Why do ants shift their foraging from extrafloral nectar to aphid honeydew?

When aphids parasitize plants with extrafloral nectaries (EFNs) and aphid colony size is small, ants frequently use EFNs but hardly tend aphids. However, as the aphid colony size increases, ants stop using EFNs and strengthen their associations with aphids. Although the shift in ant behavior is important for determining the dynamics of the ant–plant–aphid interaction, it is not known why this shift occurs. Here, we test two hypotheses to explain the mechanism responsible for this behavioral shift: (1) Extrafloral nectar secretion changes in response to aphid herbivory, or (2) plants do not change extrafloral nectar secretion, but the total reward to ants from aphids will exceed that from EFNs above a certain aphid colony size. To judge which mechanism is plausible, we investigated secretion patterns of extrafloral nectar produced by plants with and without aphids, compared the amount of sugar supplied by EFNs and aphids, and examined whether extrafloral nectar or honeydew was more attractive to ants. Our results show that there was no inducible extrafloral secretion in response to aphid herbivory, but the sugar concentration in extrafloral nectar was higher than in honeydew, and more ant workers were attracted to an artificial extrafloral nectar solution than to an artificial aphid honeydew solution. These results indicate that extrafloral nectar is a more attractive reward than aphid honeydew per unit volume. However, even an aphid colony containing only two individuals can supply a greater reward to ants than EFNs. This suggests that the ant behavioral shift may be explained by the second hypothesis.     

Do Lanchester's laws of combat describe competition in ants?

Lanchester's laws are mathematical models, originally designedto model military combat, that describe battle outcomes basedon the sizes of armies and the potencies of individual fightingunits. The "square law" describes a scenario in which small-sizedcompetitors may use a numerical advantage to overcome large-sizedcompetitors in simultaneous combat. The "linear law" describeshow a competitor with large-sized fighting units may use one-on-onecombat to favor victory when outnumbered. Lanchester's lawshave been suggested as an important regulator of interspecificcompetition in social insects, but without experimental support. In this study, experimental platforms were designed to invokethe conditions of both Lanchester's laws in a community ofants in lowland tropical wet forest in Costa Rica. I measuredbehavioral dominance at the food platforms in two separatemanners: an ordinal ranking of the number of workers present,and the monopoly of the food platforms. At the platform invokingsimultaneous combat, satisfying the square law, small-sizedants were more behaviorally dominant by numerical superiority.At the platform invoking one-on-one combat, satisfying thelinear law, larger ants were more behaviorally dominant by monopoly. These results suggest that Lanchester's laws explain,in part, the outcome of interspecific competition in ants.     

Does the availability of arboreal honeydew determine the prevalence of ecologically dominant ants in restored habitats?

Ants are extensive users of arboreal sugars, but little is known about how ecological dominance or habitat succession influences this interaction. We investigated how the availability and use of arboreal sugar resources by ants changes across a restoration chronosequence. We surveyed the use and availability of hemipteran honeydew and floral nectar on the two dominant plant genera, Eucalyptus and Acacia, in study sites in south eastern Australia. Sugars used by ants are likely to drive their role as ecosystem engineers, while sugars not used by ants remain available to other organisms. We also tested whether the use of sugars differed between ecologically dominant and non-dominant ants; taxa likely to perform different functions in ecosystems. No floral nectar was available on Acacia, but later successional eucalypts supported more floral resources and fewer mutualist hemiptera. Successional stage significantly affected how much sugar remained unexploited by ants, with similar trends for ant use of sugars. Non-dominant ants used mainly floral nectar, while hemipteran honeydew resources were used disproportionately by dominant ants, consistent with the prediction that this group monopolises persistent carbohydrate resources. This pattern was similar across successional stages, but the difference was least in habitats with the greatest availability of floral nectar, suggesting that high sugar availability may reduce the incentive to defend honeydew. Across habitat types, the proportion of dominant ants increased with the availability of hemipteran honeydew. This suggests that honeydew availability may regulate ecological dominance, thus affecting ant-driven ecosystem processes.     

