Impact of environmental pollution on ant (Camponotus japonicus) development and labial gland disease.

Metallic pollutants can have harmful impacts on ant morphology and physiology. We studied the occurrence of labial gland disease in Camponotus japonicus from two polluted areas (traffic pollution and industrial pollution) and one non-polluted area. We further analyzed the metal levels (Al, As, Cd, Co, Cr, Cu, Fe, Hg, Mn, Ni, Pb and Zn) and morphological characteristics (head width, body mass, and other morphological traits) of both diseased and healthy workers. Our results showed that labial gland disease was only present in polluted areas, indicating that pollution stress makes ants more vulnerable to infections. Our research revealed that diseased ants in polluted areas accumulate higher levels of metals in their bodies and have lower dry weight and residual body mass compared to healthy ants in non-polluted environments, negatively impacting their development. We evaluated the importance of these elements and found that Cu has the greatest impact on the health risk of C. japonicus. Our study underscores the significant impact of environmental pollution on ant morphology and physiology and raises concerns about the broader ecological implications.

Leaderless consensus decision-making determines cooperative transport direction in weaver ants.

Animal groups need to achieve and maintain consensus to minimize conflict among individuals and prevent group fragmentation. An excellent example of a consensus challenge is cooperative transport, where multiple individuals cooperate to move a large item together. This behaviour, regularly displayed by ants and humans only, requires individuals to agree on which direction to move in. Unlike humans, ants cannot use verbal communication but most likely rely on private information and/or mechanical forces sensed through the carried item to coordinate their behaviour. Here, we investigated how groups of weaver ants achieve consensus during cooperative transport using a tethered-object protocol, where ants had to transport a prey item that was tethered in place with a thin string. This protocol allows the decoupling of the movement of informed ants from that of uninformed individuals. We showed that weaver ants pool together the opinions of all group members to increase their navigational accuracy. We confirmed this result using a symmetry-breaking task, in which we challenged ants with navigating an open-ended corridor. Weaver ants are the first reported ant species to use a 'wisdom-of-the-crowd' strategy for cooperative transport, demonstrating that consensus mechanisms may differ according to the ecology of each species.

Long-term strict ant-plant mutualism identity characterises growth rate and leaf shearing resistance of an Amazonian myrmecophyte.

Over 125 million years of ant-plant interactions have culminated in one of the most intriguing evolutionary outcomes in life history. The myrmecophyte Duroia hirsuta (Rubiaceae) is known for its mutualistic association with the ant Myrmelachista schumanni and several other species, mainly Azteca, in the north-western Amazon. While both ants provide indirect defences to plants, only M. schumanni nests in plant domatia and has the unique behaviour of clearing the surroundings of its host tree from heterospecific plants, potentially increasing resource availability to its host. Using a 12-year survey, we asked how the continuous presence of either only M. schumanni or only Azteca spp. benefits the growth and defence traits of host trees. We found that the continuous presence of M. schumanni improved relative growth rates and leaf shearing resistance of Duroia better than trees with Azteca. However, leaf herbivory, dry matter content, trichome density, and secondary metabolite production were the same in all trees. Survival depended directly on ant association (> 94% of trees died when ants were absent). This study extends our understanding of the long-term effects of strict ant-plant mutualism on host plant traits in the field and reinforces the use of D. hirsuta-M. schumanni as a model system suitable for eco-co-evolutionary research on plant-animal interactions.

Ant impacts on global patterns of bird elevational diversity.

Using data on bird species elevational distributions from the world's mountain ranges, bird diets, and the distribution of the ant genus Oecophylla, we report that global patterns in bird elevational diversity show signals of competition with ants. Oecophylla is an abundant and effective predator of invertebrates, preying on the same species that invertivorous birds feed on. In mountain ranges with Oecophylla present in the foothills, the maximum species richness of invertivorous birds (but not other trophic guilds) occurs, on average, at 960 m, ca. 450 m higher than in mountain ranges without Oecophylla, resulting in a mid-elevation peak in bird species richness. Where Oecophylla is absent, bird species richness for all guilds generally show monotonic declines with increasing elevation. We argue that Oecophylla reduces prey density for invertivorous birds and that low prey abundance reduces invertivorous bird density, which in turn is correlated with lower bird species richness. These findngs suggest that competition between distantly related taxa can set range limits, leading to emergent diversity patterns over large scales.

