Biology subjects

Czaczkes, T. J.

Publications and source records attributed to Czaczkes, T. J..

At least 19 recordsLinked to original sources

bioRxiv2026

Below-ground ants follow pheromones more quickly under dark conditions, but pheromones do not affect decision accuracy nor aggression

Communication allows organisms to quickly convey information vital for survival or fitness. Chemical communication and speed-accuracy trade-offs are ubiquitous in animal decision making. Most studies have used species which forage mainly above-ground species, tested in an epigean setting, but it remains unclear whether below-ground species behave similarly. Here, we use the below-ground ant Tetramorium alpestre to assess the efficacy of above- vs. below-ground mazes, the accuracy of decisions when using natural vs. artificial pheromones, the presence of a speed-accuracy trade-off, and the pheromones effect on aggression. Ants decided more quickly under below-ground than above-ground conditions, indicating they may be distracted by above-ground stimuli. Ants followed both natural and artificial trails but in direct competition preferred artificial trails, likely due to a higher pheromone concentration. Surprisingly, no speed-accuracy trade off was observed during path decision-making. Lastly, population origin but not pheromones affected if and how aggression occurred in presence of trail and home-range marking pheromones. We argue that the design of behavioural tests should match the lifestyle of the focal organism. We further speculate that speed-accuracy trade-offs likely are highly species- and context-specific and other factors besides chemicals seem important to trigger aggression, at least in this species.

animal behavior and cognition↗

bioRxiv2025

Urban Lasius niger ants more readily accept low concentration sucrose solution than rural ants

Urbanisation causes broad changes in biotic and abiotic factors, which are often detrimental to animals. While extensive attention has been focussed on food resources for pollinators, comparatively little research has examined resources for aphidophagous insects. Many ants, such as Lasius niger, are highly aphidophagous, relying on honeydew secretions for the majority of their carbohydrate intake. Ants will also often reject sugar solution which is of lower concentration than that they were expecting. Here, we ask whether Lasius niger ants from urban environments show differential acceptance to various sucrose solution molarities. We offered outgoing ants on active foraging trails drops of sucrose solutions over a range of molarities (0.125, 0.25, 0.5, 1, and 2M), and quantified acceptance. Acceptance was scored using a simple behavioural measure of whether the ant remained in contact with the sucrose drop for over 3 seconds (full acceptance), broke away from the drop within 3 seconds but remained in the area and eventually drank to satiation (partial acceptance), or did not drink to satiation and walked away (rejection). Urban ants showed a significantly higher acceptance of all sucrose molarities than rural ants, with this difference especially pronounced at lower molarities. This may indicate that they are under nutritional stress for carbohydrates. Ant responses to sucrose solutions may allow a straightforward measure of the availability of carbohydrate resources in the environment.

ecology↗

bioRxiv2025

No decoy effect in bees: rewardless flowers do not increase bumblebees' preference for neighbouring flowers

Nectarless flowers are common among flowering plants, which often retain colour-changed, rewardless flowers instead of shedding them. Yet, how these flowers influence pollinators foraging choices within an inflorescence remains unclear. We hypothesised that rewardless flowers in an inflorescence may act as "decoys", causing the rewarding flowers in the inflorescence to be perceived as more valuable by contrast. Using artificial inflorescences, we presented individual bumblebees (Bombus terrestris) with a binary choice between two equally rewarding inflorescences, one of which included additional unrewarded, differently coloured flowers. We found that the presence of rewardless flowers did not increase bees preference for neighbouring flowers, nor did it affect their overall choice between inflorescences. However, bees quickly learned to avoid the unrewarded flowers, drastically reducing visits and probing within a few foraging bouts. We review research on decoy effects in bees, and find very little support for their presence. Our findings contribute to the growing body of evidence that rewardless flowers do not induce decoy effects in bees, and highlight the need for further research into the ecological role of nectarless flowers within floral patches. It may be time to abandon the search for classical decoy effects in pollinators.

