Small Caterpillars Research Articles

Ontogenetic Changes in the Feeding Behaviour of Helicoverpa armigera Larvae on Pigeonpea (Cajanus cajan) Flowers and Pods.

Despite substantial research examining caterpillar-plant interactions, changes in the feeding behaviour of lepidopteran larvae as they develop are poorly understood. In this study, we investigated ontogenetic changes in the behaviour of Helicoverpa armigera larvae feeding on reproductive structures of pigeonpea (Cajanus cajan). Specifically, we examined the preference for and avoidance of pigeonpea flowers and pods of first, second, third, and fourth instar H. armigera larvae. We also conducted a no-choice assay to compare the ability of third and fourth instar larvae to penetrate pigeonpea pod walls, which act as a physical defence against herbivory. When presented with a choice between pigeonpea pods and flowers, different instars behaved differently. First and second instar larvae largely avoided pigeonpea pods, instead feeding on flowers; third instar larvae initially avoided pods, but by 24 h, did not strongly discriminate between the structures; and fourth instars demonstrated a preference for pods. When initially placed on pods, first instars were slower than other instars to leave these structures, despite pods being suboptimal feeding sites for small caterpillars. We identified a clear instar-specific ability to penetrate through the pod wall to reach the seeds. Most third instar larvae were unable to penetrate the pod wall, whereas most fourth instars succeeded. Third instars suffered a physiological cost (measured by relative growth rate) when boring through the pod wall, which was not observed in fourth instars. Our study further illuminates the insect-plant interactions of the H. armigera-pigeonpea system and provides evidence for the significant changes in feeding behaviour that may occur during lepidopteran larval development.

Adult female Barred Owl (Strix varia) eating small caterpillars (family Geometridae) while guarding a recently fledged chick in Winnipeg, Manitoba

Adult female Barred Owl (Strix varia) eating small caterpillars (family Geometridae) while guarding a recently fledged chick in Winnipeg, Manitoba

A double-edged sword: parental care increases risk of offspring infection by a maternally vectored parasite.

Parental care can protect offspring from predators but can also create opportunities for parents to vector parasites to their offspring. We hypothesized that the risk of infection by maternally vectored parasites would increase with the frequency of mother–offspring contact. Ammophila spp. wasps (Hymenoptera: Sphecidae) build nests in which they rear a single offspring. Ammophila species exhibit varied offspring provisioning behaviours: some species enter the nest once to provision a single, large caterpillar, whereas others enter the nest repeatedly to provision with many smaller caterpillars. We hypothesized that each nest visit increases the risk of offspring parasitism by Paraxenos lugubris (Strepsiptera: Xenidae), whose infectious stages ride on the mother wasp (phoresy) to reach the vulnerable Ammophila offspring. We quantified parasitism risk by external examination of museum-curated Ammophila specimens—the anterior portion of P. lugubris protrudes between the adult host's abdominal sclerites and reflects infection during the larval stage. As predicted, Ammophila species that receive larger numbers of provisions incur greater risks of parasitism, with nest provisioning behaviour explaining ca 90% of the interspecific variation in mean parasitism. These findings demonstrate that parental care can augment, rather than reduce, the risk of parasite transmission to offspring.

Lepidopteran caterpillars in the Cretaceous: were they a good food source for early birds?

Caterpillars are an omnipresent component in modern terrestrial faunas. Not surprisingly, they represent an important food source for larger animals. The oldest fossils of caterpillars are known from the Cretaceous. Yet, so far, only nine possible Cretaceous specimens have been reported. Here we expand the known record by four new specimens of caterpillars preserved in 99 million-year-old Kachin amber from Myanmar. The specimens cannot easily be interpreted in a taxonomic or phylogenetic frame. A simple morphometric comparison reveals that the new specimens differ in their relative body dimensions from those of the previously known specimens, expanding the morphological diversity of Cretaceous caterpillars. All caterpillars from the Cretaceous of which sizes are known are rather small, the largest definite one being only about 5 mm. Comparison to younger ambers reveals no clear directed preservation bias towards preserving only small caterpillars. In addition, the sizes of known adult lepidopterans from the Cretaceous are compatible with rather small caterpillars, although possibly slightly larger than the known ones. While small, the observed size range appears still to be within the size range of food items known to be consumed by modern birds. It therefore seems likely that also in the Cretaceous caterpillars were adequate food items for early birds.