Male genitalia, hierarchical homology, and the anatomy of the bullet ant (Paraponera clavata; Hymenoptera, Formicidae).

The male genitalia of insects are among the most variable, complex, and informative character systems for evolutionary analysis and taxonomic purposes. Because of these general properties, many generations of systematists have struggled to develop a theory of homology and alignment of parts. This struggle continues to the present day, where fundamentally different models and nomenclatures for the male genitalia of Hymenoptera, for example, are applied. Here, we take a multimodal approach to digitalize and comprehensively document the genital skeletomuscular anatomy of the bullet ant (Paraponera clavata; Hymenoptera: Formicidae), including hand dissection, synchrotron radiation microcomputed tomography, microphotography, scanning electron microscopy, confocal laser scanning microscopy, and 3D-printing. Through this work, we generate several new concepts for the structure and form of the male genitalia of Hymenoptera, such as for the endophallic sclerite (=fibula ducti), which we were able to evaluate in detail for the first time for any species. Based on this phenomic anatomical study and comparison with other Holometabola and Hexapoda, we reconsider the homologies of insect genitalia more broadly, and propose a series of clarifications in support of the penis-gonopod theory of male genital identity. Specifically, we use the male genitalia of Paraponera and insects more broadly as an empirical case for hierarchical homology by applying and refining the 5-category classification of serial homologs from DiFrisco et al. (2023) (DLW23) to all of our formalized concepts. Through this, we find that: (1) geometry is a critical attribute to account for in ontology, especially as all individually identifiable attributes are positionally indexed hence can be recognized as homomorphic; (2) the definition of "structure" proposed by DLW23 is difficult to apply, and likely heterogeneous; and (3) formative elements, or spatially defined foldings or in- or evaginations of the epidermis and cuticle, are an important yet overlooked class of homomorphs. We propose a morphogenetic model for male and female insect genitalia, and a model analogous to gene-tree species-tree mappings for the hierarchical homology of male genitalia specifically. For all of the structures evaluated in the present study, we provide 3D-printable models - with and without musculature, and in various states of digital dissection - to facilitate the development of a tactile understanding. Our treatment of the male genitalia of P. clavata serves as a basic template for future phenomic studies of male insect genitalia, which will be substantially improved with the development of automation and collections-based data processing pipelines, that is, collectomics. The Hymenoptera Anatomy Ontology will be a critical resource to include in this effort, and in best practice concepts should be linked.

Species-specific behavioural responses to environmental variation as a potential species coexistence mechanism in ants.

A fundamental question of ecology is why species coexist in the same habitat. Coexistence can be enabled through niche differentiation, mediated by trait differentiation. Here, behaviour constitutes an often-overlooked set of traits. However, behaviours such as aggression and exploration drive intra- and interspecific competition, especially so in ants, where community structure is usually shaped by aggressive interactions. We studied behavioural variation in three ant species, which often co-occur in close proximity and occupy similar dominance ranks. We analysed how intra- and allospecific aggression, exploration and foraging activity vary under field conditions, namely with temperature and over time. Behaviours were assessed for 12 colonies per species, and four times each during several months. All behavioural traits consistently differed among colonies, but also varied over time and with temperature. These temperature-dependent and seasonal responses were highly species-specific. For example, foraging activity decreased at high temperatures in Formica rufibarbis, but not in Lasius niger; over time, it declined strongly in L. niger but much less in F. rufibarbis. Our results suggest that, owing to these species-specific responses, no species is always competitively superior. Thus, environmental and temporal variation effects a dynamic dominance hierarchy among the species, facilitating coexistence via the storage effect.

Disproportional fraction of inactive components leads to the variation in metabolic scaling.

