Many of the signals that animals use to communicate transmit relatively large distances and therefore encompass several potential signallers and receivers. This observation challenges the common characterization of animal communication systems as consisting of one signaller and one receiver. Furthermore, it suggests that the evolution of communication behaviour must be considered as occurring in the context of communication networks rather than dyads. Although considerations of selection pressures acting upon signallers in the context of communication networks have rarely been expressed in such terms, it has been noted that many signals exchanged during aggressive interactions will transmit far further than required for information transfer between the individuals directly involved, suggesting that these signals have been designed to be received by other, more distant, individuals. Here we consider the potential for receivers in communication networks to gather information, one aspect of which has been termed eavesdropping. We show that male Betta splendens monitor aggressive interactions between neighbouring conspeci¢cs and use the information on relative ¢ghting ability in subsequent aggressive interactions with the males they have observed.

Male singing behaviour correlates with extra-pair success in several passerine birds. Singing interactions during territorial contests provide relative information on the males involved. Such information may be important in female extra-pair behaviour and eavesdropping on singing interactions among males may allow females to make such relative assessments. We used interactive playback to instigate singing contests with male great tits during the peak fertile period of their mate in an attempt to alter females' assessment of mates' quality relative to neighbours (potential extra-pair partners). We escalated a contest to one male (by overlapping his songs) and then subsequently de-escalated a contest (by alternating) to a neighbour. Intrusions onto neighbouring territories by females mated to either treatment male were then monitored. Females mated to escalation treatment males were more likely to intrude following playbacks than females mated to de-escalation treatment males. Although the absolute song output of males did not di¡er between treatments, males produced more song relative to playback in de-escalation treatments and relative song output was positively correlated with female intrusions. Therefore, female great tits eavesdrop on singing interactions and change their visitation rates to neighbouring territories according to their mate's singing performance relative to neighbours.

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Communication and social behaviour are inextricably linked, with communication mediating important social behaviours such as resource defence and mate attraction. However, the social environment in which communication occurs is often ignored in discussions of communication behaviour. We argue that networks of several individuals are the common social environment for communication behaviour. The consequences for receivers and signallers of communicating in a network environment are the main subjects of this review. Eavesdropping is a receiving behaviour that is only possible in the environment of a network and therefore we concentrate on this behaviour. The main effect of communication networks on signallers is to create competition with other signallers for receiver attention. We discuss the consequences of such competition. To conclude, we explore the role of signals and signalling interactions as sources of information that animals exploit to direct their behaviour. IntroductionCommunication mediates virtually all important social behaviours. For communication to occur, at least two individuals, a signaller and a receiver, must be involved. Therefore communication can itself be considered a social behaviour. Given such obvious and strong links between communication and social behaviour, it is surprising how often the social environment of communication is ignored. For example, theoretical considerations of communication deal with a dyad of one signaller and one receiver (recent exceptions are Johnstone 1998, 2000; Hasson 2000) -social environment has literally been left out of the equation. The purpose of this review is to identify the major social environment in which communication occurs and to give a variety of examples of how this environment influences communication behaviour. The network environmentThe social environment in which much, if not all, communication occurs is a network of several signallers and receivers. For a long-range signal this assertion follows almost by definition, since such a signal will encompass several individuals during the course of transmission. The area encompassed is sometimes referred to as the signal's active space (e.g. Brenowitz 1982). This obvious association between long-range and large active space has perhaps deflected attention from the fact that it is the distance travelled by the signal in relation to the average spacing between individuals that is central to the idea of communication networks, rather than distance per se. Since many social groupings of animals are closely spaced, it is likely that many signals that are shortrange in an absolute sense can also encompass several signallers and receivers if these individuals occur sufficiently close together. Thus the commonest social environment in which communication occurs is a network (Dabelsteen 1992;McGregor 1993;McGregor and Dabelsteen 1996). Whether networks are ubiquitous still remains to be seen. Receivers and networksThe network environment creates costs and benefits for receivers in addition to those usua...