animal behavior and cognition↗

bioRxiv2025

Bumblebees do not prefer consistent floral scents over variable ones

To attract pollinators, flowering plants evolve diverse sensory traits into compelling signals. Floral scent, in particular, plays a key role in drawing bees from a distance and shaping their foraging choices. Scent composition varies widely, including across flowers of the same plant species. Yet, it is unclear whether scent variability influences bee flower choices, and thus whether plants would be under selection to minimise variation in their scent composition. Since bees typically avoid variability in rewards, we hypothesised they would favour flowers with more consistent scents. To test this, we trained individual bumblebees (Bombus terrestris) on two equally rewarding flower arrays: one with a consistent scent blend across flowers, and the other with variable scent blends between flowers. Contrary to expectations, bees showed no preference for scent consistency. They readily foraged from both arrays across bouts and did not favour either flower type in the binary choice test. To the best of our knowledge, this is the first study to examine how bees respond to scent variability. A better understanding of scent profile preferences in pollinators could offer new insights into their co-evolution with plants and the development of floral traits. More broadly, further research is needed on how pollinators respond to variability and unpredictability in neutral cues like scent or colour, a largely overlooked aspect of foraging decision-making.

animal behavior and cognition↗

bioRxiv2024

Interindividual variability in flower pickiness by foraging bumblebees

Pollinators navigate complex and heterogeneous "flower markets", where floral resources vary in quality, availability, and spatial distribution. Bumblebees, as generalist foragers, visit numerous flowers during their foraging bouts, yet the factors influencing their flower choices and the individual differences in foraging behaviour remain poorly understood. Here, we tested how bees adjust their foraging in response to different reward structures. Bombus terrestris workers completed three foraging bouts in two artificial flower environments: one simulating a favourable environment with patches alternating high- and low-quality flowers (40% vs. 20% w/w sucrose solution), and the other a challenging environment with high-quality flowers alongside unrewarded ones (40% w/w sucrose solution vs. plain water). We hypothesised that bees would improve their foraging efficiency in both environments, but more rapidly in the more extreme one, where the greater reward difference creates stronger pressure to learn quickly. In both conditions, bees increased their sucrose intake per unit time over bouts. We also observed consistent differences in flower selectivity among individuals: in the favourable environment, bees that first visited high-quality flowers focused on them and avoided low-quality ones (became "picky"), while bees that first visited low-quality flowers kept visiting both types. Despite these differences, bees across environments and pickiness levels all reached similar sucrose intake rates by the third foraging bout, either by becoming more selective, collecting more sucrose solution, or reducing time spent foraging. These findings highlight the adaptability of bee foraging and suggest that early flower experiences may contribute to lasting individual differences in foraging behaviour. Significance statementBumblebees are highly efficient pollinators of wild plants and commercial crops, yet their foraging behaviour varies notably among individuals. Understanding why bees differ in their foraging decisions is crucial, as it affects how they collect resources. We show that individual Bombus terrestris workers rapidly adapt to flower patches with different reward qualities. Remarkably, a bees first flower visits have lasting effects: individuals that started with high-quality flowers consistently favoured them and avoided low-quality ones, while those that started with low-quality flowers continued to visit both types over time. Despite these differences, bees reached similar nectar intake rates after just three foraging trips, regardless of how selective they were. Our findings show that bees can quickly adjust their foraging behaviour, and that early experiences play a key role in how they exploit resources across different environments.

animal behavior and cognition↗

bioRxiv2024

Testing the peak-end rule in bumblebees: lack of preference for a higher-reward sequence when the final reward is disappointing

The peak-end rule describes the tendency to evaluate experiences by their most intense and final moments, rather than considering the entire experience as a whole. While this cognitive bias is well-established in humans, studies on nonhuman animals are very limited. Bumblebees make foraging decisions largely based on past experiences, but whether peak-end effects influence their subsequent flower choices is still unknown. Here, we trained individual Bombus terrestris workers on two artificial flower types, alternating blue and yellow, over 12 consecutive foraging bouts. One flower type offered a sequence of three high-quality rewards (25 L drops of 50% w/w sucrose solution: "consistent" sequence), while the other provided the same sequence but ended with an additional, lower-quality reward (25 L drop of 20% w/w sucrose solution: "poor end" sequence). We then tested the bees flower type preference in a final binary choice. Bees showed a strong preference for blue flowers, both in their initial and overall visits. Across all visits during a 1-minute period, they also favoured flowers associated with the "consistent" sequence, though this preference was significant only when these flowers were yellow. Interestingly, despite offering more sucrose per foraging bout, bees did not favour the "poor end" sequence flower. This study is, to our knowledge, the first to investigate peak-end effects in an insect. How bees evaluate sequential rewards when foraging remains largely unexplored, yet could provide valuable insights into nectar distribution and plant-pollinator co-evolution.

