Greater spear-nosed bats give group-distinctive calls

Individually distinctive vocalizations are ubiquitous; however, group distinctive calls have rarely been demonstrated. Under some conditions, selection should favor calls indicating social group membership in animals that forage in groups. Greater spear-nosed bats (Phyllostomus hastatus) give calls that appear to facilitate recognition of social group mates who are unrelated. Females give loud broadband (4–18 kHz) vocalizations termed screech calls when departing on foraging trips and at foraging sites. Screech calls help to establish foraging groups among social group members, and to maintain contact over the long distances they travel while foraging. I test two hypotheses about how screech calls may be structured to convey caller identity. Individual calls may be distinct and group members may learn to recognize each individual's calls and to associate the individual with the social group. Alternatively, groups may give distinct calls and individuals within groups may share call characteristics. To test these hypotheses I conducted multivariate acoustic analysis of multiple calls from 28 bats from three social groups. Although the ubiquity of individually distinctive calls in other taxa makes this result more likely, the results reveal that group calls are highly distinctive. Individual bats within groups are statistically indistinguishable. Calls appear to decrease slightly in frequency as bats age. Call convergence among unrelated group mates implies vocal learning in this species. Received: 28 March 1996 / Accepted after revision: 6 October 1996     

Flexible bat echolocation: the influence of individual,habitat and conspecifics on sonar signal design

Acoustic signals which are used in animal communication must carry a variety of information and are therefore highly flexible. Echolocation has probably such functions and could prove as flexible. Measurable variabitlity can indicate flexibility in a behaviour. To quantify variability in bat sonar and relate to behavioural and environmental factors, I recorded echolocation calls of Euderma maculatum, Eptesicus fuscus, Lasiurus borealis and L. cinereus while the bats hunted in their natural habitat. I analysed 3390 search phase calls emitted by 16 known and 16 unknown individuals foraging in different environmental and behvioural situations. All four species used mainly multiharmonic signals that showed considerable intra- and inter-individual variability in the five signal variables I analysed (call duration, call interval, highest and lowest frequency and frequency with maximum energy) and also in the shape of the sonagram. A nested multivariate analysis of variance identified the influences of individual, hunting site, close conspecifics and of each observation on the frequency with maximum energy in the calls, and on other variables measured. Individual bats differed in multiple comparisons, most often in the main call frequency and least often in call interval. In a discriminant function analysis with resubstitution, 56–76% of a species' calls were assigned to the correct individual. Distinct individual call patterns were recorded in special situations in all species and the size of foraging areas in forested areas influenced temporal and spectral call structure. Echolocation behaviour was influenced by the presence of conspecifics. When bats were hunting together, call duration decreased and call interval increased in all species, but spectral effects were less pronounced. The role of morphometric differences as the source of individually distinct vocalizations is discussed. I also examined signal adaptations to long range echolocation and the influence of obstacle distance on echolocation call design. My results allow to discuss the problems of echo recognition and jamming avoidance in vespertilionid bats.     

Do echolocation calls of wild colony-living Bechstein's bats (Myotis bechsteinii) provide individual-specific signatures?

Social animals often use vocal communication signals that contain individual signatures. As bats emit echolocation calls several times per second to orient in space, these might seem ideal candidates for conveying the caller's individual identity as a free by-product. From a proximate perspective, however, coding of caller identity is hampered by the simple acoustic structure of echolocation signals, by their task-specific design and by propagation loss. We investigated the occurrence of individual signatures in echolocation calls in individually marked, free-living Bechstein's bats (Myotis bechsteinii) in a situation with defined social context in the field. The bats belonged to two different colonies, for both of which genetic data on relatedness structure was available. While our data clearly demonstrate situation specificity of call structure, the evidence for individual-specific signatures was relatively weak. We could not identify a robust and simple parameter that would convey the caller's identity despite the situation-specific call variability. Discriminant function analysis assigned calls to call sequences with good performance, but worsened drastically when tested with other sequences from the same bats. Therefore, we caution against concluding from a satisfactory discrimination performance with identical training and test sequences that individual bats can reliably be told apart by echolocation calls. At least the information contained in a single call sequence seems not to be sufficient for that purpose. Starting frequencies did give the best discrimination between individuals, and it was also this parameter that was correlated with genetic relatedness in one of our two study colonies. Echolocation calls could serve as an additional source of information for individual recognition in Bechstein's bats societies, while it is unlikely that a large number of individuals could be reliably identified in different situations based on echolocation alone.     

