Saving a tropical ecosystem from a destructive ant-scale (Pheidole megacephala, Pulvinaria urbicola) mutualism with support from a diverse natural enemy assemblage

Ants can disrupt the natural biological control of serious hemipteran pests by interfering with natural enemies, resulting in a change in ecosystem functioning. We focus here on interference by a highly invasive ant Pheidole megacephala on the regulation of a tree destroying hemipteran scale insect Pulvinaria urbicola on Cousine Island in the Seychelles archipelago, a tropical island ecosystem. We show how a diverse natural enemy assemblage contributes substantially to the collapse of the ant-scale mutualism following managed ant suppression. Natural enemy abundance and species richness increased significantly after ant suppression, with varying responses among the different functional guilds. Primary parasitoids coexisted with tending ants before ant suppression, but could not regulate the enormously high scale densities alone. After ant suppression, a significant increase in predators caused a collapse of the scale population. Guilds external to the mutualism were also affected, with primary parasitoids of various non-hemipteran taxa also increasing, which contributed significantly to the recovery of the community to its pre-invasion composition. Our results highlight the far-reaching and pervasive effects of the hemipteran-tending invasive ant within the natural enemy assemblage. In turn, we also illustrate the potential to restore the tropical ecosystem by encouraging an array of natural enemies through precision management of the ant.     

First record of Pulvinaria urbicola Cockerell (Hemiptera: Coccidae), a potentially damaging scale insect,on Christmas Island,Indian Ocean

Pulvinaria urbicola Cockerell, a pulvinariine soft scale (Hemiptera: Coccidae), is a broad host-plant generalist, produces honeydew and is commonly tended by ants, including the invasive yellow crazy ant Anoplolepis gracilipes Smith and big-headed ant Pheidole megacephala (Fabricius). Pu. urbicola is implicated in dieback of forest dominated by Pisonia grandis (Nyctaginaceae) on many Indo-Pacific islands. Here we report detection of Pu. urbicola on Christmas Island (Indian Ocean), describe the potential impacts of the association of this trophobiont with introduced ants, and briefly outline biosecurity and management issues. On Christmas Island, Pu. urbicola represents a threat to stands of Pi. grandis, potentially threatens the dominant forest tree Pi. umbellifera, and could exacerbate supercolony formation and impacts of the yellow crazy ant.     

Strategic Management of an Invasive Ant‐scale Mutualism Enables Recovery of a Threatened Tropical Tree Species

Mutualisms between invasive ants and honeydew‐producing insects can have widespread negative effects on natural ecosystems. This is becoming an increasingly serious problem worldwide, causing certain ecosystems to change radically. Management of these abundant and influential mutualistic species is essential if the host ecosystem is to recover to its former non‐invaded status. This negative effect is particularly prevalent on some tropical islands, including Cousine Island, Seychelles. On this island, the invasive ant Pheidole megacephala has caused serious indirect damage to the threatened native Pisonia grandis trees via a mutualism with an invasive scale insect, Pulvinaria urbicola. We aimed to suppress the ant, thereby decoupling the mutualism and enabling recovery of the Pisonia trees. We treated all areas where ant pressure was high with a selective formicidal bait, which was deployed in custom‐made bait stations designed to avoid risk of treatment to endemic fauna. In the treated area, ant foraging activity was reduced by 93 percent and was followed by a 100 percent reduction in scale insect density. Abundance of endemic herbivorous insects and herbivorous activity increased significantly, however, after the decline in mutualistic species densities. Despite the native herbivore increase, there was considerable overall improvement in Pisonia shoot condition and an observed increase in foliage density. Our results demonstrate the benefit of strategic management of highly mutualistic alien species to the native Pisonia trees. It also supports the idea that area‐wide suppression is a feasible alternative to eradication for achieving positive conservation management at the level of the forest ecosystem.     