animal behavior and cognition↗

bioRxiv2024

Palatability of Insecticides and Protein in Sugar Solutions to Argentine Ants

Invasive ant species like Linepithema humile cause significant ecological and economic harm, making effective control strategies essential. Insecticide baits are currently the most effective approach for controlling ants. Therefore, quantifying how palatable or unpalatable baits, bait additives, or toxicants are, is critical for developing effective control methods. It has recently been demonstrated that in the comparative evaluation of foods, animals that are aware of both a test food and a comparator food exhibit greatly increased sensitivity when detecting the unpalatability of liquid baits. Here, we deploy a newly developed comparative evaluation methodology to examine the palatability to L. humile workers of three toxicants used in invasive ant control: Fipronil, spinosad, and imidacloprid, as well as egg white protein. Ants showed no significant preference between pure sucrose and sucrose-toxicant solutions, indicating that they either cannot detect the toxicants or that they do not find them distasteful. Survival tests confirmed that the toxicant concentrations used were lethal, with a survival rate of 50% or below after 72 hours. However, ants found egg protein additive unpalatable, significantly preferring pure sucrose to a sucrose-egg protein mix. These findings confirm that three major toxicants are suitable for use in baits, and that reported abandonment or avoidance of toxic baits is not due to the unpalatability of these toxicants. However, the addition of egg protein to sucrose baits, even at ratios which optimise colony growth, is likely counterproductive. Future research should investigate the relative preference of invasive ants for various bait matrixes over naturally available food, ensuring more effective pest management strategies.

animal behavior and cognition↗

bioRxiv2024

Invasive ants fed spinosad collectively recruit to known food faster yet individually abandon food earlier

Current management strategies applied to invasive ants rely on slow-acting insecticides which aim to delay the ants ability to detect the poison until its effects are noticeable. Despite this, most control efforts are unsuccessful, likely due to bait abandonment and insufficient sustained consumption. Conditioned taste aversion, a learned avoidance of a particular taste, is a crucial survival mechanism which prevents animals from repeatedly ingesting toxic substances. However, whether ants are capable of this delayed association between food taste and subsequent illness remains largely unexplored. Here, we exposed colonies of the highly invasive Argentine ant, Linepithema humile, to a sublethal dose of the slow-acting insecticide spinosad. We combined measurements of individual-level feeding patterns with quantification of collective preferences and foraging dynamics to investigate the potential effects of the toxicant on behaviour. Collectively, ants preferred an odour associated with a previously experienced food, even if this contained spinosad, over a novel one. However, at the individual-level, previous exposure to spinosad resulted in reduced food consumption, as a consequence of earlier food abandonment. Moreover, while control-treated colonies recruited slower to a food source which tasted like a previously experienced one, spinosad-exposed colonies recruited equally fast to both novel and familiar foods. Although it appears that ants are unable to develop a conditioned taste aversion to sublethal doses of spinosad, ingestion of even small amounts of the toxicant strongly influences foraging behaviour. Understanding the subtle effects of slow-acting pesticides on ant cognition and behaviour can ultimately inspire the development of more efficient control methodologies.

animal behavior and cognition↗

bioRxiv2024

Three-dimensional body reconstruction enables quantification of liquid consumption in small invertebrates

Quantifying feeding patterns provides valuable insights into animal behaviour. However, small invertebrates often consume incredibly small amounts of food. This renders traditional methods, such as weighing individuals before and after food acquisition, either inaccurate or prohibitively expensive. Here, we present a non-invasive method to quantify food consumption of small invertebrates whose body expands during feeding. Using the markerless pose estimation software DeepLabCut, we three-dimensionally track the body of Argentine ants, Linepithema humile. Using these extracted markers, we developed an algorithm which computationally reconstructs the ants body, directly measuring volumetric change over time. Moreover, we provide measures of accuracy and quantify the ants feeding response to a range of sucrose concentrations, as well as a gradient of caffeine-laced sucrose solutions. Small invertebrates are often prolific invasive species and disease vectors, causing significant ecological and economical damage. Understanding their feeding behaviour could be an important step towards effective control strategies.

animal behavior and cognition↗

bioRxiv2024

A high-throughput and sensitive method for food preference assays in walking insects