Individual recognition and potential recognition errors in parent–offspring communication

The recognition of food-provisioning parents is crucial for fledglings of many bird species. Vocalizations are the most commonly used cues in avian parent–offspring communication, and it has been shown in several species that fledglings respond specifically to their parents' contact calls. However, fledglings occasionally also react to unrelated adults. Such responses may reflect recognition errors or alternatively a strategy of fledglings to obtain food or other direct benefits from unrelated adult birds. In a playback experiment, we tested whether zebra finch Taeniopygia guttata fledglings perceive variation in adult call signatures to recognize their parents and whether the propensity to respond to unrelated individuals is related to the gender of adults and to signal properties of male and female calls. Male calls are learnt and show high intra-sexual variation, which may improve the accurate recognition of the father's individual signature. In contrast, calls of adult females are innate, show lower intra-sexual variation such that the mother's call is more likely to be confused with another female call. We demonstrate that fledglings are able to recognize their parents. In addition, fledglings reacted more strongly to unrelated females compared with unrelated males. Our findings suggest that responses to unrelated adults may reflect recognition errors and indicate the importance of variation in identity signals for individual recognition processes in parent–offspring communication.     

Female preferences for multiple attributes in the acoustic signals of the Italian treefrog, <Emphasis Type="Italic">Hyla intermedia</Emphasis>

The advertisement call of frogs and toads is an example of multiple message signal because different acoustic properties encode different kinds of biologically significant information. In the Italian treefrog, Hyla intermedia, pulse rate and frequency have been found to be under stabilizing female preferences and to encode information important for mate recognition, whereas the number of calls per call group have been found to be under directional preferences and, thus, to be important for mate quality assessment. In this study, we investigate preferences for calls that differ simultaneously in frequency, pulse rate, and number of calls per call group, and we ask how these properties interact with each other in influencing female mating decisions. Results of two-choice phonotaxis experiments provide no evidence to support the hypothesis that females process multi-attribute signals in a hierarchical way. In contrast, the pattern of preferences is consistent with the ‘preference function’ hypothesis, that is, with the hypothesis that females rank signals along an ordinal scale of values and choose accordingly. Pulse rate and frequency influence mating preferences more than does the number of calls per call group. The interaction between pulse rate and frequency is not additive but multiplicative: small differences in either pulse rate or frequency that, alone, have no effects on female choice, interact synergistically so that their combination has strong influence on female preferences. A preference repeatability test shows strong among-female differences in preference for multi-attribute signals. We suggest that this result reveals not only a variation in attribute values among females, but also a variation in the way females weight and combine attribute values into a single preference score.     

The role of synchronized calling,ambient light,and ambient noise,in anti-bat-predator behavior of a treefrog

Summary Male treefrogs, Smilisca sila (Hylidae), produce calls of varying complexity and demonstrate a remarkable ability to synchronize their calls with those of neighbors. The bat Trachops cirrhosus eats frogs and uses the frogs' advertisement calls as locational cues. The bats are less likely to respond to synchronous calls than to asynchronous calls, and when given a choice prefer complex calls to simple calls.Experiments with bat models indicate that, like other frogs, S. sila probably uses visual cues to detect hunting bats. In response to bat models the frogs decreased both the number and the complexity of their calls. The calling behavior of the frogs was sampled in the field during periods with and without artificial illumination. The frogs produced fewer and less complex calls, and they tended to call from more concealed sites, during the period without illumination, when presumably it would have been more difficult for the frogs to detect hunting bats. S. sila tended to call from sites with higher ambient noise level, the noise primarily originating from waterfalls. The frequencies of the dominant energies in the waterfall sounds completely overlapped the frequency range of the S. sila call; thus waterfalls might mask the frog calls. When given a choice between calls produced near and away from waterfall sounds, bats preferred the latter.     

