Chemical convergence between a guild of facultative myrmecophilous caterpillars and host plants

1. Ants exert strong selective pressure on herbivorous insects, although some caterpillars can live in symbiosis with them using chemical defensive strategies.
2. We investigated the adaptive resemblance of cuticular hydrocarbons (CHCs) in multitrophic systems involving a guild of facultative myrmecophilous caterpillar species (Lepidoptera: Lycaenidae), tending ants (Hymenoptera: Formicidae), and host plants from three families. We hypothesised that the CHCs of the caterpillars would resemble those of their host plants (chemical camouflage).
3. We analysed CHCs using gas chromatography/mass spectrometry. Morisita's similarity index (SI) was used to compare CHC profiles of caterpillar species with different types of ant associations (commensal or mutualistic), ants, and host plants.
4. We found strong convergence between caterpillars' CHCs and plants, especially for commensal species that do not provide secretion rewards for ants. Moreover, we found unexpected chemical convergence among mutualistic (trophobiotic) caterpillar species that offer caloric reward secretions to ants.
5. These results show that the studied caterpillars acquire CHCs through their diet and that they vary according to host plant species and type of ant association (commensalism or mutualism). This ‘chemical camouflage’ of myrmecophilous caterpillars may have arisen as a defensive strategy allowing coexistence with ants on plants, whereas ‘chemical conspicuousness’ may have evolved in the context of honest signalling between mutualistic partners.
6. We suggest the existence of chemical mimicry among myrmecophilous species, especially between mutualistic caterpillars. Cuticular chemical mixtures can play a key adaptive role in decreasing ant attacks and increasing caterpillar survival in multimodal sensory systems.

     

Mortal combat between ants and caterpillars: an ominous threat to the endangered Schaus swallowtail butterfly (<Emphasis Type="Italic">Heraclides aristodemus ponceanus</Emphasis>) in the Florida Keys,USA

The federally endangered Schaus swallowtail butterfly (Heraclides aristodemus ponceanus) has reached critically low numbers. Exotic ants are a potential threat to H. a. ponceanus and other rare butterflies as they can attack immature stages. Ant surveys conducted in subtropical dry forests in Biscayne National Park documented ant species diversity and relative abundance. A caterpillar predator exclusion experiment using physical barriers in different combinations evaluated caterpillar survivorship of both early and late instar caterpillars exposed to different threats. Ant-caterpillar interactions were also documented by placing caterpillars on plants and observing physical interactions between caterpillars and ants. A total of 1418 ants comprising 25 ant species was captured and identified. In canopies of H. a. ponceanus host plants, 243 ants comprising 12 species were found. The four most common ants collected in the host plant canopies were Pseudomyrmex gracilis, Camponotus planatus, Cremastogaster ashmeadi, and Camponotus floridanus. The predator exclusion experiment revealed survivorship was significantly lower for early and late instar caterpillars without any physical barrier, as well as for early instars not protected by a mesh cage. Pseudomyrmex gracilis and C. floridanus were more aggressive towards caterpillars in comparison to other ant species; these two species ranked first and second in the “ant danger index” ranking predatory abilities of the four most common ant species. Pseudomyrmex gracilis is a common arboreal exotic ant in Biscayne National Park and presents a major threat to caterpillars during their earliest life stages.     

Behavioural ecology of defence in a risky environment: caterpillars versus ants in a Neotropical savanna

1. Predatory ants may reduce infestation by herbivorous insects, and slow‐moving Lepidopteran larvae are often vulnerable on foliage. We investigate whether caterpillars with morphological or behavioural defences have decreased risk of falling prey to ants, and if defence traits mediate host plant use in ant‐rich cerrado savanna. 2. Caterpillars were surveyed in four cerrado localities in southeast Brazil (70–460 km apart). The efficacy of caterpillar defensive traits against predation by two common ant species (Camponotus crassus, C. renggeri) was assessed through experimental trials using caterpillars of different species and captive ant colonies. 3. Although ant presence can reduce caterpillar infestation, the ants' predatory effects depend on caterpillar defence traits. Shelter construction and morphological defences can prevent ant attacks (primary defence), but once exposed or discovered by ants, caterpillars rely on their size and/or behaviour to survive (secondary defence). 4. Defence efficiency depends on ant identity: C. renggeri was more aggressive and lethal to caterpillars than C. crassus. Caterpillars without morphological defences or inside open shelters were found on plants with decreased ant numbers. No unsheltered caterpillar was found on plants with extrafloral nectaries (EFNs). Caterpillars using EFN‐bearing plants lived in closed shelters or presented morphological defences (hairs, spines), and were less frequently attacked by ants during trials. 5. The efficiency of defences against ants is thus crucial for caterpillar survival and determines host plant use by lepidopterans in cerrado. Our study highlights the effect of EFN‐mediated ant‐plant interactions on host plant use by insect herbivores, emphasizing the importance of a tritrophic viewpoint in risky environments.     

