Timing of butterfly parasitization of a plant–ant–scale symbiosis

In the Southeast Asian tropics, Arhopala lycaenid butterflies feed on Macaranga ant-plants inhabited by Crematogaster (subgenus Decacrema) ants tending Coccus-scale insects. A recent phylogenetic study showed that (1) the plants and ants have been codiversifying for the past 20–16 million years (Myr), and that (2) the tripartite symbiosis was formed 9–7 Myr ago, when the scale insects became involved in the plant–ant mutualism. To determine when the lycaenids first parasitized the Macaranga tripartite symbiosis, we constructed a molecular phylogeny of the lycaenids that feed on Macaranga by using mitochondrial and nuclear DNA sequence data and estimated their divergence times based on the cytochrome oxidase I molecular clock. The minimum age of the lycaenids was estimated by the time-calibrated phylogeny to be 2.05 Myr, about one-tenth the age of the plant–ant association, suggesting that the lycaenids are latecomers that associated themselves with the pre-existing symbiosis of plant, ant, and scale insects.     

Host‐plant use by two Orthomeria (Phasmida: Aschiphasmatini) species feeding on Macaranga myrmecophytes

Myrmecophytes depend on symbiotic ants (plant‐ants) to defend against herbivores. Although these defensive mechanisms are highly effective, some herbivorous insects can use myrmecophytes as their host‐plants. The feeding habits of these phytophages on myrmecophytes and the impacts of the plant‐ants on their feeding behavior have been poorly studied. We examined two phasmid species, Orthomeria alexis and O. cuprinus, which are known to feed on Macaranga (Euphorbiaceae) myrmecophytes in a Bornean primary forest. Our observations revealed that: (i) each phasmid species relied on two closely‐related myrmecophytic Macaranga species for its host‐plants in spite of their normal plant‐ant symbioses; and (ii) there was little overlap between their host‐plant preferences. More O. cuprinus adults and nymphs were found on new leaves, which were attended by more plant‐ants than mature leaves, while most adults and nymphs of O. alexis tended to avoid new leaves. In a feeding choice experiment under ant‐excluded conditions, O. alexis adults chose a non‐host Macaranga myrmecophyte that was more intensively defended by plant‐ants and was more palatable than their usual host‐plants almost as frequently as their usual host‐plant, suggesting that the host‐plant range of O. alexis was restricted by the presence of plant‐ants on non‐host‐plants. Phasmid behavior that appeared to minimize plant‐ant attacks is described.     

Studies of a South East Asian ant-plant association: protection of Macaranga trees by Crematogaster borneensis

Summary In the humid tropics of SE Asia there are some 14 myrmecophytic species of the pioneer tree genus Macaranga (Euphorbiaceae). In Peninsular Malaysia a close association exists between the trees and the small, non-stinging myrmicine Crematogaster borneensis. These ants feed mainly on food bodies provided by the plants and have their colonies inside the hollow internodes. In a ten months field study we were able to demonstrate for four Macaranga species (M. triloba, M. hypoleuca, M. hosei, M. hulletti) that host plants also benefit considerably from ant-occupation. Ants do not contribute to the nutrient demands of their host plant, they do, however, protect it against herbivores and plant competition. Cleaning behaviour of the ants results in the removal of potential hervivores already in their earliest developmental stages. Strong aggressiveness and a mass recruiting system enable the ants to defend the host plant against many herbivorous insects. This results in a significant decrease in leaf damage due to herbivores on ant-occupied compared to ant-free myrmecophytes as well as compared to non-myrmecophytic Macaranga species. Most important is the ants' defense of the host plant against plant competitors, especially vines, which are abundant in the well-lit pioneer habitats where Macaranga grows. Ants bite off any foreign plant part coming into contact with their host plant. Both ant-free myrmecophytes and non-myrmecophytic Macaranga species had a significantly higher incidence of vine growth than specimens with active ant colonies. This may be a factor of considerable importance allowing Macaranga plants to grow at sites of strongest competition.     

