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.

Evolutionary history of inquiline social parasitism in Plagiolepis ants.

Social parasitism, i.e. the parasitic dependence of a social species on another free-living social species, is one of the most intriguing phenomena in social insects. It has evolved to various levels, the most extreme form being inquiline social parasites which have lost the worker caste, and produce only male and female sexual offspring that are reared by the host worker force. The inquiline syndrome has been reported in 4 species within the ant genus Plagiolepis, in Europe. Whether inquiline social parasitism evolved once or multiple times within the genus remains however unknown. To address this question, we generated data for 5 inquiline social parasites - 3 species previously described and 2 unidentified species - and their free-living hosts from Europe, and we inferred their phylogenetic relationships. We tested Emery's rule, which predicts that inquiline social parasites and their hosts are close relatives. Our results show that inquiline parasitism evolved independently at least 5 times in the genus. Furthermore, we found that all inquilines were associated with one of the descendants of their most related free-living species, suggesting sympatric speciation is the main process leading to the emergence of the parasitic species, consistent with the stricter version of Emery's rule.

Alien ants spreading through Europe: Brachyponerachinensis and Nylanderiavividula in Italy.

The number of known alien ant species throughout Europe has been steadily increasing during the last few decades and Italy has been no exception, with four new taxa reported in the last five years. Here, we document new data on the Asian needle ant Brachyponerachinensis (Emery, 1895), an invasive alien species whose first establishment in Europe was detected in the southern Italian city of Naples in 2022 and which has now been found near Lake Como in northern Italy, representing the second European record, about 730 km distant from the first. Furthermore, we report for the first time the presence of Nylanderiavividula (Nylander, 1846) in the country, based on specimens collected both in Rome and near Lake Como. This is at least the second Nylanderia species established in the country after N.jaegerskioeldi, first reported in 2018. Unlike B.chinensis, N.vividula is not considered an ecological and health threat in the invaded range and is already known to occur in several other European countries. While only a few introduced ants in Europe are considered serious ecological, economic or health threats, the increasing circulation of several alien species and the poor ability to swiftly track their movements and detect their establishment can render management very difficult.

Neighbors sharing pathogens: the intricate relationship between Apis mellifera and ants (Hymenoptera: Formicidae) nesting in hives.

Ants are ubiquitous and eusocial insects that exhibit frequent physical contact among colony members, thereby increasing their susceptibility to diseases. Some species are often found in beehives and in their surroundings, where they exploit the food resources of honey bees. This intricate relationship may facilitate the interspecific transmission of honey bee pathogens to ants, although ants themselves may contribute to spillback phenomena. The objective of this study was to assess the presence and abundance of honey bee pathogens in ants sampled from Italian apiaries. A total of 37 colonies within 24 apiaries across 7 regions were monitored. In total, 6 pathogens were detected in adult ants and 3 in the brood. In particular, the study revealed a high prevalence of honey bee pathogens in ants, with DWV, BQCV, and CBPV being the most commonly encountered. The brood also tested positive for the same viruses. Notably, all analyzed viruses were found to be replicative in both adult ants and ant broods. Furthermore, co-infections were prevalent, suggesting complex pathogen interactions within ant populations. Statistical analysis indicated significant differences in pathogen prevalence and abundance among ant species and sample types. The findings highlight active infection in both the ants and the brood, suggesting a potential role of ants as reservoir hosts and vectors of honey bee pathogens emphasizing the need for further research to understand the implications of interspecific pathogen transmission on ant and bee health.

Response to Genovesi et al.: Ant biosurveillance should come before invasion.

Menchetti et al. respond to the letter of Genovesi et al. and contribute new records of the red imported fire ant in Sicily.

Nomenclatural changes on some Mediterranean Aphaenogaster Mayr, 1853 taxa (Hymenoptera, Formicidae).

