Acoustically mediated neighbor recognition in the North American bullfrog,Rana catesbeiana

Summary Playback experiments were conducted to determine whether territorial male bullfrogs (Rana catesbeiana) were capable of discriminating between advertisement calls of neighbors and strangers. Territorial males adjacent to subject males were removed and replaced with a speaker which broadcast calls of a stranger or the removed male. Ten of 11 males responded more strongly to calls of strangers than to calls of neighbors. When advertisement calls of neighbors were broadcast to subject males from positions within the removed neighbor's territory and from positions opposite the subject male, all 9 males tested responded more strongly to calls of a neighbor broadcast from a new position than to playbacks of the same call broadcast within the removed neighbor's territory. Because male bullfrogs recognize familiar calls and associate these calls with a particular direction or location, these data provide the first evidence for acoustically mediated neighbor recognition in frogs.     

Recognition of individual neighbors by song in the song sparrow,a species with song repertoires

Summary Previous theory and research have suggested that bird species with song repertoires in general, and song sparrows (Melospiza melodia) in particular, cannot readily discriminate between the songs of neighbors and strangers. In a recent study (Stoddard et al. 1991) we showed that song sparrows can in fact discriminate neighbors from strangers on the basis of song. In this study we sought to demonstrate that song sparrows can make the finer discrimination between individual neighbors and that they can do so on the basis of a single song type. We compared the response of territorial males to song playback of neighbors and strangers at three locations: the neighbor's regular boundary, the opposite boundary, and the center of the territory. The birds showed strong neighbor-stranger discrimination at the regular boundary but not at the opposite boundary, nor in the center of the territory. The differences in song discrimination between different boundary locations indicate that song sparrows associate particular songs with particular territories, effectively discriminating between individual neighbors on the basis of song. Song repertoires themselves do not interfere with neighbor recognition to the extent originally postulated. As speakers are moved inside the territory from the border, however, the degree of discrimination diminishes. We believe that differences in speaker placement may have contributed to the variability in neighbor-stranger discrimination observed in previous studies of the song sparrow and perhaps other repertoire species as well. This interpretation is consistent with data from another song sparrow population showing that half the territory takeovers are by immediate neighbors.     

Individual recognition of neighbors by song in a suboscine bird, the alder flycatcher Empidonax alnorum

The ability of territorial males to discriminate between songs of their neighbors and songs of strangers has been demonstrated in 27 species of songbirds. Such experiments test only the ability of a subject to discriminate between two classes of stimuli, familiar (neighbors) and unfamiliar (strangers) songs. Individual recognition of neighbors is a finer, more complex type of discrimination. The ability of territorial males to recognize individual neighbors by song has been documented in 12 species of oscine passerines (Passeriformes, Passeri), but has never been demonstrated in suboscine passerines (Tyranni). We investigated recognition of songs of individual neighbors in a suboscine, the alder flycatcher (Empidonax alnorum). We performed a series of song playback experiments and recorded responses of territorial males to songs of neighbors and songs of strangers broadcast from two locations, the neighbor boundary and an opposite boundary. Subjects responded more aggressively to songs of a neighbor when played from the opposite boundary than when played from the neighbor boundary. They responded with equal aggression to songs of strangers regardless of location of playback. The difference in response to neighbor songs between speaker locations and the lack of a difference in response to stranger songs indicate that territorial males associate a particular song with a particular location (territory), and thus recognize individual neighbors.Communicated by I. Hartley     

Are vocal signals used to recognize individuals during male–male competition in greater prairie-chickens (Tympanuchus cupido)?

Vocal signaling can be an important component of vertebrate communication during social interactions. If vocalizations vary among individuals but are consistent within a given individual, they may be used to discriminate among individuals. In many species, territorial males use vocalizations to discriminate between neighbors and strangers and either respond more aggressively toward strangers relative to neighbors (“dear enemy” effect) or they respond more aggressively toward neighbors relative to strangers (“nasty neighbor” effect). In the greater prairie-chicken (Tympanuchus cupido), male vocalizations are an integral part of the display males produce on leks. We investigated whether male greater prairie-chickens discriminate among familiar individuals on their own territory, familiar individuals outside their normal territory and strangers from a nearby lek. Vocal characteristics varied among males, suggesting that vocalizations may potentially be used by prairie-chickens to identify individuals. Males responded to playback of prairie-chicken calls by vocalizing at a faster rate and approaching the playback speaker, but did not vary in their response to the vocalizations based on the identity of the caller. Our results suggest that variation is present among the vocalizations of individual male greater prairie-chickens, but males do not appear to discriminate among familiar individuals and strangers based solely on their “boom” vocalizations. Greater prairie-chicken vocalization likely functions as a way of announcing that a territory is occupied and defended, but it may also serve as a way of advertising to conspecifics or as a signal that is secondary to other forms of communication.     