In biological systems, solitary organisms or eusocial groups, the metabolic rate often scales allometrically with systems' size, when they are inactive, and the scaling becomes nearly isometric when the systems are active. Here I propose a hypothesis attempting to offer a departing point for a general joint understanding of the difference in the scaling powers between inactive and active states. When the system is inactive, there exist inactive components, which consume less energy than the active ones, and the larger the system is, the larger the fraction of the inactive components, which leads to sublinear scaling. When the system is active, most inactive components are activated, which leads to nearly isometric scaling. I hypothesize that the disproportional fraction of the inactive components is caused by the diffusants screening in the complex transportation network. I.e., when metabolites or information diffuses in the system, due to the physical limitation of the network structure and the diffusant's physical feature, not all the components can equally receive the diffusants so that these components are inactive. Using the mammalian pulmonary system, ant colonies, and other few systems as examples, I discuss how the screening leads to the allometric and isometric metabolic scaling powers in inactive and active states respectively. It is noteworthy that there are a few exceptions, in which the metabolic rate of the system has an isometric scaling relationship with size at rest. I show that these exceptions not only do not disapprove the hypothesis, but actually support it.

Ant colony optimization for the identification of dysregulated gene subnetworks from expression data.

BACKGROUND: High-throughput experimental technologies can provide deeper insights into pathway perturbations in biomedical studies. Accordingly, their usage is central to the identification of molecular targets and the subsequent development of suitable treatments for various diseases. Classical interpretations of generated data, such as differential gene expression and pathway analyses, disregard interconnections between studied genes when looking for gene-disease associations. Given that these interconnections are central to cellular processes, there has been a recent interest in incorporating them in such studies. The latter allows the detection of gene modules that underlie complex phenotypes in gene interaction networks. Existing methods either impose radius-based restrictions or freely grow modules at the expense of a statistical bias towards large modules. We propose a heuristic method, inspired by Ant Colony Optimization, to apply gene-level scoring and module identification with distance-based search constraints and penalties, rather than radius-based constraints. RESULTS: We test and compare our results to other approaches using three datasets of different neurodegenerative diseases, namely Alzheimer's, Parkinson's, and Huntington's, over three independent experiments. We report the outcomes of enrichment analyses and concordance of gene-level scores for each disease. Results indicate that the proposed approach generally shows superior stability in comparison to existing methods. It produces stable and meaningful enrichment results in all three datasets which have different case to control proportions and sample sizes. CONCLUSION: The presented network-based gene expression analysis approach successfully identifies dysregulated gene modules associated with a certain disease. Using a heuristic based on Ant Colony Optimization, we perform a distance-based search with no radius constraints. Experimental results support the effectiveness and stability of our method in prioritizing modules of high relevance. Our tool is publicly available at github.com/GhadiElHasbani/ACOxGS.git.

Tracing the genealogy origin of geographic populations based on genomic variation and deep learning.

Assigning a query individual animal or plant to its derived population is a prime task in diverse applications related to organismal genealogy. Such endeavors have conventionally relied on short DNA sequences under a phylogenetic framework. These methods naturally show constraints when the inferred population sources are ambiguously phylogenetically structured, a scenario demanding substantially more informative genetic signals. Recent advances in cost-effective production of whole-genome sequences and artificial intelligence have created an unprecedented opportunity to trace the population origin for essentially any given individual, as long as the genome reference data are comprehensive and standardized. Here, we developed a convolutional neural network method to identify population origins using genomic SNPs. Three empirical datasets (an Asian honeybee, a red fire ant, and a chicken datasets) and two simulated populations are used for the proof of concepts. The performance tests indicate that our method can accurately identify the genealogy origin of query individuals, with success rates ranging from  93 % to 100 %. We further showed that the accuracy of the model can be significantly increased by refining the informative sites through FST filtering. Our method is robust to configurations related to batch sizes and epochs, whereas model learning benefits from the setting of a proper preset learning rate. Moreover, we explained the importance score of key sites for algorithm interpretability and credibility, which has been largely ignored. We anticipate that by coupling genomics and deep learning, our method will see broad potential in conservation and management applications that involve natural resources, invasive pests and weeds, and illegal trades of wildlife products.

Draft genome assemblies of the ponerine ant Odontoponera transversa and the carpenter ant Camponotus friedae (Hymenoptera: Formicidae).