Acoustic stimuli from predators trigger behavioural responses in aggregate caterpillars

AbstractPrey detect their predators through predator signals and cues and, consequently, respond with anti‐predatory behaviours to inhibit the action of their aggressors. Lepidopterans can intercept signals emitted by predators and may defend themselves through chemical, morphological or behavioural responses. In this study, we investigated the effect of acoustic stimuli of different predators on defensive behaviour of gregarious caterpillars. Our results demonstrated that Hylesia nigricans (Lepidoptera, Saturniidae) caterpillars alter their behaviour (i.e. abruptly raising the head) in response to the acoustic stimulus of the predators (i.e. predation risk signals from birds and wasps). The magnitude of this response depended on predator identity and caterpillar body size. Larger caterpillars responded more strongly to predatory stimuli than smaller caterpillars. However, regardless of the size of the caterpillars, they responded more strongly to the stimuli of wasps. In addition, we identified that H. nigricans caterpillars emit ultrasonic noise after detecting the stimuli of the predators – this noise seems to function as an alert about the risk of predation during the early stages of development (second and fifth instars). The duration of ultrasonic emission (i.e. milliseconds) increases with the number of repetitions of the stimuli (i.e. wing‐beat sounds of the wasps and insectivorous birds). These results provide novel information about predation risk in interactions among caterpillars and their predators, and indicate possible communication among invertebrates mediated by the risk of predation.

Host‐size decisions of female parasitoid wasps seeking hidden hosts

1. Choice of host size may play a critical role in parasitoid success, a task that takes on added complications when dealing with concealed hosts, but most such studies of insect behaviour have only taken place in the laboratory.2. This study investigates the success of a wasp (Alabagrus texanus: Braconidae) in finding host caterpillars Herpetogramma theseusalis (Crambidae) of the most effectively handled size hidden in shelters, in both the field and the laboratory.3. First, the study tested wasp preference and success in parasitizing large, middle‐sized and small caterpillars (> 5, 3–5, < 3 mm) presented in the open, one at a time, in the laboratory. The wasps attacked (inserted or attempted to insert their ovipositor) a higher proportion of middle‐sized (3–5 mm) caterpillars compared with either small (< 3 mm) or large (> 5 mm) caterpillars. Naïve wasps attacked large caterpillars more often than did experienced wasps. Wasps responded to increasing caterpillar size by increasing the number of legs used to pin their prey rather than by increasing handling time.4. The frequencies of visits to shelters in the field containing a majority of either large or middle‐sized caterpillars were then compared, followed by a test providing the wasps with similar choices under controlled laboratory conditions. Wasps most frequently visited shelters containing a majority of middle‐sized caterpillars both in the field and under controlled laboratory conditions.5. The combined results confirmed that the wasps can size‐select their hosts both in the field and in laboratory tests.

Distribuição vertical de Crysodeixis includens (Lepidoptera: Noctuidae) em cultivar transgênica e convencional de algodoeiro

O objetivo deste estudo foi avaliar a distribuição intra-planta de lagartas de Crysodeixis includens no cultivar convencional FM 993 e na sua isolinha transgênica FM 975 WS (WideStrike), que expressa as proteínas tóxicas Cry1Ac e Cry1F. O experimento foi realizado com o delineamento em blocos casualizados em esquema de parcelas subdivididas, com seis tratamentos (três terços da planta x duas faces foliares) e 40 repetições. As plantas foram visualmente divididas em três terços (superior, médio e inferior) e nas faces adaxial e abaxial das folhas, registrando-se o número de lagartas. Lagartas de C. includens foram identificadas em maior número no cultivar não-Bt (FM 993), enquanto que, na cultivar transgênica FM 975 WS (WideStrike), foram constatadas apenas as pequenas, em número significativamente menor que na sua isolinha não-Bt (FM 993). As lagartas, independentemente do tamanho e do cultivar, foram mais encontradas na parte inferior e mediana do dossel das plantas e, ainda, na face abaxial das folhas.

