Fight and rescue or give up and flee? Behavioural responses of different ant species tending the mutualist walnut aphid Panaphis juglandis to native and exotic lady beetles.

Mutualism between ants and honeydew-producing hemipterans is a highly successful evolutionary innovation that attains the status of ecological keystone across many terrestrial ecosystems, involving a multitude of actors through direct or cascading effects. In these relationships, ants often protect their hemipteran partners against their arthropod natural enemies, sometimes interfering with the biological control of pest species. However, the dynamics of these interactions are highly variable based on the specific identity of all the actors involved, and baseline data remain scarce. We performed a field experiment exposing colonies of the walnut aphid Panaphis juglandis attended by five European ant species (Camponotus piceus, Ca. vagus, Crematogaster scutellaris, Dolichoderus quadripunctatus, Lasius emarginatus) to a native and an exotic lady beetle (Adalia bipunctata and Harmonia axyridis), documenting the behavioural interactions between these insects and the performance of ants in the protection of the aphids. Our results reveal a significant behavioural diversity among the ant species involved, with D. quadripunctatus and L. emarginatus being the most aggressive and having the best performance as aphid defenders, and Ca. piceus being least effective and often fleeing away. Cr. scutellaris displayed a rare rescue behaviour attempting to pull away the aphids that the lady beetles grabbed. On the other hand, behavioural responses to A. bipunctata and H. axyridis were similar. Further investigations are needed to understand the eco-ethological implications of these differences, while a better understanding of ant behavioural diversity may help refine biological control strategies.

Do It by Yourself: Larval Locomotion in the Black Soldier Fly Hermetia illucens, with a Novel "Self-Harvesting" Method to Separate Prepupae.

The neotropical insect Hermetia illucens has become a cosmopolite species, and it is considered a highly promising insect in circular and sustainable economic processes. Being able to feed on a wide variety of organic substrates, it represents a source of lipids and proteins for many uses and produces recyclable waste. We investigated the characteristics and differences in the poorly-known locomotory behaviour of larvae of different instars, paying particular attention to the unique characteristics of the prepupal stage, key to farming and industrial processes. Moreover, we attempted to develop a "self-harvesting" system relying on the behavioural traits of prepupae to obtain their separation from younger larvae under rearing condition with minimum effort. Prepupae differ from younger larvae in their response to physical disturbance in the form of tonic immobility and significantly differ in their locomotory movements. Both prepupae and younger larvae reacted similarly to heat or light-induced stress, but low light and high moisture induced only prepupae to migrate away, which resulted in the development of a highly efficient separation methodology. The new data on the behaviour of H. illucens not only shed new light on some unexplored aspects of its biology, but also led to develop an inexpensive self-harvesting system that can be implemented in small-scale and industrial farming.

First Report on the Acrobat Ant Crematogaster scutellaris Storing Live Aphids in Its Oak-Gall Nests.

This study provides new data about the role of ants in mutualistic interactions with aphids mediated by galls. We focused our investigation on galls induced by the cynipid Andricus kollari by conducting a survey and a subsequent experiment in an Italian oak forest. The ants Crematogaster scutellaris, Colobopsis truncata and Temnothorax italicus frequently used the galls as nests: Crematogaster scutellaris occupied galls which were located higher on the oak trees, while C. truncata and T. italicus were located at lower positions. In addition, galls occupied by C. scutellaris showed varied internal architecture in relation to the colony composition. Importantly, field surveys revealed for the first time that C. scutellaris nest galls also contained live individuals of the non-galligenous aphid Panaphis juglandis. Field experiments suggested that the ants actively seek, collect and stock live aphids. No signs of predation and injuries were detected on the stored aphids, which were probably kept for safe overwintering, though we cannot exclude a possible occasional use as food. This report reveals a possible novel relationship which could have important consequences on the phenology and presence of aphids on the host plant.

Public Engagement Provides First Insights on Po Plain Ant Communities and Reveals the Ubiquity of the Cryptic Species Tetramorium immigrans (Hymenoptera, Formicidae).

Ants are considered a useful model for biodiversity monitoring and several of their characteristics make them promising for citizen science (CS) projects. Involving a wide range of public figures into collecting valuable data on the effect of human impact on ant biodiversity, the School of Ants (SoA) project represents one of the very few attempts to explore the potential of these insects in CS. Through the collaboration with the "BioBlitz Lombardia" project, we tested the SoA protocol on 12 Northern Italy parks, ranging from urban green to subalpine protected sites. As a result, we obtained some of the very first quantitative data characterizing the ants of this region, recording 30 species and highlighting some interesting ecological patterns. These data revealed the ubiquitous presence of the recently taxonomically defined cryptic species Tetramorium immigrans, which appears to be probably introduced in the region. We also discuss advantages and criticisms encountered applying the SoA protocol, originally intended for schools, to new categories of volunteers, from BioBlitz participants to park operators, suggesting best practices based on our experience.