Similar is not the same: Social calls of conspecifics are more effective in attracting wild bats to day roosts than those of other bat species

Many bat species regularly need to find new day roosts as they require numerous shelters each breeding season. It has been shown that bats exchange information about roosts among colony members, and use echolocation and social calls of conspecifics in order to find roosts. However, it is unclear if wild bats discriminate between social calls of conspecifics and other bat species while searching for roosts. Furthermore, the extent that bats are attracted to potential roosts by each of these two call types is unknown. We present a field experiment showing that social calls of conspecifics and other bat species both attract bats to roosts. During two summers, we played back social calls of Bechstein’s bats (Myotis bechsteinii) and Natterer’s bats (Myotis nattereri) from different bat boxes that can serve as roosts for these species. All experimental bat boxes were monitored with infrared video to identify the approaching bat species. Three species (M. bechsteinii, M. nattereri, and Plecotus auritus) approached the boxes significantly more often during nights when bat calls were played compared to nights without playbacks. Bechstein’s bats and Natterer’s bats were both more attracted to social calls of conspecifics than of the other species, whereas P. auritus did not discriminate between calls of either Myotis species. Only Bechstein’s bats entered experimental boxes and only at times when calls from conspecifics were played. Our findings show that wild bats discriminate between social calls of conspecifics and other bat species although they respond to both call types when searching for new roosts.     

Discrimination of different social companions in spectacled parrotlets (Forpus conspicillatus): evidence for individual vocal recognition

Individual recognition is generally assumed to be a prerequisite for establishing and maintaining a complex social system. Indeed, there is good evidence that highly social species have complex systems of vocal communication with individual recognition by acoustic cues. In this study, we provide experimental evidence that vocal class and individual recognition is present in a non-passerine bird, the spectacled parrotlet (Forpus conspicillatus). Spectacled parrotlets live in a complex system of social relationships. Soon after fledging, the young establish close sibling relationships which are important for successful socialization, pairing and reproduction. In a series of playback experiments we tested if spectacled parrotlets use contact calls for vocal recognition. The results showed that spectacled parrotlets discriminate between the contact calls of different social categories. Adult birds preferred to respond to the contact calls of their mates. Subadult individuals recognized the contact calls of their siblings. During the period of pair bond formation, the affiliative contacts to the siblings decrease, but the parrotlets continue to respond to the calls of their siblings. This is the first evidence that vocal sibling recognition might outlast the period of strong sibling interaction and extends into the period of pair bond formation. In cases of mate loss or divorce, the acoustic contact to their siblings might facilitate the re-establishment of close sibling relationships. Received: 29 October 1997 / Accepted after revision: 5 April 1998     

Loud calls as a mechanism of social coordination in a fission–fusion taxon, the white-bellied spider monkey (Ateles belzebuth)

Spider monkeys (Ateles spp.) live in social groups that exhibit high levels of fission–fusion dynamics, in which group members form subgroups of varying sizes and compositions. Within these fluid societies, how individuals establish contact with dispersed group members with whom they might choose to associate remains unclear. Long-range vocalizations might facilitate interactions between group members and provide a means of social coordination in fission–fusion societies. We evaluated this possibility for one spider monkey vocalization, the loud call, by examining calling behavior, the relationship between loud calls and changes in subgroup size, and the response of individuals to distant calls and playback experiments in a single study group. We found that 82 % of loud calls were emitted within 30 min of a call from a different location, suggesting that individuals frequently emit loud calls in response to the calls of distant group members. Subgroups that emitted loud calls, especially those that responded to distant calls, were much more likely to experience an increase in subgroup size within an hour after calling than those that did not. Animals also approached distant loud calls more than they avoided or ignored these calls. Finally, playbacks of male calls demonstrated that females respond preferentially to the calls of some individuals over others. Taken together, these results provide support for the hypothesis that spider monkey loud calls function to facilitate and initiate interactions between dispersed group members and suggest that vocal signals can play an important role in influencing social interactions in fission–fusion societies.     

Species-specificity and individual variation in the song of male Nathusius’ pipistrelles (<Emphasis Type="Italic">Pipistrellus nathusii</Emphasis>)