Various Chemical Strategies to Deceive Ants in Three Arhopala Species (Lepidoptera: Lycaenidae) Exploiting Macaranga Myrmecophytes

Macaranga myrmecophytes (ant-plants) are generally well protected from herbivore attacks by their symbiotic ants (plant-ants). However, larvae of Arhopala (Lepidoptera: Lycaenidae) species survive and develop on specific Macaranga ant-plant species without being attacked by the plant-ants of their host species. We hypothesized that Arhopala larvae chemically mimic or camouflage themselves with the ants on their host plant so that the larvae are accepted by the plant-ant species of their host. Chemical analyses of cuticular hydrocarbons showed that chemical congruency varied among Arhopala species; A. dajagaka matched well the host plant-ants, A. amphimuta did not match, and unexpectedly, A. zylda lacked hydrocarbons. Behaviorally, the larvae and dummies coated with cuticular chemicals of A. dajagaka were well attended by the plant-ants, especially by those of the host. A. amphimuta was often attacked by all plant-ants except for the host plant-ants toward the larvae, and those of A. zylda were ignored by all plant-ants. Our results suggested that conspicuous variations exist in the chemical strategies used by the myrmecophilous butterflies that allow them to avoid ant attack and be accepted by the plant-ant colonies.     

Phylogenetic patterns in larval host plant and ant association of Indo-Australian Arhopalini butterflies (Lycaenidae: Theclinae)

The Oriental butterfly genus Arhopala is by far the most species‐rich genus within the subfamily Theclinae. We investigated whether biotic interactions with larval host plants or ants are phylogenetically constrained in the evolutionary history of Arhopala, by collating available information on the ecology of Arhopala from the literature as well as from personal observations, and analysing and interpreting these data rigorously in a phylogenetic context. Larvae of all species in the sister‐group of Arhopala and Flos feed on Fabaceae. However, the predominant feeding preference of Arhopala s.l., with the exception of a particular monophyletic and species‐rich group, called ‘Group A’, appears to be centred on Fagaceae, with additional records of Dipterocarpaceae feeding. The preference for Fagaceae has strong phylogenetic inertia, as indicated by T‐PTP tests designed to test for phylogenetic signal. Adding all available life‐history data, using the phylogeny presented before as scaffolding, further increased the phylogenetic signal in host plant data. Feeding on Fabaceae (mainly in the outgroup) and Euphorbiaceae also showed significant phylogenetic signal, but feeding on Dipterocarpaceae did not and was found scattered across the phylogeny. Except for the Dipterocarpaceae, phylogenetic signal in feeding preferences was very clear, even despite uneven taxon sampling and apparent lability in host plant use. Ant association also demonstrated historical constraint, but based on the phylogenetic hypothesis presented here it was not clear whether increased ant association intimacy emerged independently various times in Group A, as well as in several basal groups. Our finding of distinct phylogenetic patterns in the host plant and to a lesser degree in ant association data contradicts the hypothesis that life‐history traits are very labile in the Lycaenidae. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 84 , 225–241.     

Extrafloral nectaries in the genus Macaranga (Euphorbiaceae) in Malaysia: comparative studies of their possible significance as predispositions for myrmecophytism

Some species of the paleotropical tree genus Macaranga (Euphorbiaceae) live in close association with ants. The genus comprises the full range of species from those not regularly inhabited by ants to obligate myrmecophytes. In Malaysia (Peninsular and Borneo) 23 of the 52 species are known to be ant-associated (44%). The simplest structural adaptation of plants to attract ants are extrafloral nectaries. We studied the distribution of extrafloral nectaries in the genus Macaranga to assess the significance of this character as a possible predisposition for the evolution of obligate myrmecophytism. All species have marginal glands on the leaves. However, only the glands of non- myrmecophytic species function as nectaries, whereas liquids secreted by these glands in myrmecophytic species did not contain sugar. Some non-myrmecophytic Macaranga and transitional Macaranga species in addition have extrafloral nectaries on the leaf blade near the petiole insertion. All obligatorily myrmecophytic Macaranga species, however, lack additional glands on the lamina. The non-myrmecophytic species are visited by a variety of different ant species, whereas myrmecophytic Macaranga are associated only with one specific ant-partner. Since these ants keep scale insects in the hollow stems, reduction of nectary production in ant-inhabited Macaranga seems to be biologically significant. We interpret this as a means of (a) saving the assimilates and (b) stabilization of maintenance of the association's specificity. Competition with other ant species for food rewards is avoided and thereby danger of weakening the protective function of the obligate ant- partner for the plant is reduced. A comparison with other euphorb species living in the same habitats as Macaranga showed that in genera in which extrafloral nectaries are widespread, no myrmecophytes have evolved. Possession of extrafloral nectaries does not appear to be essential for the development of symbiotic ant-plant interactions. Other predispositions such as nesting space might have played a more important role.     