Sugar feeding by the aphid parasitoid Binodoxys communis: how does honeydew compare with other sugar sources?

Parasitoids commonly forage in agricultural settings where the predominant sugar source is homopteran honeydew. The aphidiine braconid, Binodoxys communis, is an Asian parasitoid currently being released against the soybean aphid, Aphis glycines, in North American soybean fields. We conducted a number of laboratory experiments evaluating the quality of A. glycines honeydew as a sugar source for this parasitoid. Wasps readily fed on droplets of A. glycines honeydew, honey and 50% sucrose solution, but the length of feeding bouts on honey was significantly longer than on the other foods. Parasitoids lived significantly longer when fed honey or sucrose than honeydew, while starved wasps had the shortest lifespan. At 21+/-1 degrees C and 25+/-5% R.H., male B. communis that were fed honey lived for a maximum of 14 days, while females lived up to 20 days. Honeydew-fed wasps of both sexes lived approximately 3 days on average, which was 2-3 times longer than when they were only allowed access to water. Anthrone tests of whole insects showed that total sugar and glycogen levels of honey or sucrose-fed individuals were consistently higher than those fed honeydew or water. The glycogen levels of honeydew-fed wasps increased significantly after one day of feeding. HPLC analyses revealed that B. communis readily assimilates A. glycines honeydew oligosaccharides such as erlose, while others (e.g., raffinose) did not degenerate. Raffinose was present in much higher amounts in honeydew-fed wasps than in wasps fed other diets, so this sugar could be used as a 'signature' sugar for this species. Honeydew-fed wasps also had significantly lower fructose/(fructose+glucose) ratios than those from other diet treatments. Although A. glycines honeydew might be the main carbohydrate source within a soybean field, other sugar sources such as floral nectar appear to be more optimal foods for B. communis from a physiological standpoint. We discuss the results from the perspective of classical biological control of the soybean aphid in North America.     

Quarantining behaviour in ants: are Myrmica aphid milkers able to detect and get rid of fungus-contaminated aphids?

Fungal infections are highly dangerous for social insects including ants. Close trophobiotic interaction between ants and aphids promotes infection transmission, as aphids can be a disease vector. The ability of ants to detect fungus-infected aphids and get rid of them is important to the prosperity of both symbiotic partners. However, the diversity of quarantining behaviour among ants is still poorly studied. Here, the behaviour of honeydew foragers of two ant species – Myrmica rubra L. and Myrmica scabrinodis Nylander (Hymenoptera: Formicidae, Myrmicinae, Myrmicini) – was studied in laboratory towards Schizaphis graminum (Rondani) (Hemiptera: Aphididae, Aphidini) aphids contaminated with the generalist fungal pathogen Beauveria bassiana (Balsamo-Crivelli) Vuillemin (Hypocreales). This fungus attacks a wide range of hosts including aphids and ants. The removal of conidia-contaminated aphids from the host plant was found not to be typical of the ants studied. Aphid milkers of M. rubra and M. scabrinodis usually displayed non-aggressive behaviour (tolerance, antennation, honeydew collection, grooming) towards the experimental aphids regardless of whether they were covered with conidia or not. Neither ant species, nor the number of milkers had significant effects on their behaviour towards ‘infected’ aphids. However, some individuals were found to demonstrate quarantining behaviour in full. They quickly detected and removed contaminated aphids, placing them at some distance from the plant. Moreover, in addition to the simple carrying of ‘infected’ aphids down, the more effective technique of dropping them from the plant was noted as well. Ants of the genus Myrmica appear to have a tendency to perform a certain sequence of actions to remove conidia-contaminated aphids from the plant. It is likely that in larger colonies or under conditions of increased risk of infection with entomopathogenic fungi, some Myrmica ants are able to deploy and actively use the behavioural pattern of quarantining behaviour to increase their viability.     

The functional significance of E-β-Farnesene: Does it influence the populations of aphid natural enemies in the fields?