Neighbor–stranger discrimination in Audubon's shearwater (Puffinus l. lherminieri) explained by a “real enemy” effect

Neighbor–stranger discrimination (NSD) occurs when animals respond with more aggression to strangers than to territorial neighbors. NSD has been reported in many species that defend multi-purpose territories for breeding and foraging, but it is rare among species that defend other types of territories. For birds that defend only their nest sites, there is no experimental evidence for NSD, and observational studies have provided mixed results. In a colony of Audubon's shearwaters (Puffinus l. lherminieri), I played back the calls of a neighbor and a stranger to males defending nest sites. Subjects responded with longer calls to playbacks of strangers than to those of neighbors. In shearwater colonies, strangers are often birds looking for future breeding sites. In contrast, there is no evidence that established breeders compete with their neighbors for any resources. Shearwaters should benefit from NSD because strangers represent a “real enemy” and established neighbors do not.     

Social monitoring in a multilevel society: a playback study with male Guinea baboons

Keeping track of social interactions among conspecifics is a driving force for the evolution of social cognition. How social cognition, such as social knowledge, ties in with a species' social organization is, however, largely unexplored. We investigated the social knowledge of wild Guinea baboons (Papio papio) ranging in Senegal, a species that lives in a fluid multilevel society with overlapping habitat use. Using playback experiments, we tested how adult males differentiate between subjects from their own vs. a neighboring or a stranger social unit (“gang”) and assessed ranging patterns with Global Positioning System (GPS) data. While territorial species usually differentiate between group and nongroup members and often respond more strongly to strangers than neighbors (the “dear enemy” effect), subjects in this highly tolerant species should largely ignore other unit members and mainly attend to subjects from their own unit. Males responded strongly after playback of calls recorded from members of their own gang, while they attended only briefly to neighbor or stranger calls. Apparently, males benefit from monitoring the social maneuvers in their own social unit, while it remains to be resolved whether they are unmotivated or unable to keep track of the identities and actions of individuals outside their own gang. The study highlights how the allocation of social attention is tuned to the specifics of a species' social organization, while a complex social organization does not necessarily translate into the need for more elaborate social knowledge.     

Playback and speaker-replacement experiments on song-based neighbor,stranger, and self discrimination in male Red-winged Blackbirds

Summary Two experiments were conducted to deterine whether territorial male Red-winged Blackbirds discriminate among neighbor, stranger, and self songs. In the first experiment, recordings of the three classes of songs were played to territorial males. Responses to playback of stranger song, as measured by song rate, intensity of Song Spread display, and closest approach, were significantly stronger than to playback of neighbor song. In addition, stranger song elicited significantly more intense Song Spread displays than did self song.In the second experiment, territorial males were removed from their territories and each was replaced with speakers broadcasting recorded songs of a stranger, and of the removed male. Broadcasts of the removed male's own songs were more effective in discouraging trespassing by neighboring male redwings than were broadcasts of songs of a stranger. However, the two broadcasts did not differ in their ability to deter nonneighboring male redwings from flying through or trespassing within the speaker territory.We suggest that neighbor-stranger discrimination in the Red-winged Blackbird develops as a consequence of associative learing.     

Song repertoires do not hinder neighbor-stranger discrimination

Summary Through playback experiments and a comparative study we tested the hypothesis that song repertoires hinder individual recognition by song in birds. Our playback tests take advantage of the peculiar singing behavior of two species of parulid warblers: the American redstart (Setophaga ruticilla) and the yellow warbler (Dendroica petechia). Males of both species sing either the same song repeatedly (repeat mode) or a repertoire of songs in series (serial mode). Subjects were played the repeat song and the serial songs from a territorial neighbor and from a stranger some territories distant. Counter to the prediction of the hypothesis, they discriminated equally well between neighbors and strangers when the several serial songs were used as when the single repeat song was played. Thus, neither the small repertoires used by redstarts in serial mode (2–8 songs) nor the larger ones used by yellows (an average of over 20 songs) appear to hinder neighbor-stranger discrimination. In our comparative test, we examined the relationship between neighbor-stranger discrimination and repertoire size among 20 species for which data were available. Using two comparative methods, we did not find the predicted significant negative relationship.     

The red and the black: habituation and the dear-enemy phenomenon in two desert Pheidole ants

Many species of territorial animals are more aggressive toward strangers than neighbors, a pattern of aggression referred to as the ’dear-enemy phenomenon.’ In many cases, the mechanism by which neighbors are discriminated from strangers and the function of neighbor-stranger discrimination remain controversial. We investigated the spatial patterns of inter-colony aggression within and between two Pheidole species of seed-harvesting ants in the Mojave Desert of California by quantifying aggression between colonies in standardized staged encounters. We also tested whether the level of fighting between