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Analyzing the information-rich content of RNA can help uncover genetic events associated with social insect castes or other social polymorphisms. Here, we exploit a series of cDNA libraries previously derived from whole-body tissue of different castes as well as from three behaviourally distinct populations of the Eastern subterranean termite Reticulitermes flavipes. We found that the number (~0.5 M) of single nucleotide variants (SNVs) was roughly equal between nymph, worker and soldier caste libraries, but dN/dS (ratio of nonsynonymous to synonymous substitutions) analysis suggested that some of these variants confer a caste-specific advantage. Specifically, the dN/dS ratio was high (~4.3) for genes expressed in the defensively specialized soldier caste, relative to genes expressed by other castes (~1.7–1.8) and regardless of the North American population (Toronto, Raleigh, Boston) from which the castes were sampled. The populations, meanwhile, did show a large difference in SNV count but not in the manner expected from known demographic and behavioural differences; the highly invasive unicolonial population from Toronto was not the least diverse and did not show any other unique substitution patterns, suggesting any past bottleneck associated with invasion or with current unicoloniality has become obscured at the RNA level. Our study raises two important hypotheses relevant to termite sociobiology. First, the positive selection (dN/dS > 1) inferred for soldier-biased genes is presumably indirect and of the type mediated through kin selection, and second, the behavioural changes that accompany some social insect urban invasions (i.e., ‘unicoloniality’) may be detached from the loss-of-diversity expected from invasion bottlenecks.

Biological invasions are considered a major driver of biodiversity loss, particularly on islands. Invasive alien ants can often have severe consequences on native biodiversity. Here, we review published and new information on alien ant species found on the Mediterranean island of Cyprus, a biodiversity hotspot. Our checklist of alien ants of Cyprus includes a total of 17 species, of which nine are reported from Cyprus for the first time (*): Camponotus cf. vitiosus Smith, Cardiocondyla mauritanica Forel, 1890, Cardiocondyla obscurior Wheeler, W.M., 1929*, Hypoponera punctatissima (Roger, 1859)*, Monomorium bicolor Emery, 1877, Nylanderia jaegerskioeldi (Mayr, 1904), Paratrechina longicornis (Latreille, 1802), Pheidole fadli Sharaf, 2007*, Pheidole indica Mayr, 1879, Solenopsis sp. (thief ant)*, Tetramorium bicarinatum (Nylander, 1846)*, Tetramorium caldarium (Roger, 1857)*, Tetramorium immigrans Santschi, 1927*, Tetramorium lanuginosum Mayr, 1870*, Trichomyrmex destructor (Jerdon, 1851), Trichomyrmex mayri (Forel, 1902)*, and Wasmannia auropunctata (Roger, 1863). We did not include three previously reported alien species for which we could not find supporting specimens [ Monomorium pharaonis (Linnaeus, 1758), Nylanderia vividula (Nylander, 1846), Solenopsis geminata (Fabricius, 1804)], one based on a previous misidentification [ Cardiocondyla nuda (Mayr, 1866)], and two species now considered native to Cyprus [ Hypoponera eduardi (Forel, 1894), Monomorium subopacum (F. Smith, 1858)]. Literature records, specimens from field surveys and museum collections, the geographic origin of species, occupied habitats in Cyprus, and notes on invasiveness (spread and impact) are presented for each species. An identification key to distinguish alien from native ant species in Cyprus is provided, including widespread alien ants not yet known from Cyprus in order to support early detection, monitoring, and management efforts.

During the last two centuries, thousands of insect species have been transported (largely inadvertently) and established outside of their native ranges worldwide, some with catastrophic ecological and economic impacts. Global variation in numbers of invading species depends on geographic variation in propagule pressure and heterogeneity of environmental resistance to invasions. Elton’s diversity-invasibility hypothesis, proposed over sixty years ago, has been widely explored for plants but little is known on how biodiversity affects insect invasions. Here we use species inventories from 44 land areas, ranging from small oceanic islands to entire continents in various world regions, to show that numbers of established insect species are primarily driven by diversity of plants, with both native and non-native plant species richness being the strongest predictor of insect invasions. We find that at large spatial scales, plant diversity directly explains variation in non-native insect species richness among world regions, while geographic factors such as land area, climate and insularity largely affect insect invasions indirectly via their effects on local plant richness.

