Elise Nowbahari

Ants, Cataglyphis cursor, Use Precisely Directed Rescue Behavior to Free Entrapped Relatives

Although helping behavior is ubiquitous throughout the animal kingdom, actual rescue activity is particularly rare. Nonetheless, here we report the first experimental evidence that ants, Cataglyphis cursor, use precisely directed rescue behavior to free entrapped victims; equally important, they carefully discriminate between individuals in distress, offering aid only to nestmates. Our experiments simulate a natural situation, which we often observed in the field when collecting Catagyphis ants, causing sand to collapse in the process. Using a novel experimental technique that binds victims experimentally, we observed the behavior of separate, randomly chosen groups of 5 C. cursor nestmates under one of six conditions. In five of these conditions, a test stimulus (the “victim”) was ensnared with nylon thread and held partially beneath the sand. The test stimulus was either (1) an individual from the same colony; (2) an individual from a different colony of C cursor; (3) an ant from a different ant species; (4) a common prey item; or, (5) a motionless (chilled) nestmate. In the final condition, the test stimulus (6) consisted of the empty snare apparatus. Our results demonstrate that ants are able to recognize what, exactly, holds their relative in place and direct their behavior to that object, the snare, in particular. They begin by excavating sand, which exposes the nylon snare, transporting sand away from it, and then biting at the snare itself. Snare biting, a behavior never before reported in the literature, demonstrates that rescue behavior is far more sophisticated, exact and complexly organized than the simple forms of helping behavior already known, namely limb pulling and sand digging. That is, limb pulling and sand digging could be released directly by a chemical call for help and thus result from a very simple mechanism. However, its difficult to see how this same releasing mechanism could guide rescuers to the precise location of the nylon thread, and enable them to target their bites to the thread itself.

Pro-sociality without empathy

Empathy, the capacity to recognize and share feelings experienced by another individual, is an important trait in humans, but is not the same as pro-sociality, the tendency to behave so as to benefit another individual. Given the importance of understanding empathys evolutionary emergence, it is unsurprising that many studies attempt to find evidence for it in other species. To address the question of what should constitute evidence for empathy, we offer a critical comparison of two recent studies of rescuing behaviour that report similar phenomena but are interpreted very differently by their authors. In one of the studies, rescue behaviour in rats was interpreted as providing evidence for empathy, whereas in the other, rescue behaviour in ants was interpreted without reference to sharing of emotions. Evidence for empathy requires showing that actor individuals possess a representation of the receivers emotional state and are driven by the psychological goal of improving its wellbeing. Proving psychological goal-directedness by current standards involves goal-devaluation and causal sensitivity protocols, which, in our view, have not been implemented in available publications. Empathy has profound significance not only for cognitive and behavioural sciences but also for philosophy and ethics and, in our view, remains unproven outside humans.

Effect of body size on aggression in the ant, Cataglyphis niger (Hymenoptera; Formicidae)

In polymorphic ants, such as Cataglyphis niger, sterile individual workers from the same nest show some degree of variation in size and/or morphology. We studied whether worker size and size difference between opponents had an effect on aggression during conspecific encounters. Although the capacity to recognize nestmates was shared by all individuals, some patterns of agonistic behaviors were size related. Escape was mostly displayed by the small workers, and threat, associated with ritualized fights, by the large workers. As game theory predicted, ants of C. niger adjusted their level of aggression as a function of the size of the opponent. However, only large individuals used such assessment strategies, responding with escalation of aggression towards small workers and reduction of aggression towards large ones. On the contrary, small individuals behaved in the same manner whatever the opponents size. Differences between both morphological castes were discussed with reference to the resource holding assessment models. Aggr. Behav. 25:369–379, 1999.

Rescue behavior: Distinguishing between rescue, cooperation and other forms of altruistic behavior

Reports of rescue behavior in non-human animals are exceedingly rare, except in ants where rescue is well known, but has not been explored experimentally until recently. Although we predict that rescue behavior should be limited to circumstances in which the victim and the rescuer are highly related to one another, or in which unrelated individuals must cooperate very closely with one another, we also predict that it is likely to be far more common than the current literature would suggest. To address this oversight, we propose a rigorous definition of rescue behavior, one that helps researchers to focus on its necessary and sufficient components, at the same time that it helps to differentiate rescue behavior from cooperation and other forms of helping behavior. In this way we also hope to expand our understanding of altruism in particular and kin selection in general.

Learning of colonial odor in the antCataglyphis niger (Hymenoptera; Formicidae)

Ants learn the odors of members of their colony early in postnatal life, but their ability to learn to recognize noncolony conspecifics and heterospecifics has never been explored. We used a habituation—discrimination paradigm to assess individual recognition in adult Formicine ants,Cataglyphis niger. Pairs of workers from different colonies were placed together for repeated trials, and their ability to discriminate the ant that they encountered from another familiar or unfamiliar ant was observed. Some ants were isolated between encounters, and others were returned to their home colonies. Our results suggest for the first time in ants that C. niger adults learn about individual ants that they have encountered and recognize them in subsequent encounters. Ants are less aggressive toward non-nestmates after they are familiar with one another, but they are aggressive again when they encounter an unfamiliar individual. Learning about non-nestmates does not interfere with an ant’s memory of members from its own colony.