Rolling away: a novel context-dependent escape behaviour discovered in ants.

For animals facing dangers, the best option to optimize costs and benefits of defence sometimes may be avoidance. Here we report the discovery of a peculiar strategy adopted by Myrmecina graminicola, a cryptic ant living in forest floor. Experiments showed that when disturbed these ants respond with immobility. However, upon perceiving disturbance but under specific inclinations of the substrate, they shift to an active escaping strategy: rolling away. This is a context-dependent behaviour adopted only in appropriate circumstances. During rolling, the ants assume a ball-like shape using antennae and hind legs to obtain an active movement along a stable trajectory. Finally, we assessed the adaptive value of this strategy measuring its effectiveness in defence against enemies. This is the first example of locomotion by rolling discovered in ants and one of the very few among animals, offering opportunities for multidisciplinary research on the adaptations and biomechanics underlying it.

New Tools for Conservation Biological Control: Testing Ant-Attracting Artificial Nectaries to Employ Ants as Plant Defenders.

Knowledge of the role of ants in many agroecosystems is relatively scarce, and in temperate regions the possibility to exploit ants as biocontrol agents for crop protection is still largely unexplored. Drawing inspiration from mutualistic ant-plant relationships mediated by extrafloral nectaries (EFNs), we tested the use of artificial nectaries (ANs) in order to increase ant activity on pear trees and to evaluate the effects on the arthropods, plant health and fruit production. While EFNs secrete a complex solution mainly composed of sugars and amino acids, ANs were filled with water and sucrose only. The results suggest that ANs can be used as manipulative instruments to increase ant activity over long periods of time. High ant activity was significantly linked to lower incidence of the pathogen fungus Venturia pyrina (pear scab) on pear leaves, and of the presence of Cydia pomonella (codling moth) caterpillars on pear fruit production. These results further encourage exploring underrated possibilities in the development of new tools for conservation biological control (CBC).

Gall-Colonizing Ants and Their Role as Plant Defenders: From 'Bad Job' to 'Useful Service'.

Galls are neoformed structures on host plant tissues caused by the attack of insects or other organisms. They support different communities of specialized parasitic insects (the gall inducers), and can also provide refuge to other insects, such as moths, beetles and ants, referred to as secondary occupants. This study focuses on galls induced by the oak gall wasp Andricus quercustozae and secondarily colonized by ants in a mixed oak forest. A field survey and two experiments were carried out to a) study ant (species-specific) preferences for different features of the galls, b) describe differences in gall architecture due to ant activity, c) analyse the effects of the presence of gall-dwelling ants on plant health. The results show that there are differences between ant species in gall colonization and in the alteration of gall opening and inner structure. We verified that gall-dwelling ants protect their host plants efficiently, offering them an indirect defence mechanism against enemies (predators and pathogens). The data suggest a new paradigm in ant-plant relationships mediated by the presence of galls on the plants whose ecological and evolutionary implications are discussed.

Dominance-diversity relationships in ant communities differ with invasion.

The relationship between levels of dominance and species richness is highly contentious, especially in ant communities. The dominance-impoverishment rule states that high levels of dominance only occur in species-poor communities, but there appear to be many cases of high levels of dominance in highly diverse communities. The extent to which dominant species limit local richness through competitive exclusion remains unclear, but such exclusion appears more apparent for non-native rather than native dominant species. Here we perform the first global analysis of the relationship between behavioral dominance and species richness. We used data from 1,293 local assemblages of ground-dwelling ants distributed across five continents to document the generality of the dominance-impoverishment rule, and to identify the biotic and abiotic conditions under which it does and does not apply. We found that the behavioral dominance-diversity relationship varies greatly, and depends on whether dominant species are native or non-native, whether dominance is considered as occurrence or relative abundance, and on variation in mean annual temperature. There were declines in diversity with increasing dominance in invaded communities, but diversity increased with increasing dominance in native communities. These patterns occur along the global temperature gradient. However, positive and negative relationships are strongest in the hottest sites. We also found that climate regulates the degree of behavioral dominance, but differently from how it shapes species richness. Our findings imply that, despite strong competitive interactions among ants, competitive exclusion is not a major driver of local richness in native ant communities. Although the dominance-impoverishment rule applies to invaded communities, we propose an alternative dominance-diversification rule for native communities.