Invasive African big-headed ants, Pheidole megacephala, on coral cays of the southern Great Barrier Reef: distribution and impacts on other ants

Infestation of islands by exotic ants is widespread and increasing due to human activities throughout the world. Exotic ants, particularly the invasive African big-headed ant, Pheidole megacephala (Fabricius), are of great conservation concern for coral cays at the southern end of the Great Barrier Reef, Australia. Little is known, however, about the distribution and ecological impacts of invasive ants in this insular system. We surveyed the ants of 14 vegetated coral cays recording a total of 24 ant species, including at least nine exotics. Pheidole megacephala was by far the most abundant and widespread species, occurring on 11 of 14 islands, often in very large numbers. The inter-island distribution of P. megacephala was best explained by human activities, with frequently visited, and to a lesser degree disturbed islands, more likely to be infested. On large islands (≥?10?ha) P. megacephala exhibited distinct habitat preferences, occurring in significantly lower abundances within heavily-shaded Pisonia grandis forest in the centre of islands, compared to more open, fringing woodland or shrubland. On smaller islands (<10?ha) with less extensive Pisonia stands, P. megacephala penetrated throughout the forest where its abundance was similar to that in open woodland. Despite considerable differences in biotic (floristic composition) and abiotic factors (e.g. island size) as well as the spatial configuration among islands, the severity of infestation by P. megacephala best explained variation in species richness, abundance and assemblage composition of other ants. We suggest a number of strategies to manage P. megacephala infestations on these islands.     

Competition hierarchies among ants and predation by birds jointly determine the strength of multi‐species ant–aphid mutualisms

Protection mutualisms often involve multiple species of protector that vary in quality as mutualists. Because protectors may compete for access to mutualists, concordance between competitive ability and degree of benefit will determine the overall strength of multi‐species mutualisms. We compared the abilities of two similarly sized congener ants as competitors for, and mutualists of pine‐feeding aphids, and how insectivorous birds affected each ant–aphid mutualism. Formica planipilis and F. podzolica were indistinguishable in forager abundance, but the former was 13‐fold more abundant at competition baits and provided 11‐fold more benefits to aphids. These results highlight how, in a single environment, a great ecological distance can exist between two congener ants of similar size. Insectivorous birds disrupted the two mutualisms to a similar extent, reducing aphid and ant abundance by 91% and 39% respectively. Nevertheless, birds had an important influence on the relative benefits of the two ants to aphids: where F. planipilis consistently benefited aphids, F. podzolica only did so in the absence of birds. Consequently, the presence of insectivorous birds and ant species identity jointly determined whether ant–aphid mutualisms occurred in pine canopies and the strength of such interactions. Our study highlights the inter‐relatedness of competition, predation and mutualism, and how competition can serve to strengthen mutualism by filtering inferior mutualists.     

Pisonia grandis monocultures limit the spread of an invasive ant—a case of carbohydrate quality?

The mechanisms by which invasive species are able to spread into and dominate natural communities are poorly understood and remain a focus of invasion research. In this quest, studying invasions that are limited by a controlling factor will be more informative than will studies documenting unabated spread and impacts. Some ant species are very successful invaders, and research demonstrating abiotic and biotic factors limiting their success has aided the understanding of invasion ecology. We report here a study showing the highly invasive African big headed ant Pheidole megacephala having a novel distribution on coral cays within Australia’s Great Barrier Reef. These patterns displayed a clear limitation of its distribution with monocultures of the tree Pisonia grandis. This distribution was contrary to the known environmental limitations of the ant, and the limitation could not be associated with an underlying abiotic determinant of the vegetation type. We present these distributional patterns, and following consideration of all known biotic and abiotic limitations of ant invasions we discuss the potential that the peculiar ecophysiology of P. grandis is the causal factor. Specifically, we suggest that the quality of carbohydrate supply to ants is a limitation to invasive spread in much the same way that carbohydrate quantity is known to affect ant population densities in other ecosystems.     