Catch fast and kill quickly: do tiger beetles use the same strategies when hunting different types of prey?

BackgroundTiger beetles (Coleoptera: Cicindelidae) are fast running predatory insects preying on different small insects and other terrestrial arthropods. Prey is located by sight and captured after short and fast pursuit interspersed with pause-and-look behaviour. At least some tiger beetle species can recognise the size and location of prey using memory, which probably allows them to achieve greater hunting success.Material and MethodsTwo eurytopic tiger beetle species known to occur in different types of habitat were used in the study: Cicindela hybrida hybrida, a very common central European beetle found even in artificial habitats such as sandy roads or gravel pits, and Calomera littoralis nemoralis, a species widely distributed in southern European countries and occurring on sandy sea beaches, in salt marshes, as well as on sandy banks of rivers and lakes. Both species are very similar in body size. Specimens used in the study were collected in the field and later tested in the laboratory. We checked whether tiger beetles use different hunting strategies when attacking prey of different sizes and abilities to escape as well as whether the sex of the studied species makes a difference in its hunting behaviour.ResultsThe hunting strategies of both tiger beetle species consist of the following main phases: identification, pursuit (often with stops), attack, and optional release of the prey, and then the secondary attack, abandonment of the prey, or consumption of the prey. Considerable differences were noticed in hunting behaviour depending on the type of prey, its movement ability and escape potential. Caterpillars were attacked without pursuit, in the head or directly behind the head where a concentration of nerves and main muscles responsible for walking are located. Effective attacks on beetles were executed at the connection between the thorax and the abdomen. Calomera littoralis strongly preferred slow moving prey, while Cicindela hybrida preferred in equal measure slow moving prey and medium-sized fast moving prey. The experiment on the preferred size of prey indicated small beetles and small caterpillars as favoured by Calomera littoralis, while Cicindela hybrida preferred medium-sized fast moving prey and large caterpillars.DiscussionThe hunting behaviour of Calomera littoralis and Cicindela hybrida is complicated and includes a number of phases allowing to locate, capture and kill the prey. Beetles are able to discriminate between different types of prey and apply different behavioural tactics to hunt it. As the particular strategies are used to increase hunting success, and as a result allow to accumulate energy for future activity of the predator, it can be expected that such a type of hunting behaviour is characteristic also of other tiger beetle species.

Leaf-slicing behavior in the Blue-headed Parrot (Pionus menstruus) in central Amazonia is likely linked to highly selective caterpillar predation

ABSTRACT Although they eat seeds, pulp, petals, pollen, and nectar, parrots are rarely recorded as eating leaves, with only one species being fully folivorous. In terra firme forest in south-central Brazilian Amazonia, we recorded apparent folivory over a several sequential days by the Blue-headed Parrot (Pionus menstruus) on young leaflets of Swartzia polyphylla (Fabaceae). Closer examination, however, showed that the majority of processed leaves also contained small caterpillars, so that we consider this to be a record of highly selective insectivory, which is also uncommonly reported in parrots. We speculate that such events may be more common than realized and act as dietary supplements for a seed-based diet.

Size, spines and crochets: defences of luna moth caterpillars against predation by brown anoles

ABSTRACTField-testing predator-prey interactions between invasive exotic brown anole lizards and sentinel larvae of the luna moth, Actias luna, suggests that late instars of this saturniid, and perhaps many other saturniids, are well defended by their spines and crochets against predation by lizards. However, Actias luna larvae remain vulnerable to anole predation until they reach about 3 cm in length. In contrast, even relatively small caterpillars that are chemically defended, such as those of the io moth, Automeris io, and the monarch butterfly, Danaus plexippus, were shown to be well defended against anoles by their toxins. There is an obvious advantage for caterpillars to be chemically defended. However, there is also a tradeoff: Danaus plexippus are restricted to feeding on a relatively small group of plants, which limits resources and requires specialisation; in polyphagous Automeris io, the constant production of elaborate spines and toxin leads to prolonged larval development, which is at least twice as long as that of Actias luna, and exposes the caterpillars to mortality factors such as parasitism for a longer duration.