Sequences of the advertisement calls produced by male Nathusius’ pipistrelles (Pipistrellus nathusii) during the autumn mating period were recorded from individuals at two separate sites in Antrim, Northern Ireland, in August 2004. Several male roosts were found at these sites in close proximity to a single maternity roost, each containing approximately 200 adult females and their young. Analysis of measured parameters of four identified call types revealed that there were significant differences in call structure between sites and between individuals. Playback experiments, performed outside the adult female and juvenile roost sites, comprised of experimental advertisement call sequences of P. nathusii, Pipistrellus pygmaeus and Pipistrellus pipistrellus and control sound recorded without bats present (silence). Response was measured by simultaneously recording ultrasound during playbacks and counting the number of echolocation pulses identified as those of P. nathusii above a predetermined amplitude threshold. Significantly greater numbers of P. nathusii echolocation pulses were recorded during playback of male P. nathusii advertisement calls than during playback of congeners’ advertisement calls and control sound. The number of echolocation pulses recorded was similar during playback of P. pipistrellus and P. pygmaeus advertisement calls and silence. We suggest that, due to call complexity, male P. nathusii advertisement calls should be classified as ‘song’. Species-specificity and individual variation suggests that the songs of male P. nathusii have the potential to play a role in mate attraction and mate assessment.     

Individual differences and within-flock convergence in chickadee calls

Summary A detailed sound analysis of the Chick-adee call of the black-capped chickadee (Parus atricapillus) was performed in order to determine a basis for individual recognition and for imitation within winter flocks. During the winter of 1978–1979 members of five free-living black-capped chickadee flocks were uniquely marked for individual identification, and their calls were recorded in the field. Nested analysis of variance of temporal call parameters measured from sonagrams and spectral parameters from frequency spectra showed that there were significant differences between individuals within flocks for every parameter measured. There were significant differences between flocks in the frequency ranges 800–2,200 Hz and 4,000–5,300 Hz, in the spectral parameters bandwidth and maximum frequency, and in the duration of the Dee syllable and total call duration.Members of four of the five flocks were captured in December 1978 and held in aviaries segregated by original flock membership. In January 1979 the memberships of the three aviaries were rearranged to form experimental flocks. After one month, there were significant differences among the experimental flocks in the duration of the Dee syllable and total call duration. Convergence, as indicated by a significant decrease in variance among members (F-test), occurred in the experimental flock in Aviary 1 and was concentrated in the frequency ranges 1,300–1,800 and 6,200–6,900 Hz. The members of this experimental flock differed from each other in the number of 100 Hz frequency intervals within which each changed its own call.The Chick-a-dee call contains sufficient information that it can potentially be used by black-capped chickadees for individual recognition. In addition, both field and aviary data suggest that flock members converge in some call characteristics. Possible explanations of the social significance of vocal convergence in chickadee flocks are discussed.     

Behavioral responses to echolocation calls from sympatric heterospecific bats: implications for interspecific competition

Mutual recognition is the product of species coexistence, and has direct effects on survival and reproduction of animals. Bats are able to discriminate between sympatric different heterospecifics based on their echolocation calls, which has been shown both in free-flying and captive bats. To date, however, the factors that may determine the behavioral responses of bats to echolocation calls from sympatric heterospecifics have rarely been tested, especially under well-controlled conditions in captive bats. Hence, we aimed at tackling this question by performing playback experiments (habituation–dishabituation) with three horseshoe bat species within the constant-frequency bat guild, which included big-eared horseshoe bats (Rhinolophus macrotis), Blyth’s horseshoe bats (Rhinolophus lepidus), and Chinese horseshoe bats (Rhinolophus sinicus). We studied the behavioral responses of these three species to echolocation calls of conspecifics, to other two species, and to another heterospecifics bat, Stoliczka’s trident bat (Asellisus stoliczkanus), which also belongs to this guild. We found that the three rhinolophid species displayed a series of distinct behaviors to heterospecific echolocation but few to conspecific calls after habituation, suggesting that they may have been able to discriminate sympatric heterospecific echolocation calls from those of conspecifics. Interestingly, the behavioral responses to heterospecific calls were positively correlated with the interspecific overlap index in trophic niche, whereas call design had only a minor effect. This implies that the behavioral responses of these bats to heterospecific echolocation calls may be related to the degree of interspecific food competition.     