Florivorous myrmecophilous caterpillars exploit an ant–plant mutualism and distract ants from extrafloral nectaries

Females of myrmecophilous butterflies tend to oviposit in plants visited by ant species that engage in stable associations with its larvae. In Banisteriopsis malifolia, caterpillars are attended by the same ants that feed on extrafloral nectaries. A conflict may arise when both the plant and caterpillars compete for ant attention, and ants are assumed to forage on the highest quality resource. By attending caterpillars, ants can be indirectly detrimental to plant fitness because florivorous larvae feed intensively until pupation. In this study, we specifically investigated (i) whether the occurrence of facultative myrmecophilous Synargis calyce (Riodinidae) caterpillars in B. malifolia was based on ant species (Camponotus blandus or Ectatomma tuberculatum) and abundance; (ii) the monopolization of ants by the butterfly larvae and (iii) the florivory rates incurred by the caterpillars on inflorescences. The abundance of S. calyce was six‐fold greater in plants with C. blandus, compared to E. tuberculatum treatments. Caterpillars monopolized up to 50% of C. blandus on the plants, indicating that the resources offered by S. calyce were more attractive to ants than extrafloral nectaries. Florivory by riodinids incurred losses of almost 60% of flower buds. Myrmecophilous riodinids exploited an ant–plant mutualism by attracting aggressive ants that become larvae bodyguards. Thus, this ecological interaction is potentially detrimental to B. malifolia, since the ants, which can provide protection against herbivores, shift to provide defence for one of these herbivores.     

Difference in Intensity of Ant Defense among Three Species of Macaranga Myrmecophytes in a Southeast Asian Dipterocarp Forest1

To examine interspecific variation in the intensity of ant defense among three sympatric species of obligate myrme‐cophytes of Macaranga (Euphorbiaceae), we measured the ratio of ant biomass to plant biomass, ant aggressiveness to artificial damage on host plants, and increase in herbivore damage on host plants when symbiont ants were removed. Increase in herbivore damage from two‐ and four‐week ant exclusion varied significantly among the three species. The decreasing order of vulnerability to herbivory was M. winkleri, M. trachyphylla, and M. beccariana. The antip/ant biomass ratio (= rate of the dry weight of whole ant colonies to the dry weight of whole aboveground plant parts) and ant agressiveness also varied significantly among the three species; the orders of both the ant/plant biomass ratio and ant aggressiveness were the same as in the herbivory increase. These results indicated that the intensity of ant defense differs predictably among sympatric species of obligate myrmecophytes on Macaranga. In addition to the interspecific difference in the total intensity of ant defense, when symbiont ants were excluded, both patterns of within‐plant variation in the amount of herbivore damage and compositions of herbivore species that caused the damage differed among species. This suggests that the three Macaranga species have different systems of ant defense with reference to what parts of plant tissue are protected and what herbivorous species are avoided by ant defense. Thus, it is important to consider the interspecific variation in ant defense among Macaranga species to understand the herbivore community on Macaranga plants and the mechanisms that promote the coexistence of multiple Macaranga myrmecophytes.     