Aphids cause much damage to Chinese cabbage in northern China. Over reliance on pesticides have large environmental and human health costs that compel researchers to seek alternative management tactics for aphid control. The component of aphid alarm pheromone, E-β-Farnesene (EβF), extracted from Matricaria chamomilla L., which attracts natural enemies in the laboratory, may have significant implications for the design of cabbage aphid control strategies. The purpose of this paper is to understand the effects of EβF on natural enemies to cabbage aphid control in Chinese cabbage fields. Ladybeetles on Chinese cabbage leaves in EβF released plots and Aphidiidae in EβF released yellow traps were significantly higher than those of in controls. No significant differences were detected in the interactions of different treatments and the two years for all natural enemies. More important, lower aphid densities were found in EβF released plots. Our results suggested that the EβF extracted from M. chamomilla L. could attract natural enemies to reduce cabbage aphids in the Chinese cabbage fields.     

Do natural enemies really make a difference? Field scale impacts of parasitoid wasps and hoverfly larvae on cereal aphid populations

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Trophic ecology of parabiotic ants: Do the partners have similar food niches?

Organisms associated with another species may experience both costs and benefits from their partner. One of these costs is competition, which is the more likely if the two species are ecologically similar. Parabioses are associations between two ant species that share a nest and often attend the same food sources. Albeit parabioses are probably mutualistic, parabiotic partners may compete for food. We therefore investigated feeding niches and dietary overlap of two parabiotically associated ants in Borneo using cafeteria experiments and stable isotope analyses. The two species strongly differed in their food choices. While Crematogaster modiglianii mostly foraged at carbohydrate‐rich baits, Camponotus rufifemur preferred urea‐rich sources. Both species also consumed animal protein. The ¹⁵N concentration in Ca. rufifemur workers was consistently lower than in Cr. modiglianii. Camponotus rufifemur but not Cr. modiglianii possesses microbial endosymbionts, which can metabolize urea and synthesize essential amino acids. Its lower ¹⁵N signature may result from a relatively higher intake of plant‐based or otherwise ¹⁵N‐depleted nitrogen. Isotopic signatures of the two partners in the same parabiosis showed strongly parallel variation across nests. As we did not find evidence for spatial autocorrelation, this correlation suggests an overlap of food sources between the two ant species. Based on model simulations, we estimated a diet overlap of 22–66% for nitrogen sources and 45–74% for carbon sources. The overlap may arise from either joint exploitation of the same food sources or trophallactic exchange of food. This suggests an intense trophic interaction and potential for competition between the parabiotic partners.     

Aggression in imported fire ants: an explanation for shifts in their spatial distributions in Southern United States?

Abstract 1. The imported fire ants, Solenopsis invicta Buren (red), S. richteri Forel (black), and their hybrids (S. invicta × S. richteri) are sympatric congeners with overlapping but shifting spatial distributions in northern Alabama and Mississippi, United States. 2. The abilities of workers of the three ant forms (or species) to recognise non‐nestmate conspecifics and heterospecifics were compared in separate aggression bioassays using three group sizes of competitor numbers (battles): 1‐1, 5‐1, and 5‐5 resident–intruder battles. 3. Workers of all three forms showed significant aggression toward non‐nestmate conspecifics and heterospecifics. The results of the intra‐specific aggression bioassays showed that S. invicta had the lowest aggression threshold, whereas S. richteri was the least aggressive. Survival rates 1 day after intra‐specific encounters were higher for S. richteri than for S. invicta or hybrids, consistent with its lower aggressiveness. 4. In inter‐specific interactions, S. invicta workers showed the greatest aggression and were more aggressive towards S. richteri than to hybrids. Furthermore, survival 1 day after inter‐specific encounters was highest for S. invicta workers, whether they were the intruders or residents. The hybrid form was intermediate between both parental forms in aggression and post‐aggression survival. 5. The live 1‐1 and 5‐5 battles produced similar and consistent results, but differences were amplified in the 5‐5 battles, suggesting an effect of group size on aggression. These results may explain the observed shifts in the spatial distributions of the three forms and range restriction of S. richteri in southern United States.     