Extractive foraging of toxic caterpillars in wild northern pig-tailed macaques (Macaca leonina).

Extractive foraging in nonhuman primates may involve different levels of technical complexity in terms of the number of actions that must be performed and the manual dexterity involved. We describe the extractive foraging of caterpillars in wild northern pig-tailed macaques (Macaca leonina) at Khao Yai National Park, Thailand. The study group, observed from May to December 2016 (n=146days), comprised 60-70 habituated individuals, including 3-4 adult males, 20-23 adult females, and 36-47 immatures. Four adult males and five adult females, observed from September to November 2016 for a total of 24days, were selected for focal animal sampling. Northern pig-tailed macaques were observed eating at least two families (Erebidae and Limacodidae) and three genera (Macrobrochis sp., Phlossa sp. and Scopelodes sp.) of caterpillars. While the monkeys ate short and small caterpillars with stinging setae and non-setae caterpillars without processing, they performed extensive caterpillar-rubbing behavior on large and long caterpillars with stinging setae. Based on 61 extractive foraging bouts, we found that caterpillar rubbing was hierarchically organized into five stages and 12 elements. Five stages of behavior sequence started with picking the caterpillar up, transporting it to a substrate, rubbing it to remove stinging setae, ingesting it, and then cleaning hands and mouth. Only adult macaques were observed using a leaf to rub stinging caterpillars.

Sequential sampling for evaluation of caterpillars, small and large in soybean

Among the main limitations of grain yield in soybean are the insect pests, especially the defoliating caterpillars. It is necessary to reveal and quantify these insects for proper decision-making according to pre-established control levels. So, for the implementation of appropriate management of these soybean insect pests, there is a necessity to establish a sampling plan that will allow an effective, reliable and less time consuming estimation of the population density of pests. This work aims to establish a sequential sampling plan for small and large caterpillars in soybean in accordance with the sequential test of probability rates. Data was collected for two growing seasons (2010/2011 and 2012/2013). The number of small caterpillars ( 1.5 cm) of the species Anticarsia gemmatalis (Hubner, 1818), Chrysodeixis includens (Walker, 1857) and Spodoptera eridania (Cramer, 1872) was estimated using the vertical cloth-to-beat in a grid of 154 sample points marked with a 20 × 20 m spacing in an area of 6.16 ha of soybean. The distribution of small and large caterpillars in soybean is aggregated. According to the sequential sampling plan, the number of maximum samples, for the decision making of the control or not of caterpillars in soybean is 30 samples units. Key words: Pest management, sampling plan, vertical cloth-to-beat, spatial distribution.

Herbivore density mediates the indirect effect of herbivores on plants via induced resistance and apparent competition

AbstractHerbivore‐induced plant resistance and apparent competition are two indirect ways herbivores interact. If a less damaging herbivore indirectly suppresses the abundance of a more damaging herbivore via these mechanisms, then plants may ultimately benefit. Changes in herbivore density, however, can dictate the intensity of species interactions and may play a critical role in determining the outcome of plant‐ and predator‐mediated herbivore interactions. We tested the effects of herbivore density on the strength of indirect interactions among phloem‐feeding aphids and herbivorous caterpillars and the outcome of these interactions for their shared host plant, cotton (Gossypium hirsutum). We quantified the survival of caterpillars on host plants that were infested with varying densities of aphids in the presence and absence of predators (ladybeetles). We found that aphids induced defensive proteins in cotton plants and that caterpillar survival was negatively affected by induced resistance. Likewise, we found that the presence of aphids increased predation of caterpillars by ladybeetles, but that apparent competition between aphids and caterpillars was density dependent. Ladybeetles consumed relatively high numbers of small caterpillars at low to intermediate aphid densities, but essentially became aphid specialists at high aphid densities. Aphid induced defenses and apparent competition combined such that plant damage by caterpillars was lowest when predators were present at low aphid density (induced resistance + highest level of apparent competition). This suggests that herbivores can benefit plants, but the effect on host plants is mediated by herbivore density. Indirect herbivore‐plant mutualisms may increase plant quality, plant fitness, and yield of crop plants and these interactions need to be considered in ecologically based pest management plans. In addition, these interactions likely alter arthropod community structure and natural selection on anti‐herbivore defense traits in plants in natural systems.