The arms race between heliconiine butterflies and Passiflora plants – new insights on an ancient subject

Heliconiines are called passion vine butterflies because they feed exclusively on Passiflora plants during the larval stage. Many features of Passiflora and heliconiines indicate that they have radiated and speciated in association with each other, and therefore this model system was one of the first examples used to exemplify coevolution theory. Three major adaptations of Passiflora plants supported arguments in favour of their coevolution with heliconiines: unusual variation of leaf shape within the genus; the occurrence of yellow structures mimicking heliconiine eggs; and their extensive diversity of defence compounds called cyanogenic glucosides. However, the protection systems of Passiflora plants go beyond these three features. Trichomes, mimicry of pathogen infection through variegation, and production of extrafloral nectar to attract ants and other predators of their herbivores, are morphological defences reported in this plant genus. Moreover, Passiflora plants are well protected chemically, not only by cyanogenic glucosides, but also by other compounds such as alkaloids, flavonoids, saponins, tannins and phenolics. Heliconiines can synthesize cyanogenic glucosides themselves, and their ability to handle these compounds was probably one of the most crucial adaptations that allowed the ancestor of these butterflies to feed on Passiflora plants. Indeed, it has been shown that Heliconius larvae can sequester cyanogenic glucosides and alkaloids from their host plants and utilize them for their own benefit. Recently, it was discovered that Heliconius adults have highly accurate visual and chemosensory systems, and the expansion of brain structures that can process such information allows them to memorize shapes and display elaborate pre‐oviposition behaviour in order to defeat visual barriers evolved by Passiflora species. Even though the heliconiine–Passiflora model system has been intensively studied, the forces driving host‐plant preference in these butterflies remain unclear. New studies have shown that host‐plant preference seems to be genetically controlled, but in many species there is some plasticity in this choice and preferences can even be induced. Although much knowledge regarding the coevolution of Passiflora plants and heliconiine butterflies has accumulated in recent decades, there remain many exciting unanswered questions concerning this model system.     

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.

     

Diversity of ant-plant interactions: protective efficacy in Macaranga species with different degrees of ant association

The pioneer tree Macaranga in SE Asia has developed manyfold associations with ants. The genus comprises all stages of interaction with ants, from facultative relationships to obligate myrmecophytes. Only myrmecophytic Macaranga offer nesting space for ants and are associated with a specific ant partner. The nonmyrmecophytic species are visited by a variety of different ant species which are attracted by extrafloral nectaries (EFN) and food bodies. Transitional Macaranga species like M. hosei are colonized later in their development due to their stem structure. Before the colonization by their specific Crematogaster partner the young plants are visited by different ant species attracted by EFN. These nectaries are reduced and food body production starts as soon as colonization becomes possible. We demonstrated earlier that obligate ant partners can protect their Macaranga plants against herbivore damage and vine cover. In this study we focused on nonspecific interactions and studied M. tanarius and M. hosei, representing a non-myrmecophyte and a transitional species respectively. In ant exclusion experiments both M. tanarius and M. hosei suffered significantly higher mean leaf damage than controls, 37% versus 6% in M. hosei, 16% versus 7% in M. tanarius. M. tanarius offers both EFN and food bodies so that tests for different effects of these two food rewards could be conducted. Plants with food bodies removed but with EFN remaining had the lowest mean increase of herbivore damage of all experimental groups. Main herbivores on M. hosei were mites and caterpillars. Many M. tanarius plants were infested by a shootborer. Both Macaranga species were visited by various ant species, Crematogaster spp. being the most abundant. We found no evidence for any specific relationships. The results of this study strongly support the hypothesis that non-specific, facultative associations with ants can be advantageous for Macaranga plants. Food bodies appear to have lower attractive value for opportunistic ants than EFN and may require a specific dietary adaptation. This is also indicated by the fact that food body production in the transitional M. hosei does not start before stem structure allows a colonization by the obligate Crematogaster species. M. hosei thus benefits from facultative association with a variety of ants until it produces its first domatia and can be colonized by its obligate mutualist.     

Non-ant antiherbivore defenses before plant-ant colonization in Macaranga myrmecophytes