Social insects harbor diverse assemblages of bacterial microbes, which may play a crucial role in the success or failure of biological invasions. The invasive fire ant Solenopsis invicta (Formicidae, Hymenoptera) is a model system for understanding the dynamics of invasive social insects and their biological control. However, little is known about microbes as biotic factors influencing the success or failure of ant invasions. This pilot study is the first attempt to characterize and compare microbial communities associated with the introduced S. invicta and the native Solenopsis geminata in the USA. Using 16S amplicon 454 pyrosequencing, bacterial communities of workers, brood, and soil from nest walls were compared between neighboring S. invicta and S. geminata colonies at Brackenridge Field Laboratory, Austin, Texas, with the aim of identifying potential pathogenic, commensal, or mutualistic microbial associates. Two samples of S. geminata workers showed high counts of Spiroplasma bacteria, a known pathogen or mutualist of other insects. A subsequent analysis using PCR and sequencing confirmed the presence of Spiroplasma in additional colonies of both Solenopsis species. Wolbachia was found in one alate sample of S. geminata, while one brood sample of S. invicta had a high count of Lactococcus. As expected, ant samples from both species showed much lower microbial diversity than the surrounding soil. Both ant species had similar overall bacterial diversities, although little overlap in specific microbes. To properly characterize a single bacterial community associated with a Solenopsis ant sample, rarefaction analyses indicate that it is necessary to obtain 5,000-10,000 sequences. Overall, 16S amplicon 454 pyrosequencing appears to be a cost-effective approach to screen whole microbial diversity associated with invasive ant species.

Biological invasions are a threat to global biodiversity and provide unique opportunities to study ecological processes. Population bottlenecks are a common feature of biological invasions and the severity of these bottlenecks is likely to be compounded as an invasive species spreads from initial invasion sites to additional locations. Despite extensive work on the genetic consequences of bottlenecks, we know little about how they influence microbial communities of the invaders themselves. Due to serial bottlenecks, invasive species may lose microbial symbionts including pathogenic taxa (the enemy release hypothesis) and/or may accumulate natural enemies with increasing time after invasion (the pathogen accumulation and invasive decline hypothesis). We tested these alternate hypotheses by surveying bacterial communities of Argentine ants (Linepithema humile). We found evidence for serial symbiont bottlenecks: the bacterial community richness declined over the invasion pathway from Argentina to New Zealand. The abundance of some genera, such as Lactobacillus, also significantly declined over the invasion pathway. Argentine ants from populations in the United States shared the most genera with ants from their native range in Argentina, while New Zealand shared the least (120 vs. 57, respectively). Nine genera were present in all sites around the globe possibly indicating a core group of obligate microbes. In accordance with the pathogen accumulation and invasive decline hypothesis, Argentine ants acquired genera unique to each specific invaded country. The United States had the most unique genera, though even within New Zealand these ants acquired symbionts. In addition to our biogeographic sampling, we administered antibiotics to Argentine ants to determine if changes in the micro-symbiont community could influence behavior and survival in interspecific interactions. Treatment with the antibiotics spectinomycin and kanamycin only slightly increased Argentine ant interspecific aggression, but this increase significantly decreased survival in interspecific interactions. The survival of the native ant species also decreased when the symbiotic microbial community within Argentine ants was modified by antibiotics. Our work offers support for both the enemy release hypothesis and that invasive species accumulate novel microbial taxa within their invaded range. These changes appear likely to influence invader behavior and survival.

Biological invasions are one of the top drivers of the ongoing biodiversity crisis. An underestimated consequence of invasions is the enormity of their economic impacts. Knowledge gaps regarding economic costs produced by invasive alien species (IAS) are pervasive, particularly for emerging economies such as India—the fastest growing economy worldwide. To investigate, highlight and bridge this gap, we synthesised data on the economic costs of IAS in India. Specifically, we examine how IAS costs are distributed spatially, environmentally, sectorally, taxonomically, temporally, and across introduction pathways; and discuss how Indian IAS costs vary with socioeconomic indicators. We found that IAS have cost the Indian economy between at least US$ 127.3 billion to 182.6 billion (Indian Rupees ₹ 8.3 trillion to 11.9 trillion) over 1960–2020, and these costs have increased with time. Despite these massive recorded costs, most were not assigned to specific regions, environments, sectors, cost types and causal IAS, and these knowledge gaps are more pronounced in India than in the rest of the world. When costs were specifically assigned, maximum costs were incurred in West, South and North India, by invasive alien insects in semi-aquatic ecosystems; they were incurred mainly by the public and social welfare sector, and were associated with damages and losses rather than management expenses. Our findings indicate that the reported economic costs grossly underestimate the actual costs, especially considering the expected costs given India’s population size, gross domestic product and high numbers of IAS without reported costs. This cost analysis improves our knowledge of the negative economic impacts of biological invasions in India and the burden they can represent for its development. We hope this study motivates policymakers to address socio-ecological issues in India and launch a national biological invasion research programme, especially since economic growth will be accompanied by greater impacts of global change.

