Foraging behavior of Brachygastra lecheguana (Hymenoptera: Vespidae) on Banisteriopsis malifolia (Malpighiaceae): Extrafloral nectar consumption and herbivore predation in a tending ant system

In the Brazilian savanna many plant species bear regular associations with patrolling ants that are aggressive towards insect herbivores. However, not only ants but also several species of predatory wasps are attracted to plants due to the extrafloral nectaries (EFNs). Such wasps feed on both herbivores and plant exudates. In this study we describe the foraging behavior of the social Polistinae wasp Brachygastra lecheguana in the extrafloral nectaried shrub Banisteriopsis malifolia, and investigated the influence of patrolling ants Camponotus blandus on the activity of the wasp. Brachygastra lecheguana fed on the endophytic larvae of Anthonomus (Curculionidae) beetles that developed inside flower buds. The wasp lacerated the bud layers to reach the beetle larvae located at the bud core. The wasp visits to Ba. malifolia were statistically related to the abundance of flower buds and beetles. Ant exclusion experiments revealed that the hunting behavior of B. lecheguana on beetles was not related to the absence of C. blandus. However we found that wasps spent more time consuming extrafloral nectar on branches where ants were excluded. This is the first study reporting extrafloral nectar consumption by B. lecheguana, as well as the predation on herbivores in natural areas. In cerrado vegetation, ants benefit the plant by reducing insect herbivores, and our study provides evidence that the B. lecheguana – Ba. malifolia system represents a potential interaction where the wasp may also benefit the host plant. The value of this wasp species as a plant‐guard is discussed.     

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.     

Ant-related oviposition is not associated to low parasitism of the myrmecophilous butterfly Allosmaitia strophius in an extrafloral nectaried shrub

In Lycaenidae-ant mutualisms, ovipositing females select plants based on the presence and/or species of ants in order to maximize survival rates of immatures. The ants are supposed to protect the immatures from parasitoids, but there is large variation in the protection provided. Here, we experimentally investigated whether the occurrence of the facultative myrmecophilous Allosmaitia strophius (the dominant species in our study system) was ant-related. The parasitism rates of immatures collected in the field and reared in the laboratory were also investigated. Stems of the extrafloral nectaried shrub Peixotoa tomentosa were designated as either ant-present (control) or absent (treated). The occurrence of A. strophius on ant-present stems was five times greater than on treated stems. Most eggs and larvae were associated with Camponotus blandus and Ectatomma tuberculatum, two aggressive ant species in the Brazilian savanna. Egg parasitism rate was 9%, and all the parasitized eggs were on ant-present stems. Pupal parasitism on ant-present and ant-absent stems was 25.6% and 7%, respectively. The higher parasitism rate in the presence of ants might also have been density-dependent, because caterpillars were more abundant in ant-present stems. Tropical lycaenids are frequently found in association with patrolling ants. Nevertheless, there is growing evidence that parasitism is higher in the presence of ants, owing to caterpillar's density-dependent effects in plants with ants, and/or to the weak lycaenid-ant associations. This indicates that the offspring of myrmecophilous lycaenids may not benefit, at least in terms of lower parasitism, by living with ants.     

Ants,plant characteristics and habitat conservation status affect the occurrence of myrmecophilous butterflies on an extrafloral nectaried Malpighiaceae

We investigated the occurrence of myrmecophilous florivorous lycaenid larvae in Banisteriopsis malifolia (Malpighiaceae) according to habitat conservation status (disturbed and preserved savanna), plant phenology, height and the presence of tending ants. The abundance and richness of lycaenids were sixfold and fivefold greater, respectively, in the disturbed area than in the protected savanna. Lycaenids occurred mostly on plants visited by Camponotus blandus, a mutualistic partner of larvae. Habitat type was the main factor influencing lycaenid occurrence, as plants in open areas offered more food resources and tending ants. Banisteriopsis malifolia was considered useful as a host for lycaenids in disturbed sites.     