The ant genus Aphaenogaster Mayr, 1853 is especially diverse in the West-Palearctic region, where its taxonomy is still problematic for several groups. Among other issues, there is a redundancy of valid taxa compared to the number of species whose validity is currently sustained by any evidence. Names that do not represent taxa with clearly defined identities complicate specimen identification and add confusion to biodiversity databases and checklists. Based on a review of the available literature information on several taxa, and the study of both type and non-type material, the following taxonomic changes are proposed: i) A. mauritanica Dalla Torre, 1893 should be considered a senior synonym of A. gibbosa homonyma Emery, 1921 syn. nov. and of A. foreli Cagniant, 1996 syn. nov. (the Maghrebian populations of the pallida group hitherto considered to represent A. foreli should be attributed to A. dulcineae Emery, 1924); ii) A. pallida (Nylander, 1849) should be considered a senior synonym of A. leveillei Emery, 1881 syn. nov. and of A. leveillei laurenti Santschi, 1939 syn. nov.; iii) A. sangiorgii (Emery, 1901) should be considered a senior synonym of A. finzii Müller, 1921 syn. nov. and A. radchenkoi Kiran & Tezcan, 2008 syn. nov.; iv) A. sardoa Mayr, 1853 should be considered a senior synonym of A. sardoa ujhelyii Szabó, 1910 syn. nov. and A. sardoa anoemica Santschi, 1910 syn. nov.; v) A. subterraneosplendida André, 1883 should be considered a senior synonym of A. sicula Emery, 1908 syn. nov.; vi) A. subterraneoides Emery, 1881 should be considered a senior synonym of A. subterraneoides armeniaca Arnol'di, 1968 syn. nov. These changes will reduce the gap between nomenclature and actual understanding of species diversity, while synonymized names may be resurrected if supporting evidence is discovered.

Predatory Abilities of Two Mediterranean Ants on the Eggs and Larvae of the Codling Moth Cydia pomonella.

The predatory ability of ants (Hymenoptera, Formicidae) against insect pests can offer an important service to agricultural activities and may sometimes be directly exploited in biological control strategies. The codling moth Cydia pomonella (Lepidoptera, Tortricidae) is a major agricultural pest of fruit orchards, whose biological control is complicated by the fact that the larvae spend most of their life protected within the fruits they damage. In a recent experiment in Europe, pear trees in which ant activity was artificially increased by the addition of sugary liquid dispensers (artificial nectaries) suffered less damage caused by the larvae to their fruits. While some ants were already known to prey upon the mature larvae or pupae of C. pomonella in the soil, prevention of fruit damage would require predation upon eggs or newly hatched larvae, which have not yet excavated into the fruits. We verified whether two different Mediterranean ants frequently observed in fruit orchards, Crematogaster scutellaris and Tapinoma magnum, were able to prey upon C. pomonella eggs and larvae in laboratory conditions. Our experiments demonstrated that both species similarly attacked and killed young C. pomonella larvae. On the other hand, the eggs mostly attracted the attention of T. magnum but were never damaged. Further field assessments are required to understand whether ants may also interfere with oviposition by adults or whether larger ant species, although generally rarer in orchards, may also prey upon eggs.

The invasive ant Solenopsis invicta is established in Europe.

The red imported fire ant (Solenopsis invicta) is classified as one of the worst invasive alien species1 and as the fifth costliest worldwide2, impacting ecosystems, agriculture and human health3. We report the establishment of S. invicta in Europe for the first time, documenting a mature population in Sicily. We use genetic analyses to assess its putative origin, as well as wind tracking and species distribution modelling to predict its potential range on the continent. We show that half of the urban areas in Europe are already suitable and that climate warming expected under current trends will favor the expansion of this invasive ant.

Unrecognized for centuries: distribution and sexual caste descriptions of the West European Aphaenogaster species of the subterranea group (Hymenoptera, Formicidae).

There are only two Aphaenogaster species from the subterranea group in the western Mediterranean: A.ichnusa Santschi, 1925, from south-western Europe, and A.subterranea (Latreille, 1798), also occurring in central and eastern Europe. Historically, the two species have been widely misunderstood: A.ichnusa was long considered a Sardinian endemic subspecies of A.subterranea, while its continental populations were misidentified as A.subterranea s. str. Recently, A.ichnusa was elevated to species rank and its worker caste was redescribed with that of A.subterranea, allowing for their correct identification. Yet their distribution was documented in detail only for France and Sardinia. Furthermore, no morphological characters were described to distinguish the males and queens of the two species. By investigating private and museum collections, 276 new records of A.ichnusa are provided here and 154 of A.subterranea from the western Mediterranean. Additionally, qualitative and quantitative morphological characters were combined to identify their males and queens. We present the new southernmost, easternmost, and westernmost distribution limits for A.ichnusa. Based on our results, this species is widely distributed in Italy and Catalonia (Spain), also occurring on several Mediterranean islands, avoiding areas with continental climate and high altitudes. Sicily is the only island to host the less thermophilous A.subterranea, which otherwise extends westward to Galicia (Spain). Sympatric occurrence is not rare along the contact zone. Additional natural history observations are reported regarding foraging habits, associated myrmecophiles, habitat preferences, and colony structure in the two species.

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.

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).

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.