Myrmecophily in Hesperiidae. The case of Vettius tertianus in ant gardens

The larvae of the hesperiid butterfly Vettius tertianus develop by eating the leaves of Aechmea mertensii, a bromeliad epiphyte restricted to ant gardens. The relationships between ants and V. tertianus larvae highlight the preferential association of the caterpillars with Pachycondyla goeldii (Ponerinae), an ant-garden initiator. The oviposition strategy of V. tertianus may thus imply the identification of the inhabiting ant species and not only the identification of the host plant. The caterpillars neither provide secretions to the ants, nor possess defensive devices (i.e. hairs or appendices) against ants. Their activity rhythm does not isolate them from foraging workers of P. goeldii and their shelters are also attainable by the ants. Moreover, as the cuticular lipid profiles of V. tertianus larvae are clearly different from those of the ants and also from the leaf-surface of A. mertensii, acceptance is not due to mimicry between larvae and plants or ants. However, the caterpillars deposit, on the leaf they eat, silk containing a mixture of substances very similar to those found on their own cuticle. No interaction with ants was recorded during observations, even though the ant gardens were patrolled by numerous P. goeldii individuals during their activity period. But when confronted with the caterpillar, none of the tested ant species reacted aggressively. These results suggest the existence of compounds, other than cuticular lipids, responsible for the absence of aggressiveness in the ants. The case of V. tertianus is relatively new as myrmecophily within Hesperiidae has been described only once. Moreover, it preferentially involves a member of the Ponerinae, a subfamily in which interactions with other arthropods are exceptional.     

Host plant preference in the protected myrmecophilous Transylvanian Blue (<Emphasis Type="Italic">Pseudophilotes bavius hungarica</Emphasis>) butterfly (Lepidoptera: Lycaenidae) and its relationship with potential ant partners

When selecting specific host plants, caterpillars of many lycaenid butterflies, such as the protected Pseudophilotes bavius hungarica, are known to engage in various interactions with ants, which help them survive. Although P. bavius is a protected species, data about its host plant selection is very scarce, and little information is available on its myrmecophilous relationships. Our aim was to identify the host plant characteristics that determine the occurrence of the caterpillar and to clarify the specificity of its myrmecophily. We conducted a series of field surveys regarding host plant characteristics. Laboratory experiments were carried out to investigate the nature of interactions between the caterpillar and its potential ant partners. Control experiments involving non-visiting ants were also performed. On the basis of our findings, the physical characteristics of host plants do not seem to influence host plant choice, but the absence of aphids and the presence of different ant species proved important. According to the results of behavioural assays, neutral reactions to the caterpillars were recorded in the case of ant species that regularly visited the host plant (Lasius paralienus, Camponotus aethiops), in contrast to Tapinoma subboreale, which was not observed at all on the host plants and which behaved aggressively towards the larvae. Therefore, the caterpillar is expected to show a certain ant host selectivity. The study constitutes an essential contribution to our knowledge of the natural history of a protected butterfly species, which can be used as a basis for more appropriate management strategies, while also shedding light on aspects of myrmecophilous relationships in Lycaenidae in general.     

Traditional Regulation of Edible Caterpillar Exploitation in the Kopa Area of Mpika District in Northern Zambia

The Traditional Ecological Knowledge (TEK) of the Bisa people of northern Zambia on species, life cycles and host plants of the caterpillars they exploit for food and for household income generation, and the traditional control of caterpillar harvesting, were investigated through a household survey. The accessed indigenous knowledge on caterpillar biology was validated through a forest survey, caterpillar collections and identifications. Eight species of caterpillars were harvested from the surrounding miombo woodlands. Their life systems and host plants were well understood by members of local communities and this knoweldge was communicated amongst them orally. Popular commercial species were Gynanisa maya Strand and Gonimbrasia zambesina Walker. Over 20 miombo tree species hosted the two species but the mean numbers of the caterpillars on Julbernadia paniculata Troupin were significantly larger than those on other host plants (p < 0.05), indicating that J. paniculata was probably the main host plant of the two species. Traditional regulation of caterpillar harvesting involved: (i) monitoring for edible caterpillar development and abundance and for changes in caterpillar habitats, (ii) protection of host plants and moth eggs against late bush fires, through use of traditional fire technology, and (iii) temporal restriction of edible caterpillar harvesting. The possibility of the Zambian Government promoting caterpillar harvesting as an economic incentive to the Bisa people to conserve both this renewable edible caterpillar natural resource and their environment is discussed.     