Fire triggers the activity of extrafloral nectaries,but ants fail to protect the plant against herbivores in a neotropical savanna

Herbivores are attracted to young shoots and leaves because of their tender tissues. However, in extrafloral nectaried plants, young leaves also attract patrolling ants, which may chase or prey on herbivores. We examined this scenario in extrafloral nectaried shrubs of Banisteriopsis malifolia resprouting after fire, which promoted both the aseasonal production of leaves and the activity of extrafloral nectaries (EFNs). Results were compared between resprouting (burned) and unburned control plants. The aggressive ant species Camponotus crassus and the herbivorous thrips Pseudophilothrips obscuricornis were respectively rapidly attracted to resprouting plants because of the active EFNs and their less sclerophyllous leaves. The abundance of these insects was almost negligible in the control (unburned) shrubs. Ants failed to protect B. malifolia, as no thrips were preyed upon or injured by ants in resprouting plants. Consequently, on average, 37 % of leaves from resprouting shrubs had necrosis marks. Upon contact with ants, thrips released small liquid droplets from their abdomen, which rapidly displaced ants from the surroundings. This study shows that P. obscuricornis disrupted the facultative mutualism between C. crassus and B. malifolia, since ants received extrafloral nectar from plants, but were unable to deter herbivore thrips.     

Should aphids attract or repel ants? Effect of rival aphids and extrafloral nectaries on ant–aphid interactions

Among plants and herbivores, two types of conflicts occur in relation to mutualism with ants: one is competition for ant mutualism among myrmecophilous herbivores and plants, and the other is the conflict whether to attract or repel ants between myrmecophiles and nonmyrmecophiles that are damaged by ants. We investigated the extent to which two species of aphids (Megoura crassicauda and Aphis craccivora) and extrafloral nectaries on their host plant (Vicia faba var. minor) interact with one another for their relationships with ants. We designed an experiment where ants can choose to visit seedlings colonized by (1) M. crassicauda, (2) A. cracivora, (3) both aphid species, or (4) neither aphid species. Ants preferred A. craccivora to extrafloral nectaries and avoided tending M. crassicauda. We also analyzed the population growth of each aphid when it coexists with (1) ants, (2) the other aphid species, (3) ants and the other aphid species, or (4) neither of them. Under ant-free conditions, we detected an exploitative competition between the two aphid species. The ants had no significant effect on the population of A. craccivora, whereas they had negative effects on the population growth of M. crassicauda by attacking some individuals. When both aphids coexisted, M. crassicauda suffered ant attack more intensely because A. craccivora attracted more ants than extrafloral nectaries despite ant-repelling by M. crassicauda. On the other hand, the ants benefited A. craccivora by eliminating its competitor. To avoid ant attack, aphids may have been selected either to be more attractive to ants than other sympatric sugar sources or to repel the ants attracted to them. We hypothesize that competition among sympatric sugar sources including rival aphids and extrafloral nectaries is a factor restricting aphids to be myrmecophilous. Received: January 17, 2000 / Accepted: July 4, 2000     

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.     

On the inability of ants to protect their plant partners and the effect of herbivores on different stages of plant reproduction

Ants co‐occur with herbivorous thrips in several tropical plants, but their interactions are largely unexplored. Should thrips be deterred by ants, a positive effect of ants on plant fitness might be expected. Here, by using an experimental study design with ant‐present and ant‐excluded treatments, we investigated the influence of Camponotus blandus on Pseudophilothrips obscuricornis abundance and herbivory in three extrafloral nectaried species: Banisteriopsis malifolia, B. laevifolia and B. stellaris. In addition, we examined the effect of thrips herbivory on flower set and fruit development and dispersion. Thrips abundance and herbivory were higher on ant‐present stems of B. malifolia and B. laevifolia, where thrips managed to escape from ants by hiding in between clusters of flower buds (thygmotaxis behaviour). In B. stellaris the results were the opposite, as flower bud clusters did not offer hiding places, so thrips were unable to hide from ants; thus both thrips abundance and herbivory were lower on ant‐present stems. Thrips herbivory had no significant effect on flower and fruit set, but samaras (V‐shaped winged fruits of Malpighiaceae) attacked by thrips presented severe distortions and asymmetries. This caused damaged fruits to be dispersed closer to the mother plant, whereas uninjured fruits were dispersed further away. This study is evidence that ant–plant–herbivore systems have variable outcomes depending on the species involved, their behaviour and the plant structure under consideration. Unlike other herbivores, thrips negatively influence the very last stage of plant reproduction. The minute and furtive herbivorous thrips have long been ignored in natural systems, but because of their wide host range, they may be important herbivores even in extrafloral nectaried plants, which are usually fiercely protected by ants.     