Is love in the air or at first sight? Mate finding cues used by sympatric male velvet ants (Hymenoptera: Mutillidae)

Mate finding has been examined in a wide range of hymenopteran families, with the greatest knowledge from social species and parasitoids used in pest management. Velvet ants (Hymenoptera: Mutillidae) provide a unique system to study mate finding because short‐lived flying males must find solitary, wingless females that have spatially and temporally dispersed emergence. Based on limited field observations, it is thought that male velvet ants may use visual, chemical, and vibratory cues for mate finding, though there have been no controlled experiments to test these hypotheses. Using choice experiments, this study aims to determine the role of visual and airborne chemical cues used in female detection by three sympatric species of Dasymutilla males. We find that visual cues are not used for initial female detection and that airborne chemicals are sufficient for female detection. We also discover that males readily approach females of different species, indicating airborne chemicals are not used for species recognition. Despite the lack of discrimination by males, we determined that the sympatric, morphologically similar species show no genetic introgression, and therefore some other isolating mechanism must be at play after males make initial contact with the females.     

Keep in touch: Does spatial overlap correlate with contact rate frequency?

Quantifying animal contact rate is crucial both in fundamental and applied studies to understand the evolution of sociality and predict the spread of infectious diseases. Researchers typically use home-range overlap among individuals as a proxy of contact rate, assuming a positive correlation. However, very few studies have assessed how the correlation between home-range overlap and contact rate may vary with ecological context. We used proximity loggers to quantify intraspecific contact rate among raccoons (Procyon lotor) and explored the correlation between contact rate and home-range overlap in different seasons. We monitored 15 female raccoons that formed 121 dyads during summer 2010 and winter 2011. We compared contact rate with the 5 most common overlap indices: home-range overlap proportion, home-range overlap probability, utilization distribution overlap index (UDOI), volume of intersection index, and Bhattacharyya's affinity index. Our results generally supported the contention of a positive and significant correlation between home-range overlap and intraspecific contact rate in raccoons. The strength of the relationship differed among seasons and indices, being weaker during winter than summer for home-range overlap proportion and home-range overlap probability. When contact rates were high, their frequency had stronger correlations with the UDOI and volume of intersection index indices than with the other indices. Our results suggest that the UDOI performs better than other indices, as we obtained a good contact rate–home-range overlap correlation with this index with animals both aggregated and randomly distributed in space. © 2012 The Wildlife Society.     

Do abandoned nests of leaf‐cutting ants enhance plant recruitment in the Atlantic Forest?

The role played by abandoned nests of leaf‐cutting ants (Atta spp.) as a small‐scale disturbance regime that affects plant recruitment, species coexistence and forest regeneration remains poorly investigated. Here we examine whether abandoned nests of Atta cephalotes serve as regeneration niches and operate as particular plant recruitment habitats, favouring forest regeneration after ant activities cease and leading to the establishment of taxonomically/ecologically distinct plant assemblages. Soil properties, canopy openness, light availability and regenerating plant assemblages were evaluated across 18 nests and adjacent control plots in a large remnant of Atlantic Forest in north‐east Brazil from December 2004 to December 2005. Surprisingly, nests and control plots exhibited very similar light environments irrespective of nest age, but nest soils exhibited substantial reductions in carbon content (1.45 ± 0.24 vs. 1.79 ± 0.13%) and organic matter (2.50 ± 0.41 vs. 3.08 ± 0.23%), and proved to be much more resistant to penetration (30.57 ± 6.08 vs. 39.48 ± 7.53 mm). Functional signature of regenerating plant assemblages exhibited little variation across both habitat types, as they were dominated by pioneer, small‐seeded and vertebrate‐dispersed species. However, abandoned nests exhibited less dense, impoverished and more homogeneous regenerating plant assemblages at local and landscape scale; they clearly lacked nest‐dependent plant species and represented floristic subsets of the flora inhabiting the undisturbed forest. This recruitment bottleneck was transient in the long term because nest‐related effects ameliorated in older nests. Our results suggest that, unlike treefall gaps, abandoned nests represent temporary (relatively long‐lasting) islands of unsuitable substrate that reduce plant recruitment, retard forest regeneration, and fail in providing a special regeneration niche able to promote species coexistence and plant diversity.     