Host Plant Specialization and Species Turnover of Caterpillars Among Hosts in the Brazilian Cerrado

Decrease in the species composition similarity of herbivore assemblages with increasing phylogenetic distance between host plants is a widespread pattern. Here we used data for caterpillars in the Brazilian Cerrado to investigate how the similarity in caterpillar species composition decreases as the taxonomic level and genetic distance (GD) of their host plants increases. In addition, we elucidate the plant taxonomic level that provides the greatest contribution to turnover in the caterpillar species composition among host taxa. Adult Lepidoptera were reared from caterpillars collected from 52 plants over 13 yr in the same area, with each host plant sampled for 1 yr. Most species were specialists, with 66 percent of genus specialists among the nonsingleton species. The similarity in caterpillar species composition across plant taxa decreased from host species to genera, and from host genera to orders. Above this level, the similarity was consistently low. The GD between plants explained 82 percent of the variation in the similarity of caterpillar species composition. The contribution of caterpillar species turnover among host orders from the same superorder and among host superorders from the same subclass explained 70 percent of the caterpillar species richness as a whole. Our results lend support to the view that most tropical caterpillars are host specialists. Our findings further indicate that the number of orders and superorders of plants provide the greatest contribution to the total caterpillar richness compared with all of the other host taxonomic levels combined. Abstract in Portuguese is available at http://www.blackwell‐synergy.com/loi/btp .     

Lepidopteran caterpillar fauna of cerrado host plants

The cerrado biome is rich in caterpillar species although the general biology and ecology of most tropical species remains unknown. Three host plant genera (Byrsonima, Erythroxylum and Qualea) were examined for caterpillars in four cerrado sensu stricto areas near Brasi´lia (DF, Brazil), from 1991 to 1995. Altogether, more than 16000 host plants were sampled and less than 20% of them possessed caterpillars. All the caterpillars found were reared under laboratory conditions. We successfully reared 137 species of 24 lepidopteran families. The average number of caterpillar species per host species was 28.3. The faunal similarity among plant genera, as well as among study sites, was low, and not related to the distances between them. The low proportion of host plants with caterpillars and the high incidence of rare species appears to be a general pattern for the cerrado. A large proportion (74%) of the caterpillar species occurred on only one host plant family. The most frequent species were either specialists (restricted to one host plant genus) or generalists. We provide a brief outline of some taxonomic problems, some observations on general biology, and temporal and local patterns of some specific caterpillars species. The methodology used, which included weekly data collection, in restricted study areas of the same habitat, with the same sampling method for recording the caterpillar species on the same host plant species, can be used as a tool to explore biodiversity and to discuss some aspects of the conservation of cerrado insects.     

Bacterial communities within Phengaris (Maculinea) alcon caterpillars are shifted following transition from solitary living to social parasitism of Myrmica ant colonies

Bacterial symbionts are known to facilitate a wide range of physiological processes and ecological interactions for their hosts. In spite of this, caterpillars with highly diverse life histories appear to lack resident microbiota. Gut physiology, endogenous digestive enzymes, and limited social interactions may contribute to this pattern, but the consequences of shifts in social activity and diet on caterpillar microbiota are largely unknown. Phengaris alcon caterpillars undergo particularly dramatic social and dietary shifts when they parasitize Myrmica ant colonies, rapidly transitioning from solitary herbivory to ant tending (i.e., receiving protein‐rich regurgitations through trophallaxis). This unique life history provides a model for studying interactions between social living, diet, and caterpillar microbiota. Here, we characterized and compared bacterial communities within P. alcon caterpillars before and after their association with ants, using 16S rRNA amplicon sequencing and quantitative PCR. After being adopted by ants, bacterial communities within P. alcon caterpillars shifted substantially, with a significant increase in alpha diversity and greater consistency in bacterial community composition in terms of beta dissimilarity. We also characterized the bacterial communities within their host ants (Myrmica schencki), food plant (Gentiana cruciata), and soil from ant nest chambers. These data indicated that the aforementioned patterns were influenced by bacteria derived from caterpillars’ surrounding environments, rather than through transfers from ants. Thus, while bacterial communities are substantially reorganized over the life cycle of P. alcon caterpillars, it appears that they do not rely on transfers of bacteria from host ants to complete their development.     