Testing for mycorrhizal fungi-plant-ant indirect effects

Abstract

Recent work has demonstrated indirect effects between mycorrhizal fungi and insect herbivores and pollinators. The existence of indirect effects between mycorrhizal fungi and protection-for-food mutualists, such as extrafloral nectar-foraging ‘bodyguard ants’, is unknown. In this study, we examined the potential for indirect effects of arbuscular mycorrhizal fungi on aggressive ant bodyguards, mediated by changes in the expression of extrafloral nectaries of a shared host plant. We found that mycorrhizal plants grew larger and produced more extrafloral nectaries compared to their non-mycorrhizal counterparts. The difference in the number of nectaries between mycorrhizal and non-mycorrhizal plants, however, was too small to elicit differences in ant attendance. In spite of the lack of a significant indirect effect of mycorrhizal fungi on ant attendance, mycorrhizal plants suffered damage to a significantly greater proportion of their leaves compared to non-mycorrhizal plants. This result likely stems from other (non-ant-mediated) indirect effects of mycorrhizal fungi on herbivores.     

Extrafloral nectaries as a deterrent mechanism against seed predators in the chemically protected weed Crotalaria pallida (Leguminosae)

Abstract In several plants, extrafloral nectaries (EFN) are located close to the reproductive structures, suggesting that ants may act as a defence against specialized seed predators that overcome chemical defences. Alternatively, ants may also deter herbivores in a generalized manner, thereby protecting the whole plant. In this work, we examined the relationship between the chemically protected weed Crotalaria pallida Ait. (Leguminosae) that bears EFN, its specialized seed predator, the larvae of the arctiid moth Utetheisa ornatrix L. (Arctiidae) and ants. We tested two hypotheses related to the type of deterrence caused by ants. The Seed Predator Deterrence Hypothesis predicts that ant deterrence is directed primarily towards herbivores that destroy seeds and other reproductive structures, without attacking herbivores on vegetative structures. The General Deterrence Hypothesis states that ants are general in their effects, equally deterring herbivores in vegetative and reproductive structures. Our results supported the predictions of the Seed Predator Deterrence Hypothesis, namely, that (i) ant activity on EFN was related to the vulnerability of reproductive structures to attack by U. ornatrix; (ii) ant patrolling was restricted almost entirely to racemes; (iii) ants removed termites used as baits more frequently on racemes than on leaves; and (iv) U. ornatrix larvae were often expulsed from the racemes. These results indicate that EFN can act as another deterrent mechanism in chemically protected plants by promoting the expulsion of specialist seed predators.     

Induced Floral and Extrafloral Nectar Production Affect Ant‐pollinator Interactions and Plant Fitness

Thousands of plant species throughout tropical and temperate zones secrete extrafloral nectar to attract ants, whose presence provides an indirect defense against herbivores. Extrafloral nectaries are located close to flowers and may modify competition between ants and pollinators. Here, we used Lima bean (Phaseolus lunatus L.) to study the plants interaction between ants and flower visitors and its consequences for plant fitness. To test these objectives, we carried out two field experiments in which we manipulated the presence of ants and nectar production via induction with jasmonic acid (JA). We then measured floral and extrafloral nectar production, the number of patrolling ants and flower visitors as well as specific plant fitness traits. Lima bean plants under JA induction produced more nectar in both extrafloral nectaries and flowers, attracted more ants and produced more flowers and seeds than non‐induced plants. Despite an increase in floral nectar in JA plants, application of this hormone had no significant effects on flower visitor attraction. Finally, ant presence did not result in a decrease in the number of visits, but our results suggest that ants could negatively affect pollination efficiency. In particular, JA‐induced plants without ants produced a greater number of seeds compared with the JA‐treated plants with ants.     

