Organisation of foraging in ants

In social insects, foraging is often cooperative, and so requires considerable organisation. In most ants, organisation is a bottom-up process where decisions taken by individuals result in emergent colony level patterns. Individuals base their decisions on their internal state, their past experience, and their environment. By depositing trail pheromones, for example, ants can alter the environment, and thus affect the behaviour of their nestmates. The development of emergent patterns depends on both how individuals affect the environment, and how they react to changes in the environment. Chapters 4 – 9 investigate the role of trail pheromones and route memory in the ant Lasius niger. Route memories can form rapidly and be followed accurately, and when route memories and trail pheromones contradict each other, ants overwhelmingly follow route memories (chapter 4). Route memories and trail pheromones can also interact synergistically, allowing ants to forage faster without sacrificing accuracy (chapter 5). Home range markings also interact with other information sources to affect ant behaviour (chapter 6). Trail pheromones assist experienced ants when facing complex, difficult-to-learn routes (chapter 7). When facing complicated routes, ants deposit more pheromone to assist in navigation and learning (chapter 7). Deposition of trail pheromones is suppressed by ants leaving a marked path (chapter 5), strong pheromone trails (chapter 7) and trail crowding (chapter 8). Colony level ‘decisions’ can be driven by factors other than trail pheromones, such as overcrowding at a food source (chapter 9). Chapter 10 reviews the many roles of trail pheromones in ants. Chapters 11 – 14 focus on the organisation of cooperative food retrieval. Pheidole oxyops workers arrange themselves non-randomly around items to increase transport speeds (chapter 11). Groups of ants will rotate food items to reduce drag (chapter 12). Chapters 13 and 14 encompass the ecology of cooperative transport, and how it has shaped trail pheromone recruitment in P. oxyops and Paratrechina longicornis. Lastly, chapter 15 provide a comprehensive review of cooperative transport in ants and elsewhere

Effect of trail bifurcation asymmetry and pheromone presence or absence on trail choice by Lasius niger ants

During foraging, ant workers are known to make use of multiple information sources, such as private information (personal memory) and social information (trail pheromones). Environmental effects on foraging, and how these interact with other information sources, have, however, been little studied. One environmental effect is trail bifurcation asymmetry. Ants forage on branching trail networks and must often decide which branch to take at a junction (bifurcation). This is an important decision, as finding food sources relies on making the correct choices at bifurcations. Bifurcation angle may provide important information when making this choice. We used a Y-maze with a pivoting 90° bifurcation to study trail choice of Lasius niger foragers at varying branch asymmetries (0°, [both branches 45° from straight ahead], 30° [branches at 30° and 60° from straight ahead], 45°, 60° and 90° [one branch straight ahead, the other at 90°]). The experiment was carried out either with equal amounts of trail pheromone on both branches of the bifurcation or with pheromone present on only one branch. Our results show that with equal pheromone, trail asymmetry has a significant effect on trail choice. Ants preferentially follow the branch deviating least from straight, and this effect increases as asymmetry increases (47% at 0°, 54% at 30°, 57% at 45°, 66% at 60° and 73% at 90°). However, when pheromone is only present on one branch, the graded effect of asymmetry disappears. Overall, however, there is an effect of asymmetry as the preference of ants for the pheromone-marked branch over the unmarked branch is reduced from 65%, when it is the less deviating branch, to 53%, when it is the more deviating branch. These results demonstrate that trail asymmetry influences ant decision-making at bifurcations and that this information interacts with trail pheromone presence in a non-hierarchical manner

No specialist pheromone-ignoring ants in Lasius niger

In insect societies, the balance between exploitation of known resources and exploration of new ones is important to ensure sufficient resources. Mass recruiting ants, such as Lasius niger, use pheromone trails to recruit nestmates to a newly discovered food source. Pheromone following, however, shows characteristic non-following (lapse) rates among different species, with similar to 20% of L. niger foragers ignoring pheromone. These characteristic lapse rates might simply be 'noise', or they might indicate a subset of specialised explorative foragers, a scouting caste, that consistently ignores pheromone in order to explore. Here we show pheromone ignoring is not a repeatable behaviour in L. niger foragers - ants who did not follow a trail were no more likely to ignore it again an hour later than ants which did follow it. Our findings suggest that there is no subset of specialised pheromone-ignoring L. niger foragers. This may be due to their moderate colony size and strong reliance on individual memories: species with larger colony sizes or a weaker reliance on private information (i.e. memory) may have specialist non-followers. Our work raises the question: what is a scout ant? We encourage future research to investigate the presence of a scouting caste in other ant species using our straightforward methodology, as a social information-ignoring caste may be rarer than expected