Behavioral mechanisms underlie an ant-plant mutualism

Predators can reduce herbivory by consuming herbivores (a consumptive effect) and by altering herbivore behavior, life history, physiology or distribution (non-consumptive effects). The non-consumptive, or trait-mediated, effects of predators on prey may have important functions in the dynamics of communities. In a facultative ant-plant mutualism, we investigated whether these non-consumptive effects influenced the host plants of prey. Here, predaceous ants (Forelius pruinosus) consume and disturb a dominant lepidopteran folivore (Bucculatrix thurberiella) of wild cotton plants (Gossypium thurberi). Season-long ant exclusion experiments revealed that ants had a larger proportional effect on damage by B. thurberiella than on caterpillar abundance, a result that suggests ants have a strong non-consumptive effect. Behavioral experiments conducted in two populations over 2 years demonstrated that B. thurberiella caterpillars were substantially less likely to damage wild cotton leaves in the presence of ants due to ant-induced changes in caterpillar behavior. In the absence of ants caterpillars spent more time stationary (potential feeding time) and less time dropping from leaves by a thread of silk than when ants were present. Furthermore, ants altered the spatial distribution of both caterpillars and damage; caterpillars spent relatively more time on the upper surfaces of leaves and caused damage further from the leaf margin in ant exclusion treatments. Both direct encounters with ants and information conveyed when ants walked onto leaves were key events leading to the anti-predator behaviors of caterpillars. This study contributes to a small body of evidence from terrestrial systems demonstrating that the trait-mediated effects of predators can cascade to the host plants of prey.     

Larvae of Maculinea rebeli, a large-blue butterfly, and their Myrmica host ants: wild adoption and behaviour in ant-nests

The behavioural interactions between caterpillars of Maculinea rebeli Hir. and their Myrmica ant hosts were studied, both in the wild at the time of adoption, and inside captive nests of six Myrmica species.
In the wild, freshly moulted, final instar caterpillars left their food-plants at a time of day that coincided with the peak foraging activity of Myrmica (18:00-20:00 h). Once on the ground, caterpillars made no attempt to search for Myrmica but settled and waited for foraging ants to find them, which took up to 1.5 h. There was no adoption ritual: foragers of any Myrmica species picked up the caterpillars within 1–4 sec of discovery, and carried them directly to their nests.
Caterpillars grew from < 2 mg to 110 mg in laboratory ant-nests. About 60 mg was gained in autumn but 40% of this was lost during the winter, while the temperature was < 14 °C. Although caterpillars survived best with their normal host, Myrmica schencki , they could also survive in the nests of other Myrmica species. The presence of queen ants had no effect upon survival. The behaviour of the caterpillars was described and illustrated: this included the production of secretions that were drunk by the ants, begging for food and direct feeding by ants. The preferred solid food was ant eggs.
The results are discussed in terms of the social biology of Myrmica ants. It is hypothesized that Maculinea rebeli caterpillars mimic the touch pheromones of ant worker-larvae. This would explain the inability of ants to recognize caterpillars before touching them, their immediate adoption by any Myrmica species after discovery, host specificity inside wild ant-nests, the absence of queen-effect and the intimate attention of host workers.     

Caterpillars and fungal pathogens: two co-occurring parasites of an ant-plant mutualism