Insects pose significant challenges in both pest management and ecological conservation. Often, the most effective strategy is employing toxicant-laced baits, which must also be designed to specifically attract and be preferred by the targeted species for optimal species- specific effectiveness. However, traditional methods for measuring bait preference are either non-comparative, meaning that most animals only ever taste one bait, or suffer from methodological or conceptual limitations. Here we demonstrate the value of direct comparison food preference assays using the invasive and pest ant Linepithema humile (Mayr, 1868) as a model. We compare the food preference sensitivity of non-comparative (one visit to a food source) and sequential comparative (visiting one type of food then another) assays at detecting low levels of aversive quinine in sucrose solution. We then introduce and test a novel dual-choice feeder method for simultaneous comparative evaluation of bait preferences, testing its effectiveness in discerning between foods with varying quinine or sucrose levels. While the non-sequential assay could not detect aversion to 1.25mM quinine in 1M sucrose, the sequential comparative approach detected aversion to quinine levels as low as 0.94mM. The novel dual feeder method approach could detect aversion to quinine levels as low as 0.31mM, and also preference for 1M sucrose over 0.75M sucrose. The dual-feeder method combines the sensitivity of comparative evaluation with high throughput, ease of use, and avoidance of interpretational issues. This innovative approach offers a promising tool for rapid and effective testing of bait solutions, contributing to the development of targeted control strategies. The method could also be easily extended to other ant species.

animal behavior and cognition↗

bioRxiv2024

High variability in the attractiveness of municipally-planted decorative plants to pollinators

Insect populations are declining globally. A major driver of this decline is land use change, including urbanisation. However, urban environments can also offer a wide range of floral resources to pollinators, through ornamental plantings, but these can vary widely in their attractiveness to insects. Often, the largest single planter of ornamental plants in an urban area is the municipality. Here we evaluated the decorative plantings carried out by the city municipality of Regensburg, Germany, by systematically surveying insect visitations on different plant types in late summer, when forage is often limited for pollinators. We found a 130-fold increase from the least to the most attractive plants, and high variation in which insect groups were attracted to which plants. While honey bees, which are not conservation concern, were the most common insect visitors, some decorative plants attracted a very large proportion of wild bees, flies, and wasps. Our results demonstrate that there is great scope for increasing the supply of urban forage to pollinators in general, and specific groups in particular, without requiring new decorative plant types to be sourced or planted. We argue that providing local evidence-based guidance to municipalities offers a quick and potentially cost-neutral method for supporting urban insect populations.

ecology↗

bioRxiv2023

Acute exposure to caffeine improves navigation in an invasive ant

Invasive alien species are a major and growing problem, devastating ecosystems and costing billions of euros in damage and control efforts. Argentine ants, Linepithema humile, are particularly concerning, with control efforts often falling short likely due to a lack of sufficient bait consumption. Using neuroactives to manipulate ant navigation and learning could increase recruitment and consumption, ultimately leading to more efficient control strategies. Caffeine is naturally occurring, cheap, and has been found to cause motivational and cognitive improvements in bees. Here, we subject L. humile to a wide range of caffeine concentrations and a complex but ecologically relevant task: an open landscape foraging experiment. Without caffeine, we find no effect of consecutive foraging visits on the time the ants take to reach a reward, suggesting a failure to learn the rewards location. However, low (25ppm) to intermediate (250ppm) concentrations of caffeine lead to a decrease of up to 38% in the time taken to find the reward during each consecutive visit, implying that caffeine boosts learning. Interestingly, such improvements are lost at high (2000ppm) doses. In contrast, caffeine appears to have no impact on the ants homing behaviour, as the time required to reach the nest was similar across treatments. The effect of caffeine is thus not only dose-dependent, but also differentially targets neurologically distinct navigational mechanisms. Adding moderate levels of caffeine to baits could be a simple way to improve ants ability to learn its location, potentially leading to increased recruitment to, and consumption of, the toxicant.

animal behavior and cognition↗

bioRxiv2023

Can nestmate corpses cause bait neglect in invasive ants?