The Influence of Host Plant Extrafloral Nectaries on Multitrophic Interactions: An Experimental Investigation.

A field experiment was conducted with outplantings of the native perennial shrub Senna mexicana var. chapmanii in a semi-natural area adjacent to native pine rockland habitat in southern Florida. The presence of ants and the availability of extrafloral nectar were manipulated in a stratified random design. Insect communities were monitored and recorded over a period of six months with a view to addressing three main questions. Do ants provide biotic defense against key herbivores on S. chapmanii? Is the presence of ants on S. chapmanii mediated by EFN? Finally, are there ecological costs associated with the presence of ants on S. chapmanii, such as a reduction in alternative predator or parasitoid numbers? Herbivores on S. chapmanii included immature stages of three pierid butterflies, and adult weevils. Eight species of ants were associated with the plants, and other predators included spiders, ladybugs, wasps, and hemipterans. Parasitic, haemolymph-sucking midges (Ceratopogonidae) and parasitoid flies were also associated with the caterpillar herbivores, and possibly the extrafloral nectaries of the plants. The presence of ants did not appear to influence oviposition by butterflies, as numbers of lepidopterans of all developmental stages did not differ among treatments. Significantly more late instar caterpillars, however, were observed on plants with ants excluded, indicating that ants remove small caterpillars from plants. Substantially more alternative predators (spiders, ladybugs, and wasps) were observed on plants with ants excluded. Rates of parasitization did not differ among the treatments, but there were substantially fewer caterpillars succumbing to virus among those collected from control plants.We provide a rare look at facultative ant-plant mutualisms in the context of the many other interactions with which they overlap. We conclude that ants provide some biotic defense against herbivores on S. chapmanii, and plants benefit overall from the presence of ants, despite negative impacts on non-ant predators.

Body size affects the evolution of eyespots in caterpillars

Many caterpillars have conspicuous eye-like markings, called eyespots. Despite recent work demonstrating the efficacy of eyespots in deterring predator attack, a fundamental question remains: Given their protective benefits, why have eyespots not evolved in more caterpillars? Using a phylogenetically controlled analysis of hawkmoth caterpillars, we show that eyespots are associated with large body size. This relationship could arise because (i) large prey are innately conspicuous; (ii) large prey are more profitable, and thus face stronger selection to evolve such defenses; and/or (iii) eyespots are more effective on large-bodied prey. To evaluate these hypotheses, we exposed small and large caterpillar models with and without eyespots in a 2 × 2 factorial design to avian predators in the field. Overall, eyespots increased prey mortality, but the effect was particularly marked in small prey, and eyespots decreased mortality of large prey in some microhabitats. We then exposed artificial prey to naïve domestic chicks in a laboratory setting following a 2 × 3 design (small or large size × no, small, or large eyespots). Predators attacked small prey with eyespots more quickly, but were more wary of large caterpillars with large eyespots than those without eyespots or with small eyespots. Taken together, these data suggest that eyespots are effective deterrents only when both prey and eyespots are large, and that innate aversion toward eyespots is conditional. We conclude that the distribution of eyespots in nature likely results from selection against eyespots in small caterpillars and selection for eyespots in large caterpillars (at least in some microhabitats).