When are ant-attractant devices a worthwhile investment? Vicia faba extrafloral nectaries and Lasius niger ants

Most studies aiming to determine the beneficial effect of ants on plants simply consider the effects of the presence or exclusion of ants on plant yield. This approach is often inadequate, however, as ants interact with both non-tended herbivores and tended Homoptera. Moreover, the interaction with these groups of organisms is dependent on ant density, and these functional relationships are likely to be non-linear. A model is presented here that segregates plant herbivores into two categories depending on the sign of their numerical response to ants (myrmecophiles increase with ants, non-tended herbivores decline). The changes in these two components of herbivores with increasing ant density and the resulting implications for ant-plant mutualisms are considered. It emerges that a wide range of ant densities needs to be considered as the interaction sign (mutualism or parasitism) and strength is likely to change with ant density. The model is used to interpret the results of an experimental study that varied levels of Aphis fabae infestation and Lasius niger ant attendance on Vicia faba bean plants. Increasing ant density consistently reduced plant fitness and thus, in this location, the interaction between the ants and the plant can be considered parasitic. In the Vicia faba system, these costs of ants are unlikely to be offset by other beneficial agents (e.g., parasitoids), which also visit extrafloral nectaries.     

Herbivory-induced extrafloral nectar increases native and invasive ant worker survival

Ascertaining the costs and benefits of mutualistic interactions is important for predicting their stability and effect on community dynamics. Despite widespread designation of the interaction between ants and extrafloral nectaries (EFNs) as a mutualism and over 100 years of studies on ant benefits to plants, the benefits to ants have never been experimentally quantified. The success of invasive ants is thought to be linked to the availability of carbohydrate-rich resources, though reports of invasive ant visits to EFNs are mixed. In two laboratory experiments, we compared worker survival of one native (Iridomyrmex chasei) and two invasive ant species (Linepithema humile and Pheidole megacephala) exposed to herbivorized or non-herbivorized EFN-bearing plants (Acacia saligna) or positive and negative controls. We found that non-herbivorized plants did not produce any measurable extrafloral nectar, and ants with access to non-herbivorized plants had the same survival as ants with access to an artificial plant and water (unfed ants). Ants given herbivorized plants had 7–11 times greater worker survival relative to unfed ants, but there were no differences in survival between native and invasive ants exposed to herbivorized plants. Our results reveal that ants cannot induce A. saligna extrafloral nectar production, but workers of both native and invasive ant species can benefit from extrafloral nectar as much as they benefit from sucrose.     

FACULTATIVE MUTUALISM BETWEEN WEEDY VETCHES BEARING EXTRAFLORAL NECTARIES AND WEEDY ANTS IN CALIFORNIA

Vetches (Vicia spp.) were studied in the San Francisco Bay Area of California in the spring of 1978. The stipular nectaries of the vetches are visited by the Argentine ant, Iridomyrmex humilis Mayr. The nectaries were removed to exclude ants in controlled experiments to determine if these ants protect the vetches from herbivores or seed predators. Plants with excised nectaries suffered substantially greater damage to their foliage than control plants, indicating that ants protect the foliage. There was no indication that ants protect the vetches from seed predators, but fruit set was substantially lower in plants with excised nectaries. Analysis of sugar and amino acid composition of extrafloral nectar served as a basis for feeding tests with Argentine ants by using artificial nectar solutions. Ants preferred sucrose and glucose solutions over fructose. They showed no preference for any one sugar mixture over another, nor did they exhibit differential recruitment to artificial nectar solutions containing only sugars or sugars and amino acids.     