No evidence for tactile communication of direction in foraging Lasius ants

The idea that ants communicate when meeting on a trail is beguiling, but evidence for this is scarce. Physical communication in ants has been demonstrated to play a role as a modulator of behaviours such as alarm and recruitment. Honeybees can communicate the location of a resource using an advanced motor display – the waggle dance. However, no equivalent of the waggle dance has been described for any ant species, and it is widely believed that ants cannot communicate the location of resources using motor displays. One group of researchers report several demonstrations of such communication in Formica ants; however, these results have been largely ignored. More recently some evidence arose that Lasius niger foragers returning from a food source can communicate to outgoing foragers the direction that should be taken at the next bifurcation by means of physical contact on the trail. Here, we make a concerted effort to replicate these results. Although initial results seemed to indicate physical communication, once stringent controls to eliminate pheromone cues were put in place, no evidence for physical communication of food location could be found. This null result was replicated independently by a different research group on a closely related species, L. neglectus. We conclude that neither L. niger nor L. neglectus foragers communicate resource location using physical contact. Our results increase the burden of proof required for other claims of physical communication of direction in ants, but do not completely rule out this possibility

Copy when uncertain: lower light levels increase trail pheromone deposition and reliance on pheromone trails in ants

Animals may gather information from multiple sources, and these information sources may conflict. Theory predicts that, all else being equal, reliance on a particular information source will depend on its information content relative to other sources. Information conflicts are a good area in which to test such predictions. Social insects, such as ants, make extensive use of both private information (e.g. visual route memories) and social information (e.g. pheromone trails) when attempting to locate a food source. Importantly, eusocial insects collaborate on food retrieval, so both information use and information provision may be expected to vary with the information content of alternative information sources. Many ants, such as Lasius niger, are active both day and night. Variation in light levels represents an ecologically important change in the information content of visually-acquired route information. Here, we examine information use and information provision under high light levels (3200 lux, equivalent to a bright but overcast day), moderate light levels simulating dusk (10 lux) and darkness (0.007 lux, equivalent to a moonless night). Ants learn poorly, or not at all, in darkness. As light levels decrease, ants show decreasing reliance on private visual information, and a stronger reliance on social information, consistent with a ‘copy when uncertain’ strategy. In moderate light levels and darkness, pheromone deposition increases, presumably to compensate for the low information content of visual information. Varying light levels for cathemeral animals provides a powerful and ecologically meaningful method for examining information use and provision under varying levels of information content

Irrational risk aversion in an ant

Animals must often decide between exploiting safe options or risky options with a chance for large gains. Both proximate theories based on perceptual mechanisms, and evolutionary ones based on fitness benefits, have been proposed to explain decisions under risk. Eusocial insects represent a special case of risk sensitivity, as they must often make collective decisions based on resource evaluations from many individuals. Previously, colonies of the ant Lasius niger were found to be risk-neutral, but the risk preference of individual foragers was unknown. Here, we tested individual L. niger in a risk sensitivity paradigm. Ants were trained to associate one scent with 0.55 M sucrose solution and another with an equal chance of either 0.1 or 1.0 M sucrose. Preference was tested in a Y-maze. Ants were extremely risk-averse, with 91% choosing the safe option. Based on the psychophysical Weber–Fechner law, we predicted that ants evaluate resources depending on their logarithmic difference. To test this hypothesis, we designed 4 more experiments by varying the relative differences between the alternatives, making the risky option less, equally or more valuable than the safe one. Our results support the logarithmic origin of risk aversion in ants, and demonstrate that the behaviour of individual foragers can be a very poor predictor of colony-level behaviour