Insect species associated with human goods continue to be accidentally introduced into new locations. A small proportion of these introduced species become invasive, causing a range of impacts in the receiving community. It is therefore important to evaluate the patterns of which species become invasive and which strategies are most successful in managing them. This review assesses the distribution, abundance, impact and management of the invasive Vespidae worldwide. We identified 34 vespid species known to be introduced around the world, but the seven most invasive species are all eusocial. Most introduced Vespidae only occur in one or two countries, but some areas have become geographic hotspots of invasion: Hawaii (15 species), North America (eight species), New Zealand (five species), Australia (four species) and South America (four species). Two invasive species, Vespula vulgaris and V. germanica have become particularly widespread and abundant with a range of impacts on biodiversity and ecosystem function. Other successful invasive species include several Polistes spp., which affect local biodiversity through direct predation or competition for food or space. Toxic baiting has been the most successful control strategy against invasive vespids to date, although this has mostly been small scale experimental management as it has proved difficult to develop commercial control products. Development of shelf-stable lures or baits combined with suitable toxins or pathogens could overcome some of the commercial impediments. Several attempts at biological control using parasitoids have not successfully reduced invasive wasp populations, although the biocontrol agent has only established in one case. The social structure of colonies and their high reproductive efficiency have facilitated invasion by these species, but it also means management at the population level will be difficult. This emphasises the need to prevent such invasions from occurring in the first place.

Although the ecological effects of invasions often become obvious soon after introduced species become established, more gradual effects may take years to manifest and can thus require long-term data for quantification. We analyzed an 8-year record of stable isotope data on Argentine ants (Linepithema humile) from southern California to infer how the trophic position of this widespread invasive species changes over time as native ant species are displaced. We couple this longitudinal analysis with a biregional comparison of stable isotope data (δ¹⁵N) on ants from Argentina (native range) and California (introduced range) to quantify (i) how the trophic position of L. humile differs between native and introduced populations, and (ii) how relative trophic position as estimated by δ¹⁵N values of Argentine ants compare with those of other ants at the same site. Both long-term and biregional comparisons indicate that the Argentine ant's relative trophic position is reduced at sites with a longer history of occupation. Over the course of 8 years, the relative trophic position of L. humile remained high at the leading edge of an invasion front but declined, on average, behind the front as native ants disappeared. Relative to native populations, where L. humile is among the most carnivorous of ants, Argentine ants from California occupied lower trophic positions. These results support the hypothesis that Argentine ants shift their diet after establishment as a result of resource depletion and increasing reliance on plant-based resources, especially honeydew-producing Hemiptera. Our results demonstrate the value of long-term and biregional data in uncovering ecological effects of invasions.

Predicting the outcome of competitive interactions is a fundamental goal in ecology. Ecological stoichiometry, which relates nutrient balance to ecological processes, provides a framework for identifying mechanistic links among macronutrient availability, nutritional physiology and competitive performance. Because carbohydrates serve as a principal metabolic fuel, carbohydrate scarcity may impinge upon behaviours affecting competitive dominance (e.g. aggression, activity) to a greater extent than deficiencies of protein or other nutrients used preferentially for growth. Here, we tested this prediction with a diet manipulation study involving laboratory colonies of Argentine ants (Linepithema humile), a widespread and aggressive invasive species. The availability of both sucrose and insect prey influenced brood production and worker survival after three months. However, colonies became less aggressive and less active only when deprived of sucrose (but not prey). Scarcity of sucrose (but not prey) was also associated with reduced fat mass in individual workers. These data provide the first experimental support that carbohydrate scarcity compromises aggression and activity in ants, and illustrate, in principle, how access to carbohydrate-rich resources (e.g. plant exudates, hemipteran honeydew) might influence behavioural investments that contribute to competitive performance. Such investments might be especially important for invasive ants, given their aggressiveness and tendency to interact with honeydew-producing Hemiptera.

Globalization and economic growth are widely recognized as important drivers of biological invasions. Consequently, there is an increasing need for governments to address the role of international trade in their strategies to prevent species introductions. However, many of the most problematic alien species are not recent arrivals but were introduced several decades ago. Hence, current patterns of alien-species richness may better reflect historical rather than contemporary human activities, a phenomenon which might be called “invasion debt.” Here, we show that across 10 taxonomic groups (vascular plants, bryophytes, fungi, birds, mammals, reptiles, amphibians, fish, terrestrial insects, and aquatic invertebrates) in 28 European countries, current numbers of alien species established in the wild are indeed more closely related to indicators of socioeconomic activity from the year 1900 than to those from 2000, although the majority of species introductions occurred during the second half of the 20th century. The strength of the historical signal varies among taxonomic groups, with those possessing good capabilities for dispersal (birds, insects) more strongly associated with recent socioeconomic drivers. Nevertheless, our results suggest a considerable historical legacy for the majority of the taxa analyzed. The consequences of the current high levels of socioeconomic activity on the extent of biological invasions will thus probably not be completely realized until several decades into the future.