Toward a Behavioral Ecology of Rescue Behavior

Although the study of helping behavior has revolutionized the field of behavioral ecology, scientific examination of rescue behavior remains extremely rare, except perhaps in ants, having been described as early as 1874. Nonetheless, recent work in our laboratories has revealed several new patterns of rescue behavior that appear to be much more complex than previously studied forms. This precisely-directed rescue behavior bears a remarkable resemblance to what has been labeled empathy in rats, and thus raises numerous philosophical and theoretical questions: How should rescue behavior (or empathy) be defined? What distinguishes rescue from other forms of altruism? In what ways is rescue behavior in ants different from, and similar to, rescue in other non-human animals? What selection pressures dictate its appearance? In this paper, we review our own experimental studies of rescue in both laboratory and field, which, taken together, begin to reveal some of the behavioral ecological conditions that likely have given rise to rescue behavior in ants. Against this background, we also address important theoretical questions involving rescue, including those outlined above. In this way, we hope not only to encourage further experimental analysis of rescue behavior, but also to highlight important similarities and differences in very distant taxa.

Memory span for heterospecific individuals’ odors in an ant, Cataglyphis cursor

Only recently have researchers studied the ability of ants to learn and remember individual heterospecific odors. Cataglyphis cursor adults have the capacity to learn these odors, but the duration of their memory and the factors that affect its formation remain unknown. We used a habituation/discrimination paradigm to study some of these issues. C. cursor adult workers were familiarized to an anesthetized Camponotus aethiops on four successive encounters. Then they were either isolated or placed with 20 nestmates for a certain length of time before undergoing a discrimination test that consisted of reintroducing the familiar C. aethiops, as well as introducing an unknown member of the same colony. The results showed that adult C. cursor ants can retain in memory a complex individual odor for at least 30 min, as well as differentiate it from the odor of another closely related individual. However, when ants were replaced in a rich social background between the habituation and the discrimination trials, we did not observe a significant discrimination between the known and unknown C. aethiops ants. Our study shows, for the first time, the existence of long-term memory for individual odors in mature ant workers.

Rescue of newborn ants by older Cataglyphis cursor adult workers.

Cataglyphiscursor worker ants are capable of highly sophisticated rescue behaviour in which individuals are able to identify what has trapped a nestmate and to direct their behaviour towards that obstacle. Nonetheless, rescue behaviour is constrained by workers’ subcaste: whereas foragers, the oldest workers, are able both to give and to receive the most help, the youngest workers, inactives, neither give nor receive any help whatsoever; nurses give and receive intermediate levels of aid, reflecting their intermediate age. Such differences in rescue behaviour across subcastes suggest that age and experience play a critical role. In this species, as in many others in which a sensitive period for nestmate recognition exists, newly enclosed ants, called callows, are adopted by ants belonging not only to different colonies but also to different species; foreign callows receive nearly the same special care provided to resident newborns. Because callows are younger than inactives, which are incapable of soliciting rescue, we wondered whether entrapped callows would receive such aid. In the present study, we artificially ensnared individual callows from their own colony (homocolonial), from a different colony (heterocolonial), and from a different species (heterospecific), and tested each one with groups of five potential C.cursor rescuers, either all foragers or all nurses. Our results show that all three types of callows are able to elicit rescue behaviour from both foragers and nurses. Nonetheless, nurse rescuers are better able to discriminate between the three types of callow victims than are foragers.

Reproductive hierarchies and status discrimination in orphaned colonies of Pachycondyla apicalis ants

In group-living animals where dominance hierarchies occur, aggression can be reduced if individuals are able to recognize each other. To do this, and to adapt their behaviour suitably when faced with a rival, individuals may rely on two nonmutually exclusive recognition means: they could recognize group members individually and/or their social status. Within insect societies, although conflicts over reproduction resulting in hierarchy establishment are widespread, relatively little is known about the cognitive abilities involved in the regulation of agonistic interactions. We tested whether low-ranking workers of Pachycondyla apicalis ants are able to discriminate each other individually and/or if they can discriminate the status of their nestmates. We found no evidence of individual discrimination among subordinates whereas they were able to discriminate their nestmates on the basis of their social and reproductive status. Such a skill may allow them to regulate worker reproduction in queenright colonies efficiently. By considering the structure of the hierarchy and the nature of the dominance relationships in P. apicalis societies, we discuss the existence of a more accurate recognition system among the high-ranking workers.

Organization of rescue behaviour sequences in ants, Cataglyphis cursor, reflects goal-directedness, plasticity and memory

The experimental study of rescue behaviour in ants, behaviour in which individuals help entrapped nestmates in distress, has revealed that rescuers respond to victims with very precisely targeted behaviour. In Cataglyphis cursor, several different components of rescue behaviour have been observed, demonstrating the complexity of this behaviour, including sand digging and sand transport to excavate the victim, followed by pulling on the victims limbs as well as the object holding the victim in place, behaviour that serves to free the victim. Although previous work suggested that rescue was optimally organized, first to expose and then to extricate the victim under a variety of differing circumstances, experimental analysis of that organization has been lacking. Here, using experimental data, we characterize the pattern of individual rescue behaviour in C. cursor by analysing the probabilities of transitions from one behavioural component to another. The results show that the execution of each behavioural component is determined by the interplay of previous acts. In particular, we show not only that ants move sand away from the victim in an especially efficient sequence of behaviour that greatly minimizes energy expenditure, but also that ants appear to form some kind of memory of what they did in the past, a memory that directs their future behaviour.