Acute exposure to caffeine improves foraging in an invasive ant

Argentine ants, Linepithema humile, are a particularly concerning invasive species. Control efforts often fall short likely due to a lack of sustained bait consumption. Using neuroactives, such as caffeine, to improve ant learning and navigation could increase recruitment and consumption of toxic baits. Here, we exposed L. humile to a range of caffeine concentrations and a complex ecologically relevant task: an open landscape foraging experiment. Without caffeine, we found no effect of consecutive foraging visits on the time the ants take to reach a reward, suggesting a failure to learn the reward’s location. However, under low to intermediate caffeine concentrations ants were 38% faster with each consecutive visit, implying that caffeine boosts learning. Interestingly, such improvements were lost at high doses. In contrast, caffeine had no impact on the ants’ homing behavior. Adding moderate levels of caffeine to baits could improve ant’s ability to learn its location, improving bait efficacy

Does the waggle dance help honey bees to forage at greater distances than expected for their body size?

A honey bee colony has been likened to an oil company. Some members of the company or colony prospect for valuable liquid resources. When these are discovered other group members can be recruited to exploit the resource. The recruitment of nestmates to a specific location where there is a patch of flowers should change the economics of scouting, that is, the search for new resource patches. In particular, communication is predicted to make scouting at longer distances worthwhile because a profitable resource patch, once discovered, will enhance the foraging not only of the discoverer but also of nestmates that can be directed to the patch. By virtue of having large colonies and dance communication, honey bees are predicted to be able to profitably scout, and hence forage, at greater distances from the nest than either solitary bees or social bees without communication. We test this hypothesis by first examining existing data on foraging distance to evaluate whether honey bees do indeed forage at greater distances than other bees given their body size. Second, we present a simple cost-benefit analysis of scouting which indicates that communication causes longer range scouting to be more profitable. Overall, our analyses are supportive, but not conclusive, that honey bees forage further than would be expected given their size and that the waggle dance is a cause of the honey bee's exceptional foraging range

Positive and negative incentive contrasts lead to relative value perception in ants

Humans usually assess things not according to their absolute value, but relative to reference points - a main tenant of Prospect Theory. For example, people rate a new salary relative to previous salaries and salaries of their peers, rather than absolute income. We demonstrate a similar effect in an insect: ants expecting to find low-quality food showed higher acceptance of medium-quality food than ants expecting medium quality, and vice versa for high expectations. Further experiments demonstrate that these contrast effects arise from cognitive rather than mere sensory or pre-cognitive perceptual causes. Social information gained inside the nest can also serve as a reference point: the quality of food received from other ants affected the perceived value of food found later. Value judgement is a key element in decision making, and thus relative value perception strongly influences which option is chosen and ultimately how all animals make decisions

Not dear neighbours: Antennation and jerking, but not aggression, correlate with genetic relatedness and spatial distance in the ant Lasius niger

1. Neighbour–stranger response differences (NSRDs) are when individuals are either more aggressive (“Nasty Neighbour”) or less aggressive (“Dear Enemy” or “Dear Neighbour”) to direct neighbours than to other competitors perceived as “strangers” by the residents. Such effects are often reported in ants which, being fixed-location central-place foragers, may compete directly with their neighbours for resources or raid each other for brood. Overlayed onto this are potential spatial distance and relatedness effects on aggression, which are often not differentiated from NSRDs. 2. The literature on NSRDs and distance effects in ants does not reveal a systematic pattern across all ants due to their diversity of life histories, requiring each species to be evaluated individually. Lasius niger is a common Eurasian ant species, which can form very dense populations of colonies and shows pronounced nestmate recognition, so may be expected to show NSRDs. 3. Here, we take advantage of a semi-regular colony array to examine the effect of spatial distance and relatedness on aggression and probe for NSRDs. 4. Overt aggression does not vary with relatedness or spatial distance, and there is no evidence that direct neighbours represent a special case in terms of aggression. However, antennation and jerking decrease between less related and more spatially distant pairs, but are almost completely absent from allospecific interactions. 5. We tentatively propose that antennation and jerking together represent a ‘negotiation’ phase, which may either precede or reduce the need for overt aggression. While a Nasty Neighbour effect might occur, a Dear Neighbour effect is unlikely in this species, and overall NSRDs do not play a large role in the ecology of this species. More broadly, this work highlights the importance of considering non-overtly aggressive responses when studying NSRDs