Host ontogeny determines parasitoid use of a forest caterpillar

For most organisms, patterns of natural enemy-mediated mortality change over the course of development. Shifts in enemy pressure are particularly relevant for organisms that exhibit exponential growth during development, such as juvenile insects that increase their mass by several orders of magnitude. As one of the dominant groups of insect herbivores in most terrestrial plant communities, larval lepidopterans (caterpillars) are host to a diverse array of parasitoids. Previous research has described how the frequency of herbivore parasitism varies among host plants or habitats, but much less is known about how parasitism pressure changes during host development. To test whether the two major parasitoid taxa, wasps and flies, differentially attack shared hosts based on host developmental stage, we simultaneously exposed early- and late-instar Euclea delphinii Boisduval (Lepidoptera: Limacodidae) caterpillars to parasitism in the field. We found strong evidence that parasitoids partition hosts by size; adult female wasps preferentially parasitized small caterpillars, whereas adult female flies preferred to attack large caterpillars. Our results demonstrate that host ontogeny is a major determinant of parasitoid host selection. Documenting how shifts in enemy pressure vary with development is important to understanding both the population biology and evolutionary ecology of prey species and their enemies.

Physiological selectivity of insecticides to adult of Doru luteipes (Scudder, 1876) (Dermaptera: Forficulidae)

Doru luteipes (SCUDDER, 1876) is considered one of the best natural enemies of the fall armyworm Spodoptera frugiperda (J. E. Smith, 1797) (Lepidoptera: Noctuidae), feeding on their eggs and small caterpillars. For its conservation it is necessary to use selective insecticides to S. frugiperda and harmless to the predator. Therefore, objective of the present work was to evaluate the toxicity of insecticides registered to control of S. frugiperda. It was conducted bioassays with D. luteipes adults treated with insecticides directly by exposure to residues of compounds applied on glass plates and the consumption of eggs of S. frugiperda contaminated and offered each one, 24 and 48 hours after treatment. The insecticides were classified according to indices proposed by IOBC/WPRS. For adults D. luteipes treated directly with insecticides, triflumuron was harmless (class 1); chlorfenapyr and etofenprox as slightly harmful (class 2) teflubenzuron/α-cypermethrin and spinosad moderately harmful (class 3) and thiamethoxan/λ-cyhalothrin harmful (class 4). In bioassay exposure of D. luteipes residues of insecticides applied to glass plates, all products were harmful to the predator, except triflumuron which was considered slightly harmful. The survival of adults after consumption of contaminated eggs was 46.7% for the insecticide tiametoxam/λ-cialotrina considered slightly harmful to the predator. The other insecticides were innocuous. Due to the low toxicity presented by the triflumuron to the D. luteipes adults, this compound can be recommended in programs aimed at integrated pest management the preservation of this natural enemy. All other products must be evaluated in greenhouse and field to prove its toxicity.

Nectar secretion on fern fronds associated with lower levels of herbivore damage: field experiments with a widespread epiphyte of Mexican cloud forest remnants

The oldest group of plants in which nectar secretions have been observed are the Polypodiopsida (ferns sensu lato). Nectaries have been reported in a dozen extant genera. The function of these nectaries has been investigated in several fern species, and in some circumstances has been demonstrated to have an antiherbivore role, attracting and maintaining biotic defence (ants and/or other predatory arthropods). This study documents foliar nectaries in Pleopeltis crassinervata, a widespread Central American epiphyte growing on a variety of trees in cloud forest areas of Veracruz, Mexico. This is a new record for this genus and species. As previous experimental work on epiphytic species of Polypodium has demonstrated a protective role of ants for developing fronds, we conducted similar experiments (using nylon nail polish to cover nectaries rather than excluding ants with bands of sticky resin as in earlier work). The fronds of Pl. crassinervata developed over 6 weeks, at which time damage was assessed. The experiment was simultaneously conducted on a sympatric species lacking nectaries, Polypodium furfuraceum. Herbivore placement experiments were conducted with large and small caterpillars on both of these ferns. Fronds with nectaries covered suffered greater damage from herbivores over the course of their development, compared with fronds that had uncovered nectaries functioning normally. The parallel experiment on Po. furfuraceum showed no difference between manipulated and control fronds. Six species of ants (Brachymyrmex minutus, Crematogaster formosa, Paratrechina longicornis, Solenopsis geminata, S. picea and Wasmannia auropunctata) were observed visiting nectaries of Pl. crassinervata; most were effective in removing herbivore larvae placed on the fronds. The long evolutionary history of ferns may explain why some previous studies of fern nectaries have shown little or no benefit to ferns from nectary visitors, as any coevolved herbivores are those resistant to ant defence. The results suggest that ants protect Pl. crassinervata fronds against herbivory. The presence of nectaries, and the relationship with ants, may contribute to this fern's widespread occurrence and persistence in the face of disturbance, though many other factors also play a role. Ant defence may be more likely to benefit a widespread species of disturbed habitats that encounters a wide range of non-adapted herbivores.