Invasive ants, such as Linepithema humile (the Argentine ant), pose a global threat, necessitating a better understanding of their behaviour in order to improve management strategies. Traditional eradication methods, including baiting, have had limited success, but the causes of control failure are not always clear. Here we propose that ants may learn to avoid toxic baits in part due to their association with ant corpses. Ants were tested on a Y-maze after exposure to scented corpses or dummies. 69% (n = 64) of ants avoided branches bearing the scent of scented corpses. At a collective level, colonies neglected food sources associated with scented ant corpses in favour of a food source with a novel odour, with only 42% (n = 273) of foragers feeding from the corpse-scent associated food source. However, if corpses were produced by feeding ants scented toxicant, focal ants encountering these corpses did not avoid the corpse-associated scent on a Y maze (53%, n = 65). Moreover, in a dual-feeder test, ants did not avoid feeding at food sources scented with an odour associated with conspecific corpses. The study demonstrates that conspecific corpses can act as a negative stimulus for Linepithema humile, leading to avoidance of odours associated with corpses, which can lead to potential avoidance of toxic baits. Why the more realistic Y-maze trial with corpses of ants that had ingested the toxicant elicited no avoidance is unclear: it may be due to weaker odour cues from ingested food, or a counterbalancing of the negative corpse stimulus by the positive presence of food remains on the corpse. Nonetheless, this study demonstrates that conspecific corpses act as a negative stimulus for ants, and this should be kept in mind when planning control efforts. A simple solution to this issue would be adding odours to baits, and cycling baits between treatments. This would disrupt the association between bait odour and corpses.

animal behavior and cognition↗

bioRxiv2022

Optimistic ants: Positive cognitive judgement bias but no emotional contagion in the ant Lasius niger

Understanding the emotional states of animals is key for informing their ethical treatment, but very little attention has been directed towards the emotional lives of invertebrates. As emotions influence information processing, one way to assess emotional states is to look for an individuals cognitive bias, i.e., their tendency to make optimistic or pessimistic judgements. Here we developed a free-running judgment bias task for the ant Lasius niger, and applied the judgement bias to assess ants reactions towards positive and negative stimuli. After an initial learning phase in which individuals were trained to associate two odour stimuli with positive or negative reinforcement, their reaction towards ambiguous stimuli, i.e., a mixture between both odours, was assessed. We also explored our study species capacity to socially transmit emotional states ( emotional contagion) by investigating whether social information could elicit emotional responses. We find L. niger to be optimistic, showing a baseline positive judgement bias, with 65-68% of ants preferring an ambiguous 1:1 mix of positive and negative cues over no cues. Providing an unexpected food reward prior to the judgement bias task increases positive judgement bias (c. 75% positive). There was a non- significant tendency towards a negative judgement bias after experiencing a mild electric shock (c. 75% negative). Neither positive nor negative social information (trail and alarm pheromones, respectively) affected the ants judgement biases, thus providing no indication for emotional contagion. The development of a powerful, simple, and ecologically relevant cognitive judgement task, deployable in the lab and in the field, opens the door to systematic comparative studies of the evolutionary and ecological causes of judgement bias.

animal behavior and cognition↗

bioRxiv2022

Invasive ant learning is not affected by seven potential neuroactive chemicals

Nectar-feeding insects are often the victims of psychoactive manipulation, with plants lacing their nectar with secondary metabolites such as alkaloids and non-protein amino acids which often boost learning, foraging, or recruitment. However, the effect of neuroactive chemicals has seldomly been explored in ants. Argentine ants (Linepithema humile) are one of the most damaging invasive alien species worldwide. Enhancing or disrupting cognitive abilities, such as learning, has the potential to improve management efforts, for example by increasing preference for a bait, or improving ants ability to learn its characteristics or location. Here, we test the effects of seven potential neuroactive chemicals - two alkaloids: caffeine and nicotine; two biogenic amines: dopamine and octopamine, and three non-protein amino acids: {beta}-alanine, GABA and taurine - on the cognitive abilities of invasive L. humile using bifurcation mazes. Our results confirm that these ants are strong associative learners, requiring as little as one experience to develop an association. However, we show no short-term effect of any of the chemicals tested on spatial learning, and in addition no effect of caffeine on short-term olfactory learning. This lack of effect is surprising, given the extensive reports of the tested chemicals affecting learning and foraging in bees. This mismatch could be due to the heavy bias towards bees in the literature, a positive result publication bias, or differences in methodology. Graphical Abstract O_FIG O_LINKSMALLFIG WIDTH=200 HEIGHT=129 SRC="FIGDIR/small/514620v1_ufig1.gif" ALT="Figure 1"> View larger version (17K): org.highwire.dtl.DTLVardef@10a971org.highwire.dtl.DTLVardef@1e4701dorg.highwire.dtl.DTLVardef@7cb741org.highwire.dtl.DTLVardef@1e33eea_HPS_FORMAT_FIGEXP M_FIG C_FIG