Impact of an introduced ant on native rain forest invertebrates: Pheidole megacephala in monsoonal Australia

Pheidole megacephala is an exotic ant species that has severely affected native invertebrate biodiversity throughout the tropics. Its impacts have been documented extensively in relatively depauperate invertebrate communities, but not in species-rich habitats such as tropical rain forests. Here we describe the local distribution of P. megacephala and its impacts on native invertebrate assemblages in and around a rain forest patch at Howard Springs, in Australia's monsoonal tropics. P. megacephala was found to be confined to a single area of approximately 25 ha, with its distribution centered on drainage lines and the rain forest. Significant but weak correlations were found between its abundance and vegetative canopy cover (positive) and distance from the rain forest (negative). In the most heavily infested area within the rain forest, the abundance of P. megacephala was 37–110 times that of total native ant abundance found within uninfested plots, as measured by pitfall traps. The abundance and richness of native ants and other invertebrates were significantly reduced in litter samples, pitfall catches and foliage beats where P. megacephala was present, inversely relative to the abundance of P. megacephala. Only two individuals of a single native ant species were found within the most infested plot, with native ant richness being reduced to about half in the least infested plot. The most persistent functional groups of native ants in infested plots were Cryptic species, which forage primarily within soil and leaf litter, and Opportunists, which exhibit highly generalised foraging behaviour. The highest abundance of P. megacephala corresponded with a 42–85% decrease in the abundance of other native invertebrates. Insect larvae were totally absent from foliage beats collected at the most heavily infested plot. P. megacephala was found overall to be expanding its range, averaging 12 m range expansion in the dry season and contracting 7 m in the wet season. It is able to spread into surrounding savanna habitats by occupying relatively sheltered microsites, such as beneath logs and at the bases of trees. However, it is unlikely to attain high population densities in open savanna habitats because of its relative intolerance of desiccation, and the prevalence of behaviourally dominant native ant species. Howard Springs is currently the only rain forest patch in monsoonal Australia known to be infested by P. megacephala, but clearly this ant is a serious potential threat to the region's rain forest invertebrate fauna. Received: 19 August 1998 / Accepted: 12 May 1999     

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.     

Mutualisms and population regulation: mechanism matters

For both applied and theoretical ecological science, the mutualism between ants and their hemipteran partners is iconic. In this well-studied interaction, ants are assumed to provide hemipterans protection from natural enemies in exchange for nutritive honeydew. Despite decades of research and the potential importance in pest control, the precise mechanism producing this mutualism remains contested. By analyzing maximum likelihood parameter estimates of a hemipteran population model, we show that the mechanism of the mutualism is direct, via improved hemipteran growth rates, as opposed to the frequently assumed indirect mechanism, via harassment of the specialist parasites and predators of the hemipterans. Broadly, this study demonstrates that the management of mutualism-based ecosystem services requires a mechanistic understanding of mutualistic interactions. A consequence of this finding is the counter intuitive demonstration that preserving ant participation in the ant-hemipteran mutualism may be the best way of insuring pest control.     

Invasional 'meltdown' on an oceanic island

Islands can serve as model systems for understanding how biological invasions affect community structure and ecosystem function. Here we show invasion by the alien crazy ant Anoplolepis gracilipes causes a rapid, catastrophic shift in the rain forest ecosystem of a tropical oceanic island, affecting at least three trophic levels. In invaded areas, crazy ants extirpate the red land crab, the dominant endemic consumer on the forest floor. In doing so, crazy ants indirectly release seedling recruitment, enhance species richness of seedlings, and slow litter breakdown. In the forest canopy, new associations between this invasive ant and honeydew‐secreting scale insects accelerate and diversify impacts. Sustained high densities of foraging ants on canopy trees result in high population densities of host‐generalist scale insects and growth of sooty moulds, leading to canopy dieback and even deaths of canopy trees. The indirect fallout from the displacement of a native ‘keystone’ species by an ant invader, itself abetted by introduced/cryptogenic mutualists, produces synergism in impacts to precipitate invasional ‘meltdown’ in this system.     