Prey choice, provisioning behaviour, and effects of early nutrition on nestling phenotype of titmice

It is generally assumed that blue and great tits (Cyanistes caeruleus, Parus major) compete for the same type of food (Lepidoptera larvae) during the breeding season and that the former have some advantage because they are usually earlier and can exploit small caterpillars that are often more abundant and are not consumed by great tits. However, it is not clear whether, when confronted with similar circumstances (e.g., nestling demand), blue and great tits show similar preferences for a particular caterpillar type. In this 2-y study, we compare the diet of both tit species in detail by controlling for hatching date and brood size. We also examine how the contribution of caterpillars and spiders to the diet is related to nestling development. A positive relationship was found between the percentage of spiders in the diet and nestling tarsus length in both species, reinforcing the idea that neonatal nutrition could have a strong influence on nestling phenotype (e.g., offspring size). Such a correlation may arise because spiders contain a high level of taurine, an essential nutrient in the early development of young, and/or this prey type contains more calcium than other food items, which may affect the rate of nestling bone mineralization. Blue tits fed their young double the number of tortricid larvae in comparison with great tits, whereas the latter showed a clear preference for noctuids and exploited, with a low frequency, a type of larvae not consumed by blue tits, hairy caterpillars (Lasiocampidae). Our results point to resource partitioning by these species in this forest ecosystem and contribute to a better understanding of feeding ecology of titmice, which is particularly timely in a global warming context.

Non‐trophic effects of litter reduce ant predation and determine caterpillar survival and distribution

The tritrophic model featuring plants consumed by herbivores consumed by parasitoids or predators has become the primary paradigm used to describe herbivore dynamics. However, interactions involving herbivores can be habitat‐ specific and plants often provide habitat, as well as food. Structural complexity of the habitat may favor predators or may allow herbivore prey to escape detection and capture. This study considered the spatial and temporal dynamics of an arctiid caterpillar, Platyprepia virginalis. The tritrophic model that includes only a tachinid parasitoid that attacks P. virginalis and the caterpillars’ primary host‐plant, Lupinus arboreus, has failed to provide much insight into this system. Instead, we found that ants killed and removed many small caterpillars. Protecting caterpillars from ants increased their survival three‐fold and five‐fold in assays conducted during two years. Caterpillars were more likely to survive in short‐term assays at sites that naturally had a deeper cover of dead and living plant material. Experiments with baits showed that ant recruitment declined as litter depth increased on average. These survey results indicated that ant predation was an important source of mortality for young caterpillars and that the presence of thick litter reduced this mortality. These results were corroborated in an experiment that manipulated litter depth and ant access to caterpillars. Previous findings that other defoliating caterpillars increased litter depth and benefitted P. virginalis are also consistent with this hypothesis. Litter acts as an important non‐trophic resource, allowing caterpillars to avoid predation by ants such that wet sites with deep litter act as source populations for caterpillars. Our results show strong effects of both trophic and non‐trophic interactions since plants indirectly provided limiting habitat and this heterogeneous habitat strongly affected risk of predation and ultimately caterpillar abundance and distribution.