We examined changes in the intensity of non-ant defenses of three myrmecophytic Macaranga species before and after the initiation of symbiosis with ants in a Bornean dipterocarp forest. The intensities of non-ant defenses at different growth stages of each Macaranga species were estimated by measuring the survival rate of larvae of the common cutworm, Spodoptera litura, when the larvae were fed on fresh leaves from seedlings (saplings) at three growth stages of each Macaranga species. In all species, the intensity of the non-ant defenses when seedlings had not yet received symbiont foundress queens was significantly higher than that after ant defense was well established. These results support the hypothesis that myrmecophytic Macaranga may defend themselves sufficiently via non-ant defenses before beginning symbiosis with ants and that the intensity of non-ant defenses may decrease as the symbiont colony size increases. We suggest that, where the status of myrmecophytism changes as plant–ant colonies grow, the decrease in the intensity of non-ant defenses which we detected after the establishment of ant colonies might generate an optimal allocation of metabolic cost to ant and non-ant defenses under resource limitations. We also measured leaf toughness, which is considered to be one of the most important agents of non-ant defenses against herbivorous insects, at different plant stages to assess its contribution to the change in the intensity of non-ant defenses after ant colonization. However, we found no evidence that changes in leaf toughness have a significant effect on the change in balance of the two antiherbivory mechanisms. Received: February 2, 2001 / Accepted: August 21, 2001     

Characterization of microsatellite markers for plant‐ants of the genus Crematogaster subgenus Decacrema

We present primer sequences for five polymorphic microsatellite loci in ants of the genus Crematogaster subgenus Decacrema that live in obligate symbiosis with host plants of the euphorb genus Macaranga. Microsatellite loci were isolated with a highly efficient method of enrichment. The number of alleles ranged from 10 to 18 for Crematogaster morphospecies 2, for which these markers were developed, with an observed heterozygosity ranging from 0.6578 to 0.9474. The markers were designed for the study of the population as well as of the social structure of colonies.     

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.     

Carbon and nitrogen contents of food bodies in three myrmecophytic species of <Emphasis Type="Italic">Macaranga</Emphasis>: implications for antiherbivore defense mechanisms

 In Macaranga myrmecophytes, differences in the production of the food bodies (FBs), on which symbiont ants feed, may relate to the intensity of antiherbivore defense by the ants. Interspecific comparisons among Macaranga species on such a mutualistic cost give important information on their strategies and evolution of antiherbivore defense. In this study, the carbon and nitrogen contents of FBs as well as the production rate of FBs were measured in three Macaranga species, M. winkleri, M. trachyphylla, and M. beccariana. There were significant differences in the production rates of FBs among species; the investment in FBs was greater in the Macaranga species in which ant defenses were more intensive. The carbon and nitrogen contents of FBs were significantly different among the three species, although they did not match the intensity of ant defense; the nitrogen content, especially, was greatest in the species of least intensive ant defense. It is suggested that Macaranga plants may have differentiated in the dependence on ant defense by controlling the total amount of nitrogen of FBs, not simply by nitrogen content. Received: January 19, 2001 / Accepted: December 23, 2001     

Association between the African lycaenid,Anthene usamba,and an obligate acacia ant,Crematogaster mimosae

The African lycaenid butterfly, Anthene usamba, is an obligate myrmecophile of the acacia ant, Crematogaster mimosae. Female butterflies use the presence of C. mimosae as an oviposition cue. The eggs are laid on the foliage and young branches of the host plant, Acacia drepanolobium. Larvae shelter in the swollen thorns (domatia) of the host tree, where they live in close association with the acacia ants, and each larva occupies a domatium singly. Anthene usamba are tended by ants that feed from the dorsal nectary organ at regular intervals. Larvae also possess tentacle organs flanking the dorsal nectary organ and appear to signal to ants by everting these structures. Larvae were observed to spend most of their time within the domatia. Stable isotope analysis of matched host plant–ant–butterfly samples revealed that Anthene usamba are δ¹⁵N enriched relative to the ants with which they associate. These data, based on the increase in δ¹⁵N through trophic levels, indicate that the caterpillars of these butterflies are aphytophagous and either exploit the ant brood of C. mimosae within the domatia, or are fed mouth to mouth by adult workers via trophallaxis. This is the first documented case of aphytophagy in African Anthene. Pupation occurs inside the domatium and the imago emerges and departs via the hole chewed by the larva. The adult females remain closely associated with their natal patch of trees, whereas males disperse more widely across the acacia savannah. Females prefer to oviposit on trees with the specific host ant, C. mimosae, an aggressive obligate mutualist, and avoid neighbouring trees with other ant species. Adult butterflies are active during most months of the year, and there are at least two to three generations each year. Observations made over a 5‐year period indicate that a number of different lycaenid species utilize ant‐acacias in East Africa, and these observations are summarized, together with comparisons from the literature. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013 , 109 , 302–312.     