Do environmental conditions experienced in early life affect recruitment age and performance at first breeding in common goldeneye females?

Environmental conditions experienced early in life may have long‐term impacts on life history traits and reproductive performance. We investigated whether ambient temperature experienced during the first two to four weeks of life and weather severity during the first two winters affected recruitment age and relative timing of breeding in the year of recruitment in female common goldeneyes Bucephala clangula. Our sample consisted of 141 female recruits hatched in a study population in central Finland between 1985 and 2013 and captured later as breeders. About 56% of the recruited females bred for the first time when two years old (range 2–6 yr). Individuals facing colder ambient temperatures during the first two to four weeks posthatch or more severe winter conditions during the first two winters did not recruit at an older age. Nor did maternal characteristics, relative hatch date or nest site availability affect recruitment age. For females that recruited at two years old, the date of first breeding was usually late relative to the population mean that year (mean difference 6.9 d, range –7 to 21 d). Our results suggest developmental buffering enables female goldeneye ducklings to mitigate the impacts of adverse environmental conditions experienced during the first weeks of life, at least in terms of first breeding.     

Neotyphodium endophytes in grasses: deterrents or promoters of herbivory by leaf-cutting ants?

Endophytic fungi, particularly in the genus Neotyphodium, are thought to interact mutualistically with host grasses primarily by deterring herbivores and pathogens via production of alkaloidal mycotoxins. Little is known, however, about how these endophytes interact with host plants and herbivores outside the realm of agronomic forage grasses, such as tall fescue, and their livestock grazers or invertebrate pest herbivores. We tested the effects of Neotyphodium inhabiting introduced tall fescue and native Arizona fescue on preference, survival, and performance of the leaf-cutting ant, Acromyrmex versicolor, an important generalist herbivore in the southwestern United States. In a choice experiment, we determined preferences of foraging queens and workers for infected and uninfected tall fescue and Arizona fescue. In a no-choice experiment, we determined queen survival, worker production, and size of fungal gardens for foundress queens reared on diets of infected and uninfected tall fescue and Arizona fescue. Foraging workers and queens did not significantly prefer either uninfected tall fescue or Arizona fescue relative to infected grasses, although ants tended to harvest more uninfected than infected tall fescue and more infected than uninfected Arizona fescue. Queen survivorship and length of survival was greater on uninfected tall fescue, uninfected Arizona fescue, and infected Arizona fescue than on infected tall fescue or the standard diet of palo verde and mesquite leaves. No queens survived beyond 6 weeks of the study when fed the infected tall fescue diet, in contrast to the effects of the other diets. Likewise, worker production was much lower and fungal garden size much smaller on infected tall fescue than in all other treatments, including the standard diet. In general, ant colonies survived and performed better on uninfected tall fescue and infected and uninfected Arizona fescue than standard diets of palo verde and mesquite leaves. The interaction of Neotyphodium with its host grasses is highly variable and these endophytes may increase, not alter, or even decrease resistance to herbivores. The direction of the interaction depends on host and fungal genotype, herbivore species, and environmental factors. The presence of endophytes in most, if not all, host plants suggests that endophytes may alter foraging patterns, performance, and survival of herbivores, such as leaf-cutting ants, but not always in ways that increase host plant fitness. Received: 27 October 1998 / Accepted: 19 October 1998     

Congruency of hydrocarbon patterns in heterospecific groups of ants: transfer and/or biosynthesis?