OBJECTIVES: Ants are ecologically dominant insects in most terrestrial ecosystems, with more than 14,000 extant species in about 340 genera recorded to date. However, genomic resources are still scarce for most species, especially for species endemic in East or Southeast Asia, limiting the study of phylogeny, speciation and adaptation of this evolutionarily successful animal lineage. Here, we assemble and annotate the genomes of Odontoponera transversa and Camponotus friedae, two ant species with a natural distribution in China, to facilitate future study of ant evolution. DATA DESCRIPTION: We obtained a total of 16 Gb and 51 Gb PacBio HiFi data for O. transversa and C. friedae, respectively, which were assembled into the draft genomes of 339 Mb for O. transversa and 233 Mb for C. friedae. Genome assessments by multiple metrics showed good completeness and high accuracy of the two assemblies. Gene annotations assisted by RNA-seq data yielded a comparable number of protein-coding genes in the two genomes (10,892 for O. transversa and 11,296 for C. friedae), while repeat annotations revealed a remarkable difference of repeat content between these two ant species (149.4 Mb for O. transversa versus 49.7 Mb for C. friedae). Besides, complete mitochondrial genomes for the two species were assembled and annotated.

Antibacterial activity of crude extracts of Camponotus compressus (Fabricius, 1787) (Hymenoptera: Formicidae).

The current study evaluates the antibacterial activity of Camponotus compressus (Hymenoptera: Formicidae) body crude extracts. The increasing antibiotic resistance of bacteria has prompted the world to turn its attention towards insects in the search for new sources of antibacterial compounds. The body crude extract obtained with different solvents were tested against both Gram positive (Staphylococcus aureus, Bacillus subtilis) and Gram negative bacteria (Pseudomonas aeruginosa, Escherichia coli, Klebsiella pneumoniae). Standard disc diffusion method was used to perform the activity. The extracts of C. compressus were investigated for their effectiveness against all resistant pathogenic bacteria. Staphylococcus aureus was found to be the most susceptible, exhibiting a high average growth inhibition, while Bacillus subtilis showed a lower average growth inhibition zone. Our findings regarding the inhibitory effect of C. compressus extracts show the presence of a broad-spectrum antibacterial compound. This will be helpful in the search for novel natural antibiotics against robust pathogenic bacterial strains.

Random walks with spatial and temporal resets can explain individual and colony-level searching patterns in ants.

Central place foragers, such as many ants, exploit the environment around their nest. The extent of their foraging range is a function of individual movement, but how the movement patterns of large numbers of foragers result in an emergent colony foraging range remains unclear. Here, we introduce a random walk model with stochastic resetting to depict the movements of searching ants. Stochastic resetting refers to spatially resetting at random times the position of agents to a given location, here the nest of searching ants. We investigate the effect of a range of resetting mechanisms and compare the macroscopic predictions of our model to laboratory and field data. We find that all returning mechanisms very robustly ensure that scouts exploring the surroundings of a nest will be exponentially distributed with distance from the nest. We also find that a decreasing probability for searching ants to return to their nest is compatible with empirical data, resulting in scouts going further away from the nest as the number of foraging trips increases. Our findings highlight the importance of resetting random walk models for depicting the movements of central place foragers and nurture novel questions regarding the searching behaviour of ants.

Coastal dunes as drivers of genetic differentiation in the honeypot ant Myrmecocystus baja (Formicidae: Formicinae).

Coastal dunes are unique habitats, threatened by human activities. In biogeographical terms, coastal dunes are habitat islands, being discrete and distinct patches of similar habitat among themselves, separated from each other by a different type of habitat. Furthermore, coastal dunes harbor endemic species, adapted to living solely in the habitats found on specific dune systems. For example, the honeypot ant Myrmecocystus baja is endemic and restricted to coastal dunes of Mexico's Baja California Pacific coast. This ecological and biogeographical scenario led to the questions whether their geographical isolation is reflected in their genetic diversity and structuring, and how their demographic history is related with the formation of the dune system habitats. To answer these questions, population genetic, isolation-with-migration, and phylogeographical analyses were carried out, based on mitochondrial and five nuclear intronic markers. Minimal gene flow was detected only between two of the dune systems sampled; otherwise, the M. baja populations were found to be isolated and genetically structured, and their divergence generally pre-dated the modern-day dune systems. It is therefore highly likely that these ants were already present in paleodunes and that each of the populations was established from founder populations as the dunes formed. These findings highlight the importance of coastal dunes for species such as the honeypot ant from Baja California, in promoting genetic differentiation.

A soluble model for synchronized rhythmic activity in ant colonies.