Response of a Predatory ant to Volatiles Emitted by Aphid- and Caterpillar-Infested Cucumber and Potato Plants.

In response to herbivory by insects, various plants produce volatiles that attract enemies of the herbivores. Although ants are important components of natural and agro-ecosystems, the importance of herbivore-induced plant volatiles (HIPVs) as cues for ants for finding food sources have received little attention. We investigated responses of the ant Formica pratensis to volatiles emitted by uninfested and insect-infested cucumber (Cucumis sativus) and potato (Solanum tuberosum) plants. Cucumber plants were infested by the phloem-feeding aphid Aphis gossypii, the leaf chewer Mamestra brassicae or simultaneously by both insects. Potato plants were infested by either Aphis gossypii, by the leaf chewer Chrysodeixis chalcites or both. In olfactometer experiments, ants preferred volatile blends emitted by cucumber plants infested with M. brassicae caterpillars alone or combined with A. gossypii to volatiles of undamaged plants or plants damaged by A. gossypii only. No preference was recorded in choice tests between volatiles released by aphid-infested plants over undamaged plants. Volatiles emitted by potato plants infested by either C. chalcites or A. gossypii were preferred by ants over volatiles released by undamaged plants. Ants did not discriminate between potato plants infested with aphids and caterpillars over plants infested with aphids only. Plant headspace composition showed qualitative and/or quantitative differences between herbivore treatments. Multivariate analysis revealed clear separation between uninfested and infested plants and among herbivore treatments. The importance of HIPVs in indirect plant defence by ants is discussed in the context of the ecology of ant-plant interactions and possible roles of ants in pest management.

A global database of ant species abundances.

What forces structure ecological assemblages? A key limitation to general insights about assemblage structure is the availability of data that are collected at a small spatial grain (local assemblages) and a large spatial extent (global coverage). Here, we present published and unpublished data from 51 ,388 ant abundance and occurrence records of more than 2,693 species and 7,953 morphospecies from local assemblages collected at 4,212 locations around the world. Ants were selected because they are diverse and abundant globally, comprise a large fraction of animal biomass in most terrestrial communities, and are key contributors to a range of ecosystem functions. Data were collected between 1949 and 2014, and include, for each geo-referenced sampling site, both the identity of the ants collected and details of sampling design, habitat type, and degree of disturbance. The aim of compiling this data set was to provide comprehensive species abundance data in order to test relationships between assemblage structure and environmental and biogeographic factors. Data were collected using a variety of standardized methods, such as pitfall and Winkler traps, and will be valuable for studies investigating large-scale forces structuring local assemblages. Understanding such relationships is particularly critical under current rates of global change. We encourage authors holding additional data on systematically collected ant assemblages, especially those in dry and cold, and remote areas, to contact us and contribute their data to this growing data set.

Climate mediates the effects of disturbance on ant assemblage structure.

Many studies have focused on the impacts of climate change on biological assemblages, yet little is known about how climate interacts with other major anthropogenic influences on biodiversity, such as habitat disturbance. Using a unique global database of 1128 local ant assemblages, we examined whether climate mediates the effects of habitat disturbance on assemblage structure at a global scale. Species richness and evenness were associated positively with temperature, and negatively with disturbance. However, the interaction among temperature, precipitation and disturbance shaped species richness and evenness. The effect was manifested through a failure of species richness to increase substantially with temperature in transformed habitats at low precipitation. At low precipitation levels, evenness increased with temperature in undisturbed sites, peaked at medium temperatures in disturbed sites and remained low in transformed sites. In warmer climates with lower rainfall, the effects of increasing disturbance on species richness and evenness were akin to decreases in temperature of up to 9°C. Anthropogenic disturbance and ongoing climate change may interact in complicated ways to shape the structure of assemblages, with hot, arid environments likely to be at greatest risk.

Investigation by solid-phase microextraction and gas chromatography/mass spectrometry of trail pheromones in ants.

The Dufour's gland content of workers of two ant species of the genus Messor has been analyzed by solid-phase microextraction and gas chromatography with mass spectrometry (GC/MS). The structures of the compounds in the pheromonal mixtures have been determined. In both cases only one intact gland, inserted in a properly dimensioned capillary vial, is sufficient to produce a clean and fully interpretable GC/MS profile. It is worth noting that, for the first time in Messor ants, farnesol has been detected as a minor component of glandular secretion in Messor capitatus.