The alarm call system of wild black-fronted titi monkeys, <Emphasis Type="Italic">Callicebus nigrifrons</Emphasis>

Upon encountering predators, many animals produce specific vocalisations that alert others and sometimes dissuade the predators from hunting. Callicebus monkeys are known for their large vocal repertoire, but little is known about the function and meaning of most call types. We recorded a large number of natural predator responses from five different groups of black-fronted titi monkeys in their Atlantic forest habitat in South Eastern Brazil. When detecting predatory threats, adult group members responded with call sequences that initially consisted of two brief, high-pitched calls with distinct frequency contours. Call A was mainly given to raptors but also to predatory capuchin monkeys and other threats within the canopy, while call B was given to predatory or non-predatory disturbances on the ground. In later parts of the sequences, we also recorded a high-pitched unmodulated call C and various low-pitched loud calls. Results therefore suggest that calls A and B provide listeners with rapid and reliable information about the general classes of danger experienced by the caller, while obtaining more specific information through other call types and combinations and behavioural responses. We discuss these findings in relation to current evolutionary theory of primate communication.     

Fighting,assessment, and frequency alteration in Blanchard's cricket frog

Summary This study examines the use of dominant frequency for assessment of fighting ability in Blanchard's cricket frog Acris crepitans blanchardi. Like most anurans, the dominant frequency of the call is negatively correlated with size. However, unlike many anurans, dominant frequency varies within individual males, providing a less reliable signal of size. Wrestling contests between male cricket frogs were differentially won by larger males. I thus tested the hypothesis that males use dominant frequency for assessment of an opponent's fighting ability. Males retreated from, or became satellites of, broadcasts of synthetic low frequency calls, simulating the calls of a large male. In contrast, males attacked broadcasts of synthetic high frequency calls, simulating the calls of a small male. The dominant frequency of the call therefore appears to contain information about size which males use to assess the fighting ability of opponents, despite the lower degree of reliability resulting from individual variation in the signal. Individual variation in dominant frequency arises partially as a consequence of active alteration of the signal in response to social competition. Males differentially lowered their dominant frequencies in response to lower frequency broadcasts, thus in response to larger opponents. Furthermore, the higher a male's dominant frequency relative to that of an opponent, the greater was the decrease in dominant frequency. These results suggest that smaller males may conditionally lower their dominant frequencies in order to sound larger to larger opponents, thereby inflating their apparent fighting ability. Two alternative hypotheses are discussed.     

Echolocation beam shape in emballonurid bats, Saccopteryx bilineata and Cormura brevirostris

The shape of the sonar beam plays a crucial role in how echolocating bats perceive their surroundings. Signal design may thus be adapted to optimize beam shape to a given context. Studies suggest that this is indeed true for vespertilionid bats, but little is known from the remaining 16 families of echolocating bats. We investigated the echolocation beam shape of two species of emballonurid bats, Cormura brevirostris and Saccopteryx bilineata, while they navigated a large outdoor flight cage on Barro Colorado Island, Panama. C. brevirostris emitted more directional signals than did S. bilineata. The difference in directionality was due to a markedly different energy distribution in the calls. C. brevirostris emitted two call types, a multiharmonic shallowly frequency-modulated call and a multiharmonic sweep, both with most energy in the fifth harmonic around 68?kHz. S. bilineata emitted only one call type, multiharmonic shallowly frequency-modulated calls with most energy in the second harmonic (~46?kHz). When comparing same harmonic number, the directionality of the calls of the two bat species was nearly identical. However, the difference in energy distribution in the calls made the signals emitted by C. brevirostris more directional overall than those emitted by S. bilineata. We hypothesize that the upward shift in frequency exhibited by C. brevirostris serves to increase directionality, in order to generate a less cluttered auditory scene. The study indicates that emballonurid bats are forced to adjust their relative harmonic energy instead of adjusting the fundamental frequency, as the vespertilionids do, presumably due to a less flexible sound production.     

Male calling behavior,female discrimination and acoustic interference in the Neotropical treefrog Hyla microcephala under realistic acoustic conditions

Summary Anuran choruses are acoustically complex assemblages of calling males. Little is known about the behavior of males or females in such natural sound environments. I studied calling behavior of males of Hyla microcephala in nature by using an interactive computer-based system that allowed me to simulate call interruptions by a number of males. I also monitored the calling behavior of groups of four to six males. When a male is interrupted by the call of another frog, he increases the spacing between the notes of his call. Responses of this kind are strongest to the loudest neighbor, and some males may ignore interruptions by all but a single close male. Interruptions using synthetic calls with silent gaps indicated that males respond vocally to reductions in sound intensity as brief as 20 ms. This ability helps to explain how males can rapidly alternate notes during pairwise interactions. Amounts of acoustic overlap between pairs of males in the choruses were usually below 10% of an individual's total calling time during bouts. The time a male spent calling that was free of acoustic interference by any other male ranged from 34–92% of his total calling time. When group size was decreased, this unobstructed calling time increased. Previous research showed that females of H. microcephala discriminate against calls that overlap so that the call pulse-train structure is degraded. Here I show that a 6 dB difference in intensity between the overlapped calls is sufficient to reduce the degradative effect of call interference. Females were also given a choice between interfering calls broadcast from two adjacent and two widely separated speakers. An angular separation between speakers of 120° was insufficient to elicit a preference for the separated sources. Together, data on behavior of males and females indicated that males actively reduce acoustic interference with those loud individuals most likely to degrade seriously the temporal structure of their calls.     