Invasive Argentine ants reduce fitness of red maple via a mutualism with an endemic coccid

Many invasive ant species form mutualisms with honeydew-producing Hemiptera and their aggressive presence deters the natural enemies of the Hemiptera. Invasive ant species like the Argentine ant have often been associated with hemipteran outbreaks in urban, agricultural and natural ecosystems. We investigated the effects of a mutualism between the invasive Argentine ant and the endemic terrapin scale on coccid density and the fitness of the host of this mutualism, the endemic red maple, situated in a commercial park. The terrapin scale has numerous natural enemies and we predicted that the high terrapin scale numbers associated with tending Argentine ants would collapse once Argentine ants were excluded from the host tree canopy. We predicted that excluding the Argentine ant from the tree canopy would result in an indirect net fitness benefit to the host. Terrapin scale numbers collapsed when Argentine ants were excluded from the host tree canopy. Red maples with Argentine ants excluded from their canopy had higher seed mass and larger early leaves indicating that this invasive ant-endemic scale mutualism imposed a net fitness cost to the host tree. The Argentine ant has yet to invade closed-canopy forest within its introduced range. The red maple is common in adjacent closed-canopy forest fragments and recent work has shown that invasion of these forest fragments by the Argentine ant is limited by a steady carbohydrate resource. We discuss the implications to forest invasion posed by a mutualism involving the Argentine ant and an endemic coccid.     

Disruption of ant-seed dispersal mutualisms by the invasive Asian needle ant (<Emphasis Type="Italic">Pachycondyla chinensis</Emphasis>)

By disrupting the structure of native ant assemblages, invasive ants can have effects across trophic levels. Most studies to date, however, have focused on the impacts just two species (Linepithema humile and Solenopsis invicta). The impacts of many other invasive ant species on ecological processes in their introduced range are unknown. In this study we tested the hypothesis that the invasive ant Pachycondyla chinensis disrupts ant-seed dispersal mutualisms by displacing native ant species, especially the keystone mutualist Aphaenogaster rudis, while failing to disperse seeds itself. In a paired design we measured the impact of P. chinensis on the native ant-plant seed dispersal mutualism. The number of A. rudis workers was 96% lower in invaded than in intact plots, and the number of seeds removed was 70% lower in these plots. Finally, in invaded plots the abundance of Hexastylis arifolia, a locally abundant myrmecochorous plant, was 50% lower than in plots where P. chinensis was absent. A parsimonious interpretation of our results is that P. chinensis causes precipitous declines in the abundance of A. rudis within invaded communities, thereby disrupting the ant-plant seed dispersal mutualisms and reducing abundances of ant-dispersed plants. In sum, the magnitude of the effects of P. chinensis on seed dispersal is quantitatively similar to that documented for the intensively studied invasive Argentine ant. We suggest that more studies on the impacts of less-studied invasive ant species on seed dispersal mutualisms may increase our knowledge of the effects of these invaders on ecosystem function.     

A facultative mutualism between aphids and an invasive ant increases plant reproduction

1. The consequences to plants of ant–aphid mutualisms, particularly those involving invasive ants, are poorly studied. Ant–aphid mutualisms may increase or decrease plant fitness depending on the relative cost of herbivory by ant‐tended aphids versus the relative benefit of increased ant suppression of other (non‐aphid) herbivores. 2. We conducted field and greenhouse experiments in which we manipulated the presence and absence of cotton aphids (Aphis gossypii) on cotton plants to test the hypothesis that a mutualism between cotton aphids and an invasive ant, the red imported fire ant (Solenopsis invicta), benefits cotton plants by increasing fire ant suppression of caterpillars. We also manipulated caterpillar abundance to test whether the benefit of the mutualism varied with caterpillar density. 3. We found that more fire ants foraged on plants with cotton aphids than on plants without cotton aphids, which resulted in a significant reduction in caterpillar survival and caterpillar herbivory of leaves, flower buds, and bolls on plants with aphids. Consequently, cotton aphids indirectly increased cotton reproduction: plants with cotton aphids produced 16% more bolls, 25% more seeds, and 10% greater seedcotton mass than plants without aphids. The indirect benefit of cotton aphids, however, varied with caterpillar density: the number of bolls per plant at harvest was 32% greater on plants with aphids than on plants without aphids at high caterpillar density, versus just 3% greater at low caterpillar density. 4. Our results highlight the potential benefit to plants that host ant–hemipteran mutualisms and provide the first experimental evidence that the consequences to plants of an ant–aphid mutualism vary at different densities of non‐aphid herbivores.     