Ant foraging on extrafloral nectaries of Qualea grandiflora (Vochysiaceae) in cerrado vegetation: ants as potential antiherbivore agents

Summary Qualea grandiflora is a typical tree of Brazilian cerrados (savanna-like vegetation) that bears paired extrafloral nectaries (EFNs) along its stems. Results show that possession of EFNs increases ant density on Q. grandiflora shrubs over that of neighbouring non-nectariferous plants. Frequency of ant occupancy and mean number of ants per plant were much higher on Qualea than on plants lacking EFNs. These differences resulted in many more live termitebaits being attacked by foraging ants on Qualea than on neighbours without EFNs. Termites were attacked in equal numbers and with equal speeds on different-aged leaves of Qualea. The greatest potential for herbivore deterrence was presented by Camponotus ants (C. crassus, C. rufipes and C. aff. blandus), which together attacked significantly more termites than nine other ant species grouped. EFNs are regarded as important promoters of ant activity on cerado plants.     

The function of extrafloral nectaries in the pollination and reproduction of Sambucus javanica

Sambucus javanica is a perennial herb with extrafloral nectaries (EFNs) on its inflorescences. To explore the ecological functions of EFNs, a factorial combination experiment of ant (access or exclusion) and EFNs (with or without) at the plant level was created in two populations. The role of EFNs in the attraction of ants and flying pollinators, the defensive role of ants against foliar herbivores, the effects of ants on pollinator visitation and the effects of ant–pollinator interactions on fruit production in one or both populations were assessed. Ants were common on the ant-access plants with EFNs, but absent from the ant-access plants without EFNs. Foliar herbivory was independent of ant and EFN treatments and their interactions. The visitation frequency of flying pollinators (honeybees and syrphid flies) and fruit set were significantly higher for plants with EFNs than plants without EFNs, but were not affected by ant treatments or their interactions with EFN treatments. These results suggest that EFNs in S. javanica attracted both ants and flying pollinators, but ants did not present a defensive role against herbivores, did not deter flying pollinators from visiting inflorescences and had no effects on fruit production. In addition, ants were not significant pollen vectors.     

The influence of ants on patterns of colonization and establishment within a set of coexisting lycaenid butterflies in a south-east Asian tropical rain forest

In Peninsular Malaysia ten species of lycaenid butterflies use leaf flushes or inflorescences of the legume tree Saraca thaipingensis as larval hostplant. Resource partitioning among these species is regulated by a complex mixture of patterns of interaction with ants. Females of obligately myrmecophilous species lay their eggs exclusively on trees colonized by their specific host ants. On trees colonized by weaver ants, only specialist mutualists adapted to these territorial ants are able to survive, while larvae of other species are killed. The formicine ant Cladomyrma petalae, which inhabits hollow twigs of the myrmecophytic hostplant, likewise precludes oviposition by female butterflies. Lycaenid larvae confronted with this ant species never survive, but one concealed feeding species (Jamides caeruleus) escapes removal due to the cryptic life-habits of the larvae. Two facultative myrmecophiles associate in a mutualistic way with a wide and largely overlapping range of ant genera which forage at the extrafloral nectaries of leaf flushes. One species (Cheritra freja) is not myrmecophilous, but is tolerated by all but the most territorial ants. Ant-dependent hostplant selection and egg-clustering characterize the obligate mutualists, whereas facultative myrmecophiles and the non-myrmecophile distribute their eggs singly over appropriate hostplants. Signals mediating caterpillar-ant communication are highly specialized in one obligate myrmecophile (Drupadia theda), but rather unspecific in four other species tested. Altogether our observations indicate that colonization and establishment of lycaenid butterflies on S. thaipingensis trees are governed by specializations as well as opportunistic use of resources (ants and hostplant parts). Therefore, the diversity of this species assemblage is maintained by deterministic as well as stochastic factors.     