In mutualisms, each interacting species obtains resources from its partner that it would obtain less efficiently if alone, and so derives a net fitness benefit. In exchange for shelter (domatia) and food, mutualistic plant-ants protect their host myrmecophytes from herbivores, encroaching vines and fungal pathogens. Although selective filters enable myrmecophytes to host those ant species most favorable to their fitness, some insects can by-pass these filters, exploiting the rewards supplied whilst providing nothing in return. This is the case in French Guiana for Cecropia obtusa (Cecropiaceae) as Pseudocabima guianalis caterpillars (Lepidoptera, Pyralidae) can colonize saplings before the installation of their mutualistic Azteca ants. The caterpillars shelter in the domatia and feed on food bodies (FBs) whose production increases as a result. They delay colonization by ants by weaving a silk shield above the youngest trichilium, where the FBs are produced, blocking access to them. This probable temporal priority effect also allows female moths to lay new eggs on trees that already shelter caterpillars, and so to occupy the niche longer and exploit Cecropia resources before colonization by ants. However, once incipient ant colonies are able to develop, they prevent further colonization by the caterpillars. Although no higher herbivory rates were noted, these caterpillars are ineffective in protecting their host trees from a pathogenic fungus, Fusarium moniliforme (Deuteromycetes), that develops on the trichilium in the absence of mutualistic ants. Therefore, the Cecropia treelets can be parasitized by two often overlooked species: the caterpillars that shelter in the domatia and feed on FBs, delaying colonization by mutualistic ants, and the fungal pathogen that develops on old trichilia. The cost of greater FB production plus the presence of the pathogenic fungus likely affect tree growth.     

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.     

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.     

Ants benefit from attending facultatively myrmecophilous Lycaenidae caterpillars: evidence from a survival study

Workers of three ant species (Lasius niger, Lasius flavus, Myrmica rubra) were caged in the laboratory together with caterpillars and pupae of five species of lycaenid butterflies. Mortality of ants was 3–5 times higher when the ants were confined with larvae lacking a dorsal nectar organ (Lycaena phlaeas, Lycaena tityrus) rather than with caterpillars which possess a nectar gland (Aricia agestis, Polyommatus bellargus, P. icarus). For all five species, ant survival was always lower at the pupal stage (where a nectar organ is always absent) than at the caterpillar stage and was largely equivalent for the butterfly species tested. The experimental data confirm earlier estimates that ants can derive nutritive benefits from tending facultatively myrmecophilous lycaenid caterpillars, even though these caterpillars produce nectarlike secretions at low rates.     

Chemical disguise as particular caste of host ants in the ant inquiline parasite Niphanda fusca (Lepidoptera: Lycaenidae)

The exploitation of parental care is common in avian and insect 'cuckoos' and these species engage in a coevolutionary arms race. Caterpillars of the lycaenid butterfly Niphanda fusca develop as parasites inside the nests of host ants (Camponotus japonicus) where they grow by feeding on the worker trophallaxis. We hypothesized that N. fusca caterpillars chemically mimic host larvae, or some particular castes of the host ant, so that the caterpillars are accepted and cared for by the host workers. Behaviourally, it was observed that the host workers enthusiastically tended glass dummies coated with the cuticular chemicals of larvae or males and those of N. fusca caterpillars living together. Cuticular chemical analyses revealed that N. fusca caterpillars grown in a host ant nest acquired a colony-specific blend of cuticular hydrocarbons (CHCs). Furthermore, the CHC profiles of the N. fusca caterpillars were particularly close to those of the males rather than those of the host larvae and the others. We suggest that N. fusca caterpillars exploit worker care by matching their cuticular profile to that of the host males, since the males are fed by trophallaxis with workers in their natal nests for approximately ten months.     

A single lycaenid caterpillar gets an ant‐constructed shelter and uninterrupted ant attendance

Ant‐lycaenid associations range from mutualism to parasitism and the caterpillars of some species of lycaenids are reported to enter ant nests for shelter, diapause, or pupation. The present study aimed to examine the nature of the association between Euchrysops cnejus (Fabricius) (Lepidoptera: Lycaenidae) and Camponotus compressus (Fabricius) (Hymenoptera: Formicidae) worker ants on the extrafloral nectary‐bearing cowpea plant, Vigna unguiculata (L.) Walp. (Fabaceae). The abundance patterns of the ants and the lycaenid caterpillars together with the spatial patrolling patterns of the ants on the plants revealed that ant abundance increased with the occurrence of the lycaenid caterpillars and the ants preferred the lycaenids over the extrafloral nectar. Camponotus compressus worker ants constructed a shelter at the cowpea plant base after interaction with one or more lycaenid caterpillar(s) and tended the caterpillars and pupae till the emergence of the butterfly. The ant‐constructed shelters (ACSs) inhabited by the minor caste workers (13 ± 1.3 ants per ACS), were utilized by the caterpillars to undergo pupation. The ants confined their activities predominantly to tending the pod‐feeding caterpillars and the solitary pupa within each ACS. It appears that the behavior of the tending worker ants is modulated by the lycaenid vulnerable stages.     