Elevated temperatures alter an ant‐aphid mutualism

Ant‐hemipteran mutualisms are keystone interactions that can be variously affected by warming: these mutualisms can be strengthened or weakened, or the species can transition to new mutualist partners. We examined the effects of elevated temperatures on an ant‐aphid mutualism in the subalpine zone of the Rocky Mountains in Colorado, USA. In this system, inflorescences of the host plant, Ligusticum porteri Coult. & Rose (Apiaceae), are colonized by the ant‐tended aphid Aphis asclepiadis Fitch or less frequently by the non‐ant tended aphid Cavariella aegopodii (Scopoli) (both Hemiptera: Aphididae). Using an 8‐year observational study, we tested for two key mechanisms by which ant‐hemipteran mutualisms may be altered by climate change: shifts in species identity and phenological mismatch. Whereas the aphid species colonizing the host plant is not changing in response to year‐to‐year variation in temperature, we found evidence that a phenological mismatch between ants and aphids could occur. In warmer years, colonization of host plant inflorescences by ants is decreased, whereas for A. asclepiadis aphids, host plant colonization is mostly responsive to date of snowmelt. We also experimentally established A. asclepiadis colonies on replicate host plants at ambient and elevated temperatures. Ant abundance did not differ between aphid colonies at ambient vs. elevated temperatures, but ants were less likely to engage in tending behaviors on aphid colonies at elevated temperatures. Sugar composition of aphid honeydew was also altered by experimental warming. Despite reduced tending by ants, aphid colonies at elevated temperatures had fewer intraguild predators. Altogether, our results suggest that higher temperatures may disrupt this ant‐aphid mutualism through both phenological mismatch and by altering benefits exchanged in the interaction.     

Influence of native ants on arthropod communities in a vineyard

• 1 Ants can have a range of effects on arthropods in crops, including suppressing herbivores such as caterpillars. However, ants can also increase hemipteran densities while reducing natural enemy numbers. In vineyard ecosystem, the effects of native ants and their interactions with other arthropods are poorly understood.
• 2 An ant‐exclusion experiment was designed to test the impact of native ants on both canopy and ground arthropods concurrently. The potential influence of ants on predation and parasitism of light brown apple moth (LBAM) eggs, a grape pest, was also examined. Adult grapevine scale insects and earwigs under bark were counted after a season of ant‐exclusion.
• 3 Among 23 ground ant species collected, six were found to forage in the canopy, with two Iridomyrmex species being the most commonly encountered.
• 4 There was no difference in the abundance of most arthropod orders and feeding groups between ant‐excluded and control vines, although ground spiders were more abundant under ant‐excluded vines, despite increased ground ant foraging pressure. LBAM egg parasitism and predation were low and probably affected by weather and other arthropods. Ant exclusion did not reduce survival of scale insects, although the distribution and abundance of scale insects were negatively associated with earwigs.
• 5 In conclusion, native ants did not consistently suppress arthropod assemblages, including natural enemies, and they did not promote the survival of scale insects. Interactions among native ant species within a vineyard might minimize their effects on other arthropods, although this needs further study.