animal behavior and cognition↗

bioRxiv2022

Bundling and segregation affects "liking", but not "wanting", in an insect

Behavioural economists have identified many psychological manipulations which affect perceived value, although value in humans is not a unitary experience, with "liking" and "wanting" being neurologically separate processes. A prominent example of this is bundling, in which several small gains (or costs) are experienced as more valuable (or costly) than if the same total amount is presented together. While extensively demonstrated in humans, to our knowledge this effect has never been investigated in an animal, let alone an invertebrate. We trained individual Lasius niger workers to two of three conditions in which either costs (travel distance), gains (sucrose reward), or both were either bundled or segregated: A) both costs and gains bundled, B) both segregated, and C) only gains segregated. We recorded pheromone deposition on the ants return trips to the nest as measure of "liking". After training, we offer the ants a binary choice between odours associated with the treatments, as a measure of "wanting". While bundling treatment did not affect choice, i.e. "wanting", it strongly influenced pheromone deposition, i.e. "liking". Ants deposited c. 80% more pheromone when rewards were segregated but costs bundled as compared with both costs and rewards being bundled. This pattern is further complicated by the pairwise experience each animal made, and which of the treatments it experiences first during training. The current study is the first to demonstrate a bundling effect in an animal, and the first to report a dichotomy between "liking" and "wanting" in an insect. We propose that the deviation between "wanting" and "liking" in this case is due to the unique nature of distance perception in ants, which is recorded linearly, while almost all other sensory perception in animals is logarithmic.

ecology↗

bioRxiv2022

Bees differentiate sucrose solution from water at a distance

Bees are perhaps the most important model for studying complex cognition in invertebrates, showing a variety of impressive abilities. Many experiments employ a training procedure in which the animal associates a characteristic of a "flower" (neutral stimulus) to sucrose solution (positive stimulus) over multiple foraging bouts. We hypothesized that sucrose solution might appear different from water to the bee sensorium, rendering it superfluous for the animal to learn the intended experimental cues. To test this hypothesis, we presented bumblebees simultaneously with 1.6M sucrose and water on artificial flowers. We presented the solutions in three conditions, to identify the most likely recognition mechanism: in drop form, inside cotton-plugged centrifuge tubes, and soaked into raised cigarette filters. The bumblebees chose to contact the sucrose solution significantly more often in all three conditions, confirming their ability to discriminate sucrose from water at a distance, with spectral differences being the most likely mechanism for this differentiation. These results have large implications for the design of training procedures, as the presence of an alternative cue may mask the learning of challenging tasks. This may force us to re-examine much of the literature and unpublished negative results on bee cognition: weakening some claims, but strengthening others.

animal behavior and cognition↗

bioRxiv2022

A systematic examination of learning in the invasive ant Linepithema humile reveals very rapid development of short and long-term memories

The Argentine ant (Linepithema humile) is one of the most damaging and widespread invasive ant species worldwide. However, control attempts often fail due to bait rejection or insufficient bait uptake. Increasing preference for, and consumption of, bait is thus an important requirement for successful control. Learning and within-nest information transfer might be a potential tool for achieving this goal. We conducted a systematic investigation of olfactory learning and route learning in Argentine ants. The ants showed very strong and rapid route learning, choosing the correct arm in a Y-maze 65% of time after just one visit, and 84% correct after two. Odour learning was even more rapid, reaching up to 85% correct choices after just one exposure to flavoured food. Learning is long-lasting, with 73% correct choices after 48h. Food flavour information is transferred efficiently between nestmates in the nest, driving preference: naive ants housed with ants fed on flavoured food show a strong preference (77%) for that odour after 24h. Overall, Linepithema humile are outstanding learners. This, coupled with efficient intranidal information transfer and strong pheromonal recruitment, may help explain their ability to discover and then dominate resources. However, these strengths could potentially be used against them, by exploiting learning and information transfer to increase toxic bait uptake. Steering ant preference by leveraging learning might be an underappreciated tool in invasive alien species control.

animal behavior and cognition↗