Animal culture, defined as "information or behavior-shared within a community-which is acquired from conspecifics through some form of social learning" (1), can have important consequences for the survival and reproduction of individuals, social groups, and potentially, entire populations (1, 2). Yet, until recently, conservation strategies and policies have focused primarily on broad demographic responses and the preservation of genetically defined, evolutionarily significant units. A burgeoning body of evidence on cultural transmission and other aspects of sociality (3) is now affording critical insights into what should be conserved (going beyond the protection of genetic diversity, to consider adaptive aspects of phenotypic variation), and why specific conservation programs succeed (e.g., through facilitating the resilience of cultural diversity) while others fail (e.g., by neglecting key repositories of socially transmitted knowledge). Here, we highlight how international legal instruments, such as the Convention on the Conservation of Migratory Species of Wild Animals (CMS), can facilitate smart, targeted conservation of a wide range of taxa, by explicitly considering aspects of their sociality and cultures. CONSEQUENCES OF SOCIAL KNOWLEDGE An important aspect of social learning is the speed with which new behaviors can potentially spread through populations, with effects that may be positive (e.g., adaptive exploitation of a new food source) or negative (e.g., increasing conflict with humans, such as when sperm whales learn to remove fish from longlines) (2). Transmission can be mediated by an inherent propensity to adopt innovations (e.g., "lobtail" feeding in humpback whales (1)), or curbed by cultural conservatism (e.g., southern resident killer whales' persistent foraging specialization on Chinook salmon (2)). Social learning can result in the emergence of subpopulations with distinctive behavioral profiles, erecting social barriers, as observed in distinct vocal clans of sperm whales (see the Figure). Culturally mediated population structure has important implications for conservation efforts (4), as it can influence species-wide phenotypic diversity and adaptability to changing conditions (5). In some cases, such as humpback or blue whale song, cultural variation can reflect demography and facilitate more efficient, or less invasive, assays of contemporary genetic population structure (1, 4). Most profoundly, culture can play a causal role in establishing and maintaining distinct evolutionary trajectories (6). Another consequence of social learning can be the increased importance of key individuals as repositories of accumulated knowledge, making their targeted protection particularly important for the persistence of social units. For example, the experience of African elephant matriarchs (see

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Reef manta rays (Mobula alfredi) are one of the ocean's largest and most charismatic species. Pressure from targeted and bycatch fisheries coupled with their conservative life‐history traits including slow growth, late maturity, and low fecundity has led to catastrophic declines of the global population. The species is now listed as Vulnerable to Extinction on IUCN Red List of Threatened Species. The global M. alfredi population is widely distributed in highly fragmented subpopulations. The Maldives supports the world's largest known subpopulation that undergoes seasonal migrations which are thought to be linked to peaks in ocean productivity induced by the South Asian Monsoon. Although the species is protected from targeted fisheries in the region, increasing pressures from habitat degradation and unsustainable tourism activities mean their effective conservation relies upon knowledge of the species' habitat use, seasonal distribution, and the environmental influences on such movements. Photo‐ID sighting records collected between 2005 and 2017 were used to identify key aggregation sites throughout the archipelago, and multiple linear regression and prediction analysis identified the environmental variables affecting variations in the intra‐annual sighting frequency of M. alfredi. Mobula alfredi were recorded at 273 different sites, 48 of which, with >100 sightings at each, were classified as key areas of habitat use. South‐west monsoon winds and chlorophyll‐a concentration predominantly affected the monthly percentage of M. alfredi sighted on the down‐current side of the atolls. In a country where climate change and touristic pressure are increasingly threatening this species and its habitat, the identification of key areas of habitat use and temporal changes in the use of these sites highlight the areas that should be prioritized for protection enabling more effective conservation management.

Animal communication generally occurs in the environment of a network of several potential signallers and receivers. Within a network environment, it is possible to gain relative information about conspeci¢cs by eavesdropping on signalling interactions. We presented male great tits with the opportunity to gain such information by simulating singing interactions using two loudspeakers. Interactions were presented so that relevant information was not available in the absolute singing behaviour of either individual, only in the relative timing of their songs in the interaction as a whole. We then assayed the information extracted by focal males by subsequently introducing one of the`interactants' (i.e. loudspeakers) into the territory of the focal male. Focal males responded with a reduced song output to males that had just`lost' an interaction. Focal males did not respond signi¢cantly di¡erently to`winners' as compared with intruders recently involved in an interaction that contained no consistent information. Focal males also responded by switching song types more often when encountering males that had recently been involved in a low-intensity interaction. These results provide the clearest evidence yet that male songbirds extract information from signal interactions between conspeci¢cs in the ¢eld.