Dynamic adjustment of biosonar intensity to habitat clutter in the bat Macrophyllum macrophyllum (Phyllostomidae)

Echolocating bats adjust the time–frequency structure such as sweep rate and pulse interval of their sonar calls when they move from open space to vegetation-dense environments. Emitted call intensity is equally important for echolocation, but adjustment of signal intensity to different habitats has never been systematically studied in any bat species. To address this question, we recorded sonar calls of the Neotropical trawling insectivorous bat Macrophyllum macrophyllum (Phyllostomidae) at three sites with different obstacle densities (clutter). We found a clear correlation between emitted intensity and degree of clutter, with intensity proportional to decreasing clutter. In highly cluttered, semicluttered, and open spaces, M. macrophyllum emitted calls with mean source levels (sound pressure level (SPL) 10 cm from the bat’s mouth) of 100, 105, and 111 dB SPL root mean square (rms), respectively. To our knowledge, this is the first documentation of dynamic intensity adjustments in bats. Phyllostomid bats were previously considered silent, but the 111-dB SPL rms emitted by free-ranging M. macrophyllum in open space is comparable to output in aerial insectivorous bats from other families. Our results suggest that the acoustic constraints of habitats are better predictors of call intensity than phylogeny and therefore likely to be major drivers shaping the sonar system of bats in the course of evolution.     

Interindividual use of echolocation calls: Eavesdropping by bats

Summary The use of other individual's echolocation calls by little brown bats, Myotis lucifugus, was tested by observing the response of free-flying bats to presentations of recorded echolocation calls and artificial sounds. Bats responded by approaching conspecific calls while searching for food, night roosts, nursery colonies and mating/hibernation sites. Response was low or non-existant to other sounds. While searching for prey, M. lucifugus also responded to the echolocation calls of Eptesicus fuscus, a sympatric species with overlapping diet but distinctly different echolocation calls. Subadults were especially responsive to conspecific calls.All four situations in which the bats responded involve patchily distributed resources at which bats accumulate. Concentrations of echolocation calls thus likely serve as cues regarding the location of resources. Individuals approaching feeding groups, for example, could increase prey detection range by up to 50 times over individuals relying solely on their own echolocation.Although the costs associated with eavesdropping may be negligible for M. lucifugus, for other species, particularly territorial ones, being conspicuous may be a disadvantage and the possibility of being over-heard by other bats may have been one factor involved in the evolution of echolocation call design.     

Effects of multiple-speaker playbacks on aggressive calling behavior in the treefrog <Emphasis Type="Italic">Dendropsophus ebraccatus</Emphasis>

In addition to producing signals, males of chorusing species also act as receivers by adjusting properties of their vocalizations in response to those of other nearby individuals. Although it is likely that males are responsive to more than one other individual, most playback studies investigating male response have involved dyads in which vocal responses are measured to stimuli presented from a single speaker. In this study, I explored changes in both the propensity to give aggressive calls and the temporal properties of those calls in response to the playback of multiple aggressive call stimuli in the treefrog Dendropsophus ebraccatus. I found that males were sensitive to both the number of simulated aggressive callers and their specific call characteristics. Males generally gave a highly aggressive response to the first stimulus presented, but their response to the modification of this stimulus by the addition or subtraction of a simulated competitor depended on the degree of aggressiveness of the stimuli. Males tended to decrease their aggressive responses when either a more aggressive call was silenced or a less aggressive call was added and to increase their aggressive responses in the opposite situation. Aggressive calling in this species is clearly affected by complex changes in the social environment and I suggest that future studies explore these issues in other species to improve the understanding of communication interactions.