Ants on plants: a meta-analysis of the role of ants as plant biotic defenses

We reviewed the evidence on the role of ants as plant biotic defenses, by conducting meta-analyses for the effects of experimental removal of ants on plant herbivory and fitness with data pooled from 81 studies. Effects reviewed were plant herbivory, herbivore abundance, hemipteran abundance, predator abundance, plant biomass and reproduction in studies where ants were experimentally removed (n = 273 independent comparisons). Ant removal exhibited strong effects on herbivory rates, as plants without ants suffered almost twice as much damage and exhibited 50% more herbivores than plants with ants. Ants also influenced several parameters of plant fitness, as plants without ants suffered a reduction in biomass (−23.7%), leaf production (−51.8%), and reproduction (−24.3%). Effects were much stronger in tropical regions compared to temperate ones. Tropical plants suffered almost threefold higher herbivore damage than plants from temperate regions and exhibited three times more herbivores. Ant removal in tropical plants resulted in a decrease in plant fitness of about 59%, whereas in temperate plants this reduction was not statistically significant. Ant removal effects were also more important in obligate ant–plants (=myrmecophytes) compared to plants exhibiting facultative relationships with hemiptera or those plants with extrafloral nectaries and food bodies. When only tropical plants were considered and the strength of the association between ants and plants taken into account, plants with obligate association with ants exhibited almost four times higher herbivory compared to plants with facultative associations with ants, but similar reductions in plant reproduction. The removal of a single ant species increased plant herbivory by almost three times compared to the removal of several ant species. Altogether, these results suggest that ants do act as plant biotic defenses, but the effects of their presence are more pronounced in tropical systems, especially in myrmecophytic plants. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. N. P. de U. Barbosa, L. Diniz, Y. Oki and F. Pezzini contributed equally to this work and are listed in alphabetical order.     

The effect of disturbance on an ant–plant mutualism

Protective ant–plant mutualisms—where plants provide food or shelter to ants and ants protect the plants from herbivores—are a common feature in many ecological communities, but few studies have examined the effect of disturbance on these interactions. Disturbance may affect the relationship between plants and their associated ant mutualists by increasing the plants’ susceptibility to herbivores, changing the amount of reward provided for the ants, and altering the abundance of ants and other predators. Pruning was used to simulate the damage to buttonwood mangrove (Conocarpus erectus) caused by hurricanes. Pruned plants grew faster than unpruned plants, produced lower levels of physical anti-herbivore defenses (trichomes, toughness), and higher levels of chemical defenses (tannins) and extrafloral nectaries. Thus, simulated hurricane damage increased plant growth and the amount of reward provided to ant mutualists, but did not have consistent effects on other anti-herbivore defenses. Both herbivores and ants increased in abundance on pruned plants, indicating that the effects of simulated hurricane damage on plant traits were propagated to higher trophic levels. Ant-exclusion led to higher leaf damage on both pruned and upruned plants. The effect of ant-exclusion did not differ between pruned and unpruned plants, despite the fact that pruned plants had higher ant and herbivore densities, produced more extrafloral nectaries, and had fewer physical defenses. Another common predator, clubionid spiders, increased in abundance on pruned plants from which ants had been excluded. I suggest that compensatory predation by these spiders diminished the effect of ant-exclusion on pruned plants.     

The influence of extrafloral nectaries on parasitism of an insect herbivore

The extrafloral nectaries of many plants promote ant defense against insect herbivores. We examined the influence of extrafloral nectaries on the levels of parasitism of a generalist insect herbivore, the gypsy moth (Lymantria dispar L.). Larvae and pupae of the moth were collected from trees with and without extrafloral nectaries growing in the same forests in South Korea and reared to evaluate parasitism. More parasitism occurred on plants with extrafloral nectaries in seven of the nine season-long collections at the six sites and in four out of five collecting periods. Parasitism was higher on the four main genera of plants with extrafloral nectaries than on any of five main genera of plants without extrafloral nectaries. There was no difference in parasitoid richness; nine species occurred in each group, eight of which were the same. There was a positive and almost significant correlation between the abundance of plants with extrafloral nectaries and the parasitism of gypsy moth at the sites. Extrafloral nectaries may reduce herbivory by inducing more parasitism of the insect herbivores that attack plants bearing the glands.