Synchronization is one of the most striking instances of collective behavior, occurring in many natural phenomena. For example, in some ant species, ants are inactive within the nest most of the time, but their bursts of activity are highly synchronized and involve the entire nest population. Here we revisit a simulation model that generates this synchronized rhythmic activity through autocatalytic behavior, i.e., active ants can activate inactive ants, followed by a period of rest. We derive a set of delay differential equations that provide an accurate description of the simulations for large ant colonies. Analysis of the fixed-point solutions, complemented by numerical integration of the equations, indicates the existence of stable limit-cycle solutions when the rest period is greater than a threshold and the event of spontaneous activation of inactive ants is very unlikely, so that most of the arousal of ants is done by active ants. Furthermore, we argue that the persistent oscillations observed in the simulations for colonies of finite size are due to resonant amplification of demographic noise.

Wolbachia-infected pharaoh ant colonies have higher egg production, metabolic rate and worker survival.

Wolbachia is a widespread maternally transmitted endosymbiotic bacteria with diverse phenotypic effects on its insect hosts, ranging from parasitic to mutualistic. Wolbachia commonly infects social insects, where it faces unique challenges associated with its host's caste-based reproductive division of labor and colony living. Here, we dissect the benefits and costs of Wolbachia infection on life-history traits of the invasive pharaoh ant, Monomorium pharaonis, which are relatively short lived and show natural variation in Wolbachia infection status between colonies. We quantified the effects of Wolbachia infection on the lifespan of queen and worker castes, the egg-laying rate of queens across queen lifespan, and the metabolic rates of whole colonies and colony members. Infected queens laid more eggs than uninfected queens but had similar metabolic rates and lifespans. Interestingly, infected workers outlived uninfected workers. At the colony level, infected colonies were more productive as a consequence of increased queen egg-laying rates and worker longevity, and infected colonies had higher metabolic rates during peak colony productivity. While some effects of infection, such as elevated colony-level metabolic rates, may be detrimental in more stressful natural conditions, we did not find any costs of infection under relatively benign laboratory conditions. Overall, our study emphasizes that Wolbachia infection can have beneficial effects on ant colony growth and worker survival in at least some environments.

Co-inheritance of recombined chromatids maintains heterozygosity in a parthenogenetic ant.

According to Mendel's second law, chromosomes segregate randomly in meiosis. Non-random segregation is primarily known for cases of selfish meiotic drive in females, in which particular alleles bias their own transmission into the oocyte. Here we report a rare example of unselfish meiotic drive for crossover inheritance in the clonal raider ant, Ooceraea biroi, in which both alleles are co-inherited at all loci across the entire genome. This species produces diploid offspring parthenogenetically via fusion of two haploid nuclei from the same meiosis. This process should cause rapid genotypic degeneration due to loss of heterozygosity, which results if crossover recombination is followed by random (Mendelian) segregation of chromosomes. However, by comparing whole genomes of mothers and daughters, we show that loss of heterozygosity is exceedingly rare, raising the possibility that crossovers are infrequent or absent in O. biroi meiosis. Using a combination of cytology and whole-genome sequencing, we show that crossover recombination is, in fact, common but that loss of heterozygosity is avoided because crossover products are faithfully co-inherited. This results from a programmed violation of Mendel's law of segregation, such that crossover products segregate together rather than randomly. This discovery highlights an extreme example of cellular 'memory' of crossovers, which could be a common yet cryptic feature of chromosomal segregation.

Testing the joint effects of arbuscular mycorrhizal fungi and ants on insect herbivory on potato plants.