Summary In homospecific groups of ants, each species has its own hydrocarbon profile, on the epicuticle and in the postpharyngeal gland (PPG). When reared together in bispecific groups, workers of both species possess each other's hydrocarbons in both locations. The present study investigated two alternative mechanisms by which a mixed odour in artificial groups ofFormica selysi/Manica rubida can be created. Using [1-¹⁴C] sodium acetate as a precursor,de novo biosynthesis of hydrocarbons was demonstrated for both species whether reared in homospecific or mixed-species groups. The newly synthesized hydrocarbons occurred on the epicuticle, internally, and in particularly large amounts in the PPG. As expected from their PPG and epicuticular hydrocarbons composition, workersF. selysi synthesized alkanes and alkenes in comparable amounts irrespective of their rearing scheme. Likewise,M. rubida reared in bispecific groups synthesized mostly alkanes with only negligible amounts of alkenes, according to a ratio characteristic toM. rubida workers from homospecific groups and not toF. selysi workers. During dyadic encounters, a transfer of labeled hydrocarbons between nestmates (conspecific in homospecific groups and allospecific in mixed groups) was observed. These results suggest that the formation of the mixed hydrocarbon profile in artificial groups of ants is the result of a transfer of these chemicals between nestmates rather thande novo biosynthesis of the allospecific hydrocarbons. Behaviours like trophallaxis, grooming and body contact that occurred during the encounters mediated such a transfer.     

Interaction of the PhiHSIC virus with its host: lysogeny or pseudolysogeny?

The marine phage PhiHSIC has been previously reported to enter into a lysogenic relationship with its host, HSIC, identified as Listonella pelagia. This phage produces a variety of plaques on its host, including turbid and haloed plaques, from which lysogens were previously isolated. These lysogens were unstable during long-term storage at -80( degrees ) C and were lost. When HSIC was reinfected with phage PhiHSIC, pseudolysogen-like interactions between the phage and its host were observed. The cells (termed HSIC-2 or HSIC-2e) produced high viral titers (10(11) ml(-1)) in the absence of inoculating phage and yet reached culture densities of nearly 10(9) ml(-1). Prophages were not induced by mitomycin C or the polyaromatic hydrocarbon naphthalene in cells harboring such infections. However, such cells were homoimmune to superinfection. Colonies hybridized strongly with a gene probe from a 100-bp fragment of the PhiHSIC genome, while the host did not. Analysis of chromosomal DNA preparations suggested the presence of a chromosomally integrated prophage. Phage adsorption experiments suggested that HSIC-2 was adsorption impaired. Because of the chromosomal prophage integration and homoimmunity, we interpret these results to indicate that PhiHSIC establishes a lysogenic relationship with its host that involves an extremely high level of spontaneous induction. This could be caused by a weak repressor of phage production. Additionally, poor phage adsorption of HSIC-2 compared to the wild type probably helped maintain this pseudolysogen-like relationship. In many ways, pseudolysogenic phage-host interactions may provide a paradigm for phage-host interactions in the marine environment.     

Origin of an insect outbreak: escape in space or time from natural enemies?

Initiation of insect outbreaks is poorly understood, and may involve sporadic events that temporarily release insect populations from predation or parasitism. While studying a declining outbreak of the western tussock moth (Orgyia vetusta) on bush lupine (Lupinus arboreus), we witnessed the onset of a new tussock moth outbreak, separated by 1,000 m in space and 2 months in phenological timing from the original population. This new population underwent explosive growth for 2 years and then collapsed because of a massive die-off of lupines. We tested whether during its growth phase, this new outbreak benefited by escaping in either space or time from the natural enemies attacking the original population. In experimental populations on single bushes, we compared predation and parasitism at the sites of the new and the old outbreak. At the site of the old outbreak, we compared predation and parasitism early and late in the season. Parasitism was significantly lower and population growth significantly higher at the new outbreak site than the old one. Neither seasonal timing, predator exclusion, nor their interaction significantly affected survival at either site. Thus the new outbreak appeared to escape in space from parasitism. These results corroborate our previous experimental findings, which suggest that as predicted by theory, the interaction between the tussock moth and its parasitoids can produce large-scale spatial patterning in population densities.