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.     

Ant-Repelling Pollinators of the Myrmecophytic <Emphasis Type="Italic">Macaranga winkleri</Emphasis> (Euphorbiaceae)

Many plants have mutualistic relationships with ants, whereby plants provide food and/or nesting sites for the symbiotic ants, and in turn the ants protect the host plants by excluding herbivores. While the ants are useful as guards, they may negatively affect host reproduction by excluding pollinators. Here we studied this potential conflict in the myrmecophytic Macaranga winkleri pollinated by the thrips Dolichothrips fialae. Behavioural responses of ant guards to pollinator thrips and their chemicals, and related chemical analyses, provide evidence that thrips deter ant-guards by secreting droplets containing ant-repelling n-decanoic acid from their anuses. This is the first report of insect pollinators repelling their host’s symbiotic guard ants to perform pollination. This is a novel strategy by which a plant host avoids interference with pollination by ant-guards in an ant–plant mutualism. The acquisition of a pollination system that is resistant to ant attacks may have facilitated the evolution of myrmecophytes in the genus Macaranga.     

Pruning of host plant neighbours as defence against enemy ant invasions: <Emphasis Type="Italic">Crematogaster </Emphasis>ant partners of <Emphasis Type="Italic">Macaranga</Emphasis> protected by "wax barriers" prune less than their congeners

Many ant partners of tropical ant-plants prune the leaves and shoot tips of other plants growing around their hosts. According to the hypothesis proposed by Davidson et al. (Ecology 69:801-808), this specialized behaviour not only protects the host plants against overgrowth, but it also conveys a direct benefit to the ant colony as it removes contact points to the neighbouring vegetation where invasions of enemy ants could occur. Here we test this hypothesis by comparing pruning intensity in five closely related Crematogaster (subgenus Decacrema) plant-ant species (and one species of Technomyrmex) that differ in their exposure to competition by other ants. Pruning intensity was quantified by measuring the area loss of paper tape pieces wrapped around the stems of Macaranga host plants. All Crematogaster (Decacrema) ants tested but not Technomyrmex sp. pruned, but the intensity of the behaviour varied strongly between and within species. Pruning was significantly weaker in the three tested Crematogaster species inhabiting Macaranga host plants with a slippery, waxy stem surface, which functions as a mechanical barrier protecting the specific ant partners against generalist competitors. Pruning was generally stronger on more densely ant-populated trees. Even though the number of ants per twig length was lower in associations of ants with glaucous Macaranga hosts, only part of the variation of pruning activity could be explained by "ant density". When corrected for ant density, "wax-running" Crematogaster (Decacrema) ants still pruned more weakly than their congeners inhabiting non-glaucous Macaranga hosts. Pruning is obviously most important when an ant-plant is potentially accessible to intruders, but less necessary when the ant colony is isolated by a protective wax barrier. Our results support the hypothesis that "selfish" defence against invasions is the major selective pressure that has led to the development and maintenance of pruning behaviour in weakly competitive plant-ants.     

No Experimental Evidence for Host Ant Related Oviposition in a Parasitic Butterfly

The ability of adult butterflies of the genus Maculinea to locate their host ants prior to oviposition has been the subject of much discussion. We studied the egg laying behavior of the dusky large blue Maculinea nausithous whose larvae parasitize colonies of the ant Myrmica rubra. Flowerheads of the initial food plant were sprinkled with soil from ant nests, which contain chemicals involved in the nest recognition behavior of ants. The experiment was conducted to determine whether ant-released chemicals may act as oviposition cues and whether intraspecific competition for suitable plants may force female butterflies to alternative decisions. Host plant choice was not influenced by the presence of nest-derived host-ant cues. Density dependent shifts to less suitable host plants could not be ascertained nor changes in egg laying behavior across the flight period. The observed egg distribution could be primarily explained by host plant characteristics and environmental variability among sites. The result confirms the theory that host ant dependent oviposition appears to be a disadvantageous strategy in the face of resource limitation within ant colonies and the immobility of caterpillars.