     

An invasion revisited: the African big-headed ant (<Emphasis Type="Italic">Pheidole megacephala</Emphasis>) in northern Australia

Long-term studies provide the best information for invasion ecology as multi-temporal sampling can illuminate invasion dynamics, such as population changes and rate of spread. In 1996, Hoffmann et al. surveyed an infestation of African big-headed ant, Pheidole megacephala, in semi-natural rainforest of northern Australia. Here, we re-survey this infestation 9 years later to assess the dynamics of this invasion against the key finding of the initial study. Importantly, we re-sample a site sampled prior to invasion to demonstrate the causal link between an infestation and changes in a native invertebrate community. We found the area infested had almost doubled, and P. megacephala biomass in infested sites was up to 18 times greater than that of native ants in uninfested sites. In the two sites with the youngest infestations in 1996, P. megacephala abundance had increased more than 20-fold than that measured in 1996. Native ant abundance and species richness in infested sites were notably lower in 2005, with only one native ant specimen found in the most recently infested site. The abundance of other macro-invertebrates was the lowest in the three oldest infested sites. Coleoptera and Orthoptera were less abundant in infested sites. This study supports the findings of the first ‘snapshot’ study, and has shown that the area infested has almost doubled, populations are increasing, and ecological impacts remain severe. It reinforces that P. megacephala represents a serious ecological threat, and we argue that there is a greater need for management of this ant globally.     

The Effects of Invasive Ants on Prospective Ant Mutualists

Ants are recognized for their abilities both to engage in mutualistic interactions with diverse taxa, and to invade and dominate habitats outside their native geographic range. Here, we review the effects of invasive ants on three guilds of mutualists: ant-dispersed plants, ant-tended arthropods, and ant-tended plants. We contrast how those three guilds are affected by invasions, how invasive ants differ from native ants in their interactions with those guilds, and how the seven most invasive ant species differ amongst themselves in those interactions. Ant-dispersed plants typically suffer from interactions with invasive ants, a result we attribute to the small size of those ants relative to native seed-dispersing ants. Effects on the ant-tended arthropods and plants were more frequently positive or non-significant, although it is unclear how often these interactions are reciprocally beneficial. For example, invasive ants frequently attack the natural enemies of these prospective mutualists even in the absence of rewards, and may attack those prospective mutualists. Many studies address whether invasive ants provide some benefit to the partner, but few have asked how invasives rank within a hierarchy of prospective mutualists that includes other ant species. Because ant invasions typically result in the extirpation of native ants, this distinction is highly relevant to predicting and managing the effects of such invasions. Interspecific comparisons suggest that invasive ants are poorer partners of ant-dispersed plants than are most other ants, equally effective partners of ant-tended arthropods, and perhaps better partners of ant-tended plants. Last, we note that the invasive ant taxa differ amongst themselves in how they affect these three mutualist guilds, and in how frequently their interactions with prospective mutualists have been studied. The red imported fire ant, Solenopsis invicta, appears particularly likely to disrupt all three mutualistic interactions, relative to the other six invasive species included in this review.     

Population growth of Aphis gossypii and Myzus persicae (Hemiptera: Aphididae) in the presence of Linepithema humile and Tapinoma sessile (Hymenoptera: Formicidae)

Invasive ant species can have dramatic impacts on native ants, through direct predation and by usurping common resources. Most invasive ants and many native ants use honeydew, produced by phloem-sucking hemipterans. Because colonies of invasive ants can become very large after establishment, these ants may facilitate greater hemipteran trophobiont population growth compared with their sympatric native ant counterparts. We examined the population growth of an aphid mutualist, Aphis gossypii, and a nonmutualist, Myzus persicae, exposed to two Dolichoderine ants, Linepithema humile, a globally widespread invasive species, and Tapinoma sessile, a widespread co-occurring native ant, in North America in an enemy-free laboratory study. L. humile worker foraging activity was at least twice that of T. sessile, and populations of the myrmecophile, A. gossypii, were greater when exposed to L. humile than T. sessile, possibly caused, in part, by more frequent encounters with L. humile. L. humile ignored M. persicae when A. gossypii was absent, whereas T. sessile preyed on it. Both ant species preyed on M. persicae when A. gossypii was also present. This suggested that both ants may assess nutritional gains from aphid species (i.e., honeydew versus body tissue), eliminating less productive aphids competing for host plant space. Through their impact on populations of hemipteran mutualists, we suggest that colonies of L. humile and perhaps other invasive ants may acquire more honeydew than native ants, thereby fueling colony growth that leads to numerical dominance and widespread success in introduced environments.     

