A positive effect of cumulative intergroup threat on reproductive success.

Outgroup conflict is a powerful selective force across all social taxa. While it is well documented that individual outgroup contests can have a range of direct and indirect fitness consequences, the cumulative pressure of outgroup threats could also potentially impact reproductive success. Here, we use long-term life-history data from a wild population of dwarf mongooses (Helogale parvula) to investigate how intergroup interaction (IGI) rate might influence breeding and offspring survival. IGI rate did not predict the number of litters produced in a season or the inter-litter interval. Unexpectedly, IGI rate was positively associated with the number of pups alive three months after emergence from the breeding burrow. This was not due to a difference in how many pups emerged but because those in groups experiencing more IGIs had a higher survival likelihood post-emergence. Detailed natural observations revealed that both IGI occurrence and the threat of intergroup conflict led to more sentinel behaviour by adults, probably reducing the predation risk to young. Our results contrast the previously documented negative effects of outgroup interactions on reproductive success and highlight the need to assess cumulative threat, rather than just the impact of physical contests, when considering outgroup conflict as a social driver of fitness.

Behavioural, demographic and fitness consequences of social instability in cooperatively breeding dwarf mongoose groups.

Social instability frequently arises in group-living species, but the potential costs have rarely been investigated in free-living cooperative breeders, especially across different timeframes. Using natural observations, body mass measurements and life-history data from dwarf mongooses (Helogale parvula), we determined the short- and long-term consequences of a change in one of the dominant breeding pairs. We found that a new breeder led to alterations in both collective and individual behaviours (i.e. increases in communal scent-marking, engagement in intergroup interactions, sentinel activity and within-group grooming), as well as reduced body mass gain, further demographic changes and decreased reproductive success (i.e. fewer pups surviving to adulthood). The effects were particularly apparent when it was the female breeder who changed; new female breeders were younger than more experienced counterparts. Our findings support the idea that stability and cooperation are strongly linked and provide potential reasons for previously documented health and fitness benefits of social stability.

Fitness consequences of outgroup conflict.

In social species across the animal kingdom, conspecific outsiders threaten the valuable resources of groups and their members. This outgroup conflict is recognised as a powerful selection pressure, but we argue that studies explicitly quantifying the fitness consequences need to be broader in scope: more attention should be paid to delayed, cumulative, and third-party fitness consequences, not just those arising immediately to group members involved in physical contests. In the first part of this review, we begin by documenting how single contests can have survival and reproductive consequences either immediately or with a delay. Then, we step beyond contests to describe fitness consequences that can also result from interactions with cues of rival presence and the general landscape of outgroup threat, and beyond single interactions to describe cumulative effects of territorial pressure and elevated outgroup-induced stress. Using examples from a range of taxa, we discuss which individuals are affected negatively and positively, considering both interaction participants and third-party group members of the same or the next generation. In the second part of the review, we provide suggestions about how to move forward. We highlight the importance of considering how different types of outgroup conflict can generate different selection pressures and of investigating variation in fitness consequences within and between species. We finish by discussing the value of theoretical modelling and long-term studies of natural populations, experimental manipulations, and meta-analyses to develop further our understanding of this crucial aspect of sociality.

Variation between species, populations, groups and individuals in the fitness consequences of out-group conflict.

Out-group conflict is rife in the natural world, occurring from primates to ants. Traditionally, research on this aspect of sociality has focused on the interactions between groups and their conspecific rivals, investigating contest function and characteristics, which group members participate and what determines who wins. In recent years, however, there has been increasing interest in the consequences of out-group conflict. In this review, we first set the scene by outlining the fitness consequences that can arise immediately to contest participants, as well as a broader range of delayed, cumulative and third-party effects of out-group conflict on survival and reproductive success. For the majority of the review, we then focus on variation in these fitness consequences of out-group conflict, describing known examples both between species and between populations, groups and individuals of the same species. Throughout, we suggest possible reasons for the variation, provide examples from a diverse array of taxa, and suggest what is needed to advance this burgeoning area of social evolution. This article is part of the theme issue 'Intergroup conflict across taxa'.