Social predation by ants as a mortality source for an arboreal gregarious forest pest

Biocontrol of caterpillars by ants is highly variable, and we investigate how the strength of the trophic relationship between ants and an important outbreaking forest pest depends on phenological synchrony and on social foraging. We test the hypothesis that early spring foraging by ants, coupled with eusocial recruitment behavior, could undermine the caterpillar's strategies to achieve either enemy-free space or predator satiation.We use a series of field surveys and experiments in trembling aspen stands (Populus tremuloides) in the boreal forest of eastern Canada to assess the role of ants in early-instar mortality of the outbreaking, gregarious forest tent caterpillar (Malacosoma disstria). We also investigate individual-level mechanisms related to phenology and social behavior that underlie the effectiveness of ants as biocontrol on caterpillars. Our results show that ants climb trees early in the spring and harvest young forest tent caterpillars, suggesting that early phenology does not provide an entirely enemy-free space for caterpillars. Our findings further show that recruitment-based social foraging enables ants to deplete groups of gregarious prey, suggesting that these eusocial insects are particularly effective at generating predation pressure on gregarious herbivores since they do not satiate easily. Finally, a manipulative predator exclusion experiment confirms that ant predation is a significant mortality source for early-instar forest tent caterpillars. Taken together, these results suggest that phenology and sociality could modulate the role of ants as effective caterpillar predators and thus showcase the importance of considering natural history and behavioral traits when studying trophic interactions and their role in population dynamics.     

High host‐plant nitrogen content: a prerequisite for the evolution of ant–caterpillar mutualism?

The amount of nitrogen required to complete an insect's life cycle may vary greatly among species that have evolved distinct life history traits. Myrmecophilous caterpillars in the Lycaenidae family produce nitrogen-rich exudates from their dorsal glands to attract ants for protection, and this phenomenon has been postulated to shape the caterpillar's host-plant choice. Accordingly, it was postulated that evolution towards myrmecophily in Lycaenidae is correlated with the utilization of nitrogen-rich host plants. Although our results were consistent with the evolutionary shifts towards high-nutrient host plants serving as exaptation for the evolution of myrmecophily in lycaenids, the selection of nitrogen-rich host plants was not confined to lycaenids. Butterfly species in the nonmyrmecophilous family Pieridae also preferred nitrogen-rich host plants. Thus, we conclude that nitrogen is an overall important component in the caterpillar diet, independent of the level of myrmecophily, as nitrogen can enhance the overall insect fitness and survival. However, when nitrogen can be obtained through alternative means, as in socially parasitic lycaenid species feeding on ant brood, the selective pressure for maintaining the use of nutrient-rich host plants is relaxed, enabling the colonization of nitrogen-poor host plants.     

Nucleopolyhedroviruses of forest and western tent caterpillars: cross-infectivity and evidence for activation of latent virus in high-density field populations

Abstract. 1. Cyclic population dynamics of forest caterpillars are often associated with epizootics of nucleopolyhedrovirus, but it is not known how these viruses persist between generations or through the fluctuations in host population density. 2. To explore the question of virus persistence at different phases of the population cycle, the nucleopolyhedroviruses of two species of tent caterpillar that co‐occur in British Columbia, Canada, Malacosoma californicum pluviale (western tent caterpillar) and Malacosoma disstria (forest tent caterpillar), were characterised. The cross‐infectivity of the viruses in these two host species was investigated to determine whether there might be a route for virus persistence via the alternative host species. Any virus produced in the cross‐infections was characterised to confirm true cross‐infection or to ascertain whether cross‐inoculation triggered latent virus persisting within the population. 3. The virus associated with forest tent caterpillars (MadiNPV) did not infect western tent caterpillars from low‐density populations, nor did it trigger a latent virus infection; however, inoculation of forest tent caterpillars from high‐density populations with virus from western tent caterpillars (McplNPV) resulted in viral infection, but without a dose–response relationship. 4. Analysis of DNA profiles of virus resulting from cross‐infection of the forest tent caterpillar with McplNPV, revealed that 88% of these infections were caused by MadiNPV rather than McplNPV; however the virus from all 44 infected individuals was identical and differed in DNA profile from the stock MadiNPV used for cross‐infection. This suggests strongly that forest tent caterpillars from high‐density field populations harbour a latent, persistent, or sublethal form of MadiNPV that was triggered by exposure to nucleopolyhedrovirus from the western tent caterpillar. 5. Virus was not activated in western tent caterpillars collected over 2 years of late population decline and the first year of population increase.