Impacts of the invasive Argentine ant on native ants and other invertebrates in coastal scrub in south‐eastern Australia

Invasive ants threaten native biodiversity and ecosystem function worldwide. Although their principal direct impact is usually the displacement of native ants, they may also affect other invertebrates. The Argentine ant, Linepithema humile (Dolichoderinae), one of the most widespread invasive ant species, has invaded native habitat where it abuts peri‐urban development in coastal Victoria in south‐eastern Australia. Here we infer impacts of the Argentine ant on native ants and other litter and ground‐dwelling invertebrates by comparing their abundance and taxonomic composition in coastal scrub forest either invaded or uninvaded by the Argentine ant. Species composition of native ants at bait stations and extracted from litter differed significantly between Argentine ant‐invaded and uninvaded sites and this was consistent across years. Argentine ants had a strong effect on epigeic ants, which were either displaced or reduced in abundance. The native ant Rhytidoponera victoriae (Ponerinae), numerically dominant at uninvaded sites, was completely absent from sites invaded by the Argentine ant. However, small hypogeic ants, including Solenopsis sp. (Myrmicinae) and Heteroponera imbellis (Heteroponerinae), were little affected. Linepithema humile had no detectable effect upon the abundance and richness of other litter invertebrates. However, invertebrate group composition differed significantly between invaded and uninvaded sites, owing to the varied response of several influential groups (e.g. Collembola and Acarina). Floristics, habitat structure and measured environmental factors did not differ significantly between sites either invaded or uninvaded by Argentine ants, supporting the contention that differences in native ant abundance and species composition are related to invasion. Changes in the native ant community wrought by Argentine ant invasion have important implications for invertebrate communities in southern Australia and may affect key processes, including seed dispersal.     

Ant–hemipteran association decreases parasitism of Phenacoccus solenopsis by endoparasitoid Aenasius bambawalei

1. Mutualism between ants and honeydew-excreting hemipterans is ubiquitous in the ecosystem. It is widely accepted that ant tending facilitates the colony growth of hemipterans by protecting them from predators and parasitoids. However, few studies have explored how ant tending helps defend against natural enemies. 2. Ghost ant Tapinoma melanocephalum and the invasive mealybug Phenacoccus solenopsis have a close mutual relationship. Previous studies have shown that ghost ant tending can definitely reduce parasitism and visit frequency of Aenasius bambawalei, the dominant endoparasitoid of P. solenopsis. However, the ghost ant workers seldom attack the parasitoids. It is still unclear how the ghost ant adversely affects parasitoids. This study explored the mechanism underlying the impacts of ants on natural enemies of the mealybugs. 3. Honeydew produced by P. solenopsis was an attractant to A. bambawalei. Parasitoids exhibited less searching activity, shorter longevity and lower parasitism when supplied with less honeydew. Aenasius bambawalei showed significant avoidance of pygidial gland secretions and visual cues of ghost ants. Parasitism in plants treated with 6-methyl-5-hepten-2-one, actinidine, and gland extracts was significantly lower than that in plants treated only with solvents (paraffin oil or double-distilled water). 4. It is concluded that honeydew consumption by ghost ants could negatively influence the performance of parasitoids. The pygidial gland secretions and visual cues of ghost ants also significantly inhibit the parasitism. These results may contribute to a better understanding of the regulation mechanism in ant–hemipteran–enemy interactions.