MAIN CONCLUSION: Ants, but not mycorrhizae, significantly affected insect leaf-chewing herbivory on potato plants. However, there was no evidence of mutualistic interactive effects on herbivory. Plants associate with both aboveground and belowground mutualists, two prominent examples being ants and arbuscular mycorrhizal fungi (AMF), respectively. While both of these mutualisms have been extensively studied, joint manipulations testing their independent and interactive (non-additive) effects on plants are rare. To address this gap, we conducted a joint test of ant and AMF effects on herbivory by leaf-chewing insects attacking potato (Solanum tuberosum) plants, and further measured plant traits likely mediating mutualist effects on herbivory. In a field experiment, we factorially manipulated the presence of AMF (two levels: control and mycorrhization) and ants (two levels: exclusion and presence) and quantified the concentration of leaf phenolic compounds acting as direct defenses, as well as plant volatile organic compound (VOC) emissions potentially mediating direct (e.g., herbivore repellents) or indirect (e.g., ant attractants) defense. Moreover, we measured ant abundance and performed a dual-choice greenhouse experiment testing for effects of VOC blends (mimicking those emitted by control vs. AMF-inoculated plants) on ant attraction as a mechanism for indirect defense. Ant presence significantly reduced herbivory whereas mycorrhization had no detectable influence on herbivory and mutualist effects operated independently. Plant trait measurements indicated that mycorrhization had no effect on leaf phenolics but significantly increased VOC emissions. However, mycorrhization did not affect ant abundance and there was no evidence of AMF effects on herbivory operating via ant-mediated defense. Consistently, the dual-choice assay showed no effect of AMF-induced volatile blends on ant attraction. Together, these results suggest that herbivory on potato plants responds mainly to top-down (ant-mediated) rather than bottom-up (AMF-mediated) control, an asymmetry in effects which could have precluded mutualist non-additive effects on herbivory. Further research on this, as well as other plant systems, is needed to examine the ecological contexts under which mutualist interactive effects are more or less likely to emerge and their impacts on plant fitness and associated communities.

Paths from simple game-targeted pro-environmental behaviors to more diverse pro-environmental behaviors: Empirical evidence from the Ant Forest app.

Game interventions have drawn much attention for their playful features and "painless" promotion of simple game-targeted pro-environmental behaviors (GPEBs), which could spill over to more "costly" prosocial behaviors for pro-environmental causes (PSBPs). The literature places much emphasis on game-targeted mechanisms (e.g., enjoyment and competition) to explain the impacts of game interventions, but pays little attention to non-game-targeted mechanisms to explore their spillover effect. Using online survey data from 1246 Chinese residents, this paper first compares players' and nonplayers' levels of engagement with GPEBs and with two common PSBPs, finding a positive spillover effect from game-targeted simple behaviors to more diverse PSBPs. Based upon goal theory, self-perception theory, and learning theory, the paper then examines the underlying mediating mechanisms of the positive relationship between GPEBs and PSBPs for subsamples of players and nonplayers respectively. The results demonstrate that individual awareness of problem and self-efficacy could be enhanced through engaging in simple GPEBs, which could spill over to more challenging and "costly" PSBPs, including charitable giving and volunteering for pro-environmental causes. The study provides important insights on helping to encourage specific types of more challenging PSBPs not directly targeted by the game.

Ant social network structure is highly conserved across species.

The ecological success of social insects makes their colony organization fascinating to scientists studying collective systems. In recent years, the combination of automated behavioural tracking and social network analysis has deepened our understanding of many aspects of colony organization. However, because studies have typically worked with single species, we know little about interspecific variation in network structure. Here, we conduct a comparative network analysis across five ant species from five subfamilies, separated by more than 100 Myr of evolution. We find that social network structure is highly conserved across subfamilies. All species studied form modular networks, with two social communities, a similar distribution of individuals between the two communities, and equivalent mapping of task performance onto the communities. Against this backdrop of organizational similarity, queens of the different species occupied qualitatively distinct network positions. The deep conservation of the two community structure implies that the most fundamental behavioural division of labour in social insects is between workers that stay in the nest to rear brood, and those that leave the nest to forage. This division has parallels across the animal kingdom in systems of biparental care and probably represents the most readily evolvable form of behavioural division of labour.

Distribution of brood of the acorn ant Temnothorax crassispinus in artificial nests after forced migration.

Nest sites are important for social insects, as they provide refuge against enemies and ensure optimal conditions for the brood development. In large nests, the different chambers can be used for different reasons; for example, for food storage or as a brood chamber. Acorn ants from the genus Temnothorax dwell in small cavities in acorns and wood; however, even such small chambers can have a high degree of spatial heterogeneity. During this study, the distribution of brood items of the acorn ant Temnothorax crassispinus inside artificial nest cavities composed of three chambers in a linear system was analysed. 29 ant colonies were photographed 13 times during a period of approximately one month: during three consecutive days, and after forced migrations. I found that the distribution of the brood inside the nest cavity was similar during the consecutive days; however, after the forced migration, the distribution typically changed. Almost all the brood items were kept farther from the entrance. Keeping the brood farther from the entrance could be explained as a safer option.