality and the third trophic level on herbivores in this system.     

Fitness consequences of host use in the field: temporal variation in performance and a life history tradeoff in the moth <Emphasis Type="Italic">Rothschildia lebeau</Emphasis> (Saturniidae)

That fitness varies as a function of using different hosts is a basic premise of theory addressing the ecology and evolution of oviposition behavior and host selection. Few data exist demonstrating: (1) the effects of different hosts on fitness in the field, and (2) how these effects vary spatially or temporally. Cohorts of caterpillars were followed from hatching to adulthood to test the hypotheses that: (1) hosts have significant effects on herbivore performance in nature, and (2) host "quality" for performance varies predictably (i.e., the rank order is consistent) across herbivore generations. In total, the fates of >2,000 caterpillars were followed on 238 individual host trees. Host species had significant effects on most, but not all, measured components of caterpillar performance in the field. Variation among generations was mainly quantitative rather than qualitative, with few changes in the rank order of hosts in their effects on performance. There was also a strong seasonal effect on performance such that caterpillar growth and survival were higher in the early wet season compared to the late wet season. Using estimates derived from these data, correlations among larval growth rate, larval survival, total development time, and final adult size were examined at the level of host plant species. Across generations, larval survival was consistently poor, development time was long, but final adult size attained was large on the host Spondias mombin. The converse was true for the host Exostema mexicanum. Relative performance on the host Casearia nitida was variable between the other two hosts. Overall, the data suggest that host use involves a predictable tradeoff between larval survival and final adult size, but argue that which is the "better" host from the female perspective will depend on the fitness consequences of producing a few, relatively large offspring versus producing more, relatively small offspring.     

Herbivory meets fungivory: insect herbivores feed on plant pathogenic fungi for their own benefit

Plants are regularly colonised by fungi and bacteria, but plant‐inhabiting microbes are rarely considered in studies on plant–herbivore interactions. Here we show that young gypsy moth (Lymantria dispar) caterpillars prefer to feed on black poplar (Populus nigra) foliage infected by the rust fungus Melampsora larici‐populina instead of uninfected control foliage, and selectively consume fungal spores. This consumption, also observed in a related lepidopteran species, is stimulated by the sugar alcohol mannitol, found in much higher concentration in fungal tissue and infected leaves than uninfected plant foliage. Gypsy moth larvae developed more rapidly on rust‐infected leaves, which cannot be attributed to mannitol but rather to greater levels of total nitrogen, essential amino acids and B vitamins in fungal tissue and fungus‐infected leaves. Herbivore consumption of fungi and other microbes may be much more widespread than commonly believed with important consequences for the ecology and evolution of plant–herbivore interactions.     

Tree diversity and functional leaf traits drive herbivore‐associated microbiomes in subtropical China

Herbivorous insects acquire microorganisms from host plants or soil, but it remains unclear how the diversity and functional composition of host plants contribute to structuring herbivore microbiomes. Within a controlled tree diversity setting, we used DNA metabarcoding of 16S rRNA to assess the contribution of Lepidoptera species and their local environment (particularly, tree diversity, host tree species, and leaf traits) to the composition of associated bacterial communities. In total, we obtained 7,909 bacterial OTUs from 634 caterpillar individuals comprising 146 species. Tree diversity was found to drive the diversity of caterpillar‐associated bacteria both directly and indirectly via effects on caterpillar communities, and tree diversity was a stronger predictor of bacterial diversity than diversity of caterpillars. Leaf toughness and dry matter content were important traits of the host plant determining bacterial species composition, while leaf calcium and potassium concentration influenced bacterial richness. Our study reveals previously unknown linkages between trees and their characteristics, herbivore insects, and their associated microbes, which contributes to developing a more nuanced understanding of functional dependencies between herbivores and their environment, and has implications for the consequences of plant diversity loss for trophic interactions.     

A biotrophic fungal infection of the great burnet Sanguisorba officinalis indirectly affects caterpillar performance of the endangered scarce large blue butterfly Phengaris teleius

Interactions between ecological communities of herbivores and microbes are commonly mediated by a shared plant. A tripartite interaction between a pathogenic fungus-host plant-herbivorous insect is an example of such mutual influences. In such a system a fungal pathogen commonly has a negative influence on the morphology and biochemistry of the host plant, with consequences for insect herbivore performance. Here we studied whether the biotrophic fbngus Podosphaera ferruginea, attacking the great burnet Sanguisorba officinalis, affects caterpillar performance of the endangered scarce large blue butterfly Phengaris teleius. Our results showed that the pathogenic ftmgus affected the number and size of inflorescences produced by food-plants and, more importantly, had in direct, plant-mediated effects on the abun dance, body mass and immune response of caterpillars. Specifically, we found the relationship between caterpillar abundance and variability in inflorescence size on a plant to be positive among healthy food-plants, and negative among infected food-plants. Caterpillars that fed on healthy food-plants were smaller than those that fed on infected food-plants in one studied season, while there was no such difference in the other season. We observed the relationship between caterpillar immune response and the proportion of infected great burnets within a habitat patch to be positive when caterpillars fed on healthy food-plants, and negative when caterpillars fed on infected food-plants. Our results suggest that this biotrophic fungal infection of the great burnet may impose a significant indirect influence on P. teleius caterpillar performance with potential consequences for the population dynamics and structure of this endangered butterfly.     

Density dependence in insect performance within individual plants: induced resistance to Spodoptera exigua in tomato

Net intraspecific density dependence experienced by insect herbivores at the scale of single plants can be a function both of induced resistance in the plant and other interactions among individual herbivores. Theory suggests that non‐linearity in the form of this density dependence can influence the effects of plants on herbivore population dynamics. This study examined both net density dependence at the scale of single plants, and changes in plant quality with herbivore density for Spodoptera exigua caterpillars on tomato plants. One experiment measured the growth of caterpillars moving freely about the plant at different densities, the distribution of damage by these caterpillars, and the quality of the plant as food for caterpillars (growth of caterpillars on undamaged leaf tissue excised from the plant). A second experiment measured plant quality for plants with different amounts of damage by caterpillars confined to particular leaves in mesh bags. Growth of S. exigua caterpillars was found to be negatively density dependent, and this was in part due to decreases in plant quality both as herbivore density increased and as the amount of damage increased. The response of plant quality to herbivores was found to have non‐linear features; there was both a threshold below which no significant decreases in quality (as measured by herbivore growth) occurred, and the decrease in herbivore performance saturated at the highest damage levels. In addition, it was found that caterpillar damage was significantly more aggregated than expected when multiple caterpillars occupy a single plant. This study confirms that host plants have the potential to be a source of density dependence that affects herbivore performance.