Extended and cumulative effects of experimentally induced intergroup conflict in a cooperatively breeding mammal.

Conflict between rival groups is rife in nature. While recent work has begun exploring the behavioural consequences of this intergroup conflict, studies have primarily considered just the 1-2 h immediately after single interactions with rivals or their cues. Using a habituated population of wild dwarf mongooses (Helogale parvula), we conducted week-long manipulations to investigate longer-term impacts of intergroup conflict. Compared to a single presentation of control herbivore faeces, one rival-group faecal presentation (simulating a territorial intrusion) resulted in more within-group grooming the following day, beyond the likely period of conflict-induced stress. Repeated presentations of outsider cues led to further changes in baseline behaviour by the end of the week: compared to control weeks, mongooses spent less time foraging and foraged closer to their groupmates, even when there had been no recent simulated intrusion. Moreover, there was more baseline territorial scent-marking and a higher likelihood of group fissioning in intrusion weeks. Consequently, individuals gained less body mass at the end of weeks with repeated simulated intrusions. Our experimental findings provide evidence for longer-term, extended and cumulative, effects of an elevated intergroup threat, which may lead to fitness consequences and underpin this powerful selective pressure.

Experimental evidence for delayed post-conflict management behaviour in wild dwarf mongooses.

In many species, within-group conflict leads to immediate avoidance of potential aggressors or increases in affiliation, but no studies have investigated delayed post-conflict management behaviour. Here, we experimentally test that possibility using a wild but habituated population of dwarf mongooses (Helogale parvula). First, we used natural and playback-simulated foraging displacements to demonstrate that bystanders take notice of the vocalisations produced during such within-group conflict events but that they do not engage in any immediate post-conflict affiliative behaviour with the protagonists or other bystanders. We then used another playback experiment to assess delayed effects of within-group conflict on grooming interactions: we examined affiliative behaviour at the evening sleeping burrow, 30-60 min after the most recent simulated foraging displacement. Overall, fewer individuals groomed on evenings following an afternoon of simulated conflict, but those that did groomed more than on control evenings. Subordinate bystanders groomed with the simulated aggressor significantly less, and groomed more with one another, on conflict compared to control evenings. Our study provides experimental evidence that dwarf mongooses acoustically obtain information about within-group contests (including protagonist identity), retain that information, and use it to inform conflict-management decisions with a temporal delay.

Social animals that live in groups often have disagreements over access to mates and food. Even fleeting in-group disputes can be costly, disrupting relationships, wasting time and energy, or causing injury if aggression escalates. So, much like humans, many social animals, including primates, birds and dogs, have evolved conflict management strategies to prevent and resolve in-group disagreements. In the immediate aftermath of a conflict, this usually involves changes in the interactions between those involved in the disagreement, or between bystander groupmates and either the victim or aggressor. Less is known about whether social animals can recall past disputes and if they can use conflict management strategies some time after a quarrel has occurred. That is, do aggressive interactions between groupmates influence later social decisions of bystanders in the group? To investigate, Morris-Drake et al. studied groups of wild dwarf mongooses (Helogale parvula) that have become accustomed to living alongside humans in Limpopo Province, South Africa. Dwarf mongooses live in groups of up to 30 individuals, with one dominant breeding pair and lower-ranked helpers. When disagreements arise over food, an aggressor growls deeply and hip-slams the victim away from their foraging patch, stealing the victim's prey in the process. Victims often produce high-pitched squeals in retreat. Using recordings of these calls, Morris-Drake et al. devised a field experiment to investigate how mongooses responded to nearby conflicts between other group members. Recordings simulating a conflict over food were played to groups of foraging mongooses over the course of an afternoon, so that group members effectively heard what sounded like repeated squabbles between two out-of-sight individuals. Similar to natural conflicts, the mongooses did not engage in any obvious conflict management behaviour immediately after hearing these disputes. But when the group returned to their sleeping burrow that evening, subordinate group members shunned the perceived aggressors from grooming, a key social activity. This work provides evidence that dwarf mongooses keep tabs on conflicts that occur between groupmates. It shows these animals can extract information about conflicts from vocal cues alone and that bystanders use this information when making later social decisions impacting group dynamics. It also adds to growing evidence from baboons, monkeys and chimpanzees that social animals can remember past events and take these into account when interacting with groupmates.

Experimental field evidence that out-group threats influence within-group behavior.

In social species, conspecific outsiders present various threats to groups and their members. These out-group threats are predicted to affect subsequent within-group interactions (e.g., affiliation and aggression) and individual behavior (e.g., foraging and vigilance decisions). However, experimental investigations of such consequences are rare, especially in natural conditions. We used field-based call playbacks and fecal presentations on habituated wild dwarf mongooses (Helogale parvula)-a cooperatively breeding, territorial species-to examine postinteraction responses to the simulated threat of a rival group. Dwarf mongooses invested more in grooming of groupmates, foraged closer together, and more regularly acted as sentinels (a raised guard) after encountering indicators of rival-group presence compared to control conditions. These behavioral changes likely arise from greater anxiety and, in the case of increased vigilance, the need to seek additional information about the threat. The influence of an out-group threat lasted at least 1 h but individuals of different dominance status and sex responded similarly, potentially because all group members suffer costs if a contest with rivals is lost. Our results provide field-based experimental evidence from wild animals that out-group threats can influence within-group behavior and decision making, and suggest the need for greater consideration of the lasting impacts of social conflict.

Anthropogenic noise alters dwarf mongoose responses to heterospecific alarm calls.

Anthropogenic noise is an evolutionarily novel and widespread pollutant in both terrestrial and aquatic habitats. Despite increasing evidence that the additional noise generated by human activities can affect vocal communication, the majority of research has focused on the use of conspecific acoustic information, especially sexual signals. Many animals are known to eavesdrop on the alarm calls produced by other species, enhancing their likelihood of avoiding predation, but how this use of heterospecific information is affected by anthropogenic noise has received little empirical attention. Here, we use two field-based playback experiments on a habituated wild population of dwarf mongooses (Helogale parvula) to determine how anthropogenic noise influences the response of foragers to heterospecific alarm calls. We begin by demonstrating that dwarf mongooses respond appropriately to the alarm calls of sympatric chacma baboons (Papio ursinus) and tree squirrels (Paraxerus cepapi); fleeing only to the latter. We then show that mongoose foragers are less likely to exhibit this flee response to tree squirrel alarm calls during road-noise playback compared to ambient-sound playback. One explanation for the change in response is that noise-induced distraction or stress result in maladaptive behaviour. However, further analysis revealed that road-noise playback results in increased vigilance and that mongooses showing the greatest vigilance increase are those that do not subsequently exhibit a flee response to the alarm call. These individuals may therefore be acting appropriately: if the greater gathering of personal information indicates the absence of an actual predator despite an alarm call, the need to undertake costly fleeing behaviour can be avoided. Either way, our study indicates the potential for anthropogenic noise to interfere with the use of acoustic information from other species, and suggests the importance of considering how heterospecific networks are affected by this global pollutant.

Cross-modal impacts of anthropogenic noise on information use.

Anthropogenic noise is a global pollutant, and there is rapidly accumulating evidence of impacts on a range of animal taxa [1,2]. While many studies have considered how additional noise may affect information provision and use, they have focused on the masking and consequent alteration of acoustic signals and cues; so-called unimodal effects [3]. Using field-based experimental trials on habituated wild dwarf mongooses (Helogale parvula) [4], we combine sound playbacks and faecal presentations to demonstrate that anthropogenic noise can disrupt responses to information from different sensory modalities. The adaptive, stronger response exhibited towards predator faeces compared with control faeces in ambient-noise conditions was detrimentally affected by road-noise playback. Specifically, having taken longer to detect the faeces, the mongooses interacted less with the predator cue, did not show increased vigilance following its detection, and spent less time in the safe vicinity of a burrow refuge, thus suffering a potentially increased predation risk. Our results are the first to show that anthropogenic noise could alter responses to olfactory cues, strongly indicating the possibility of cross-modal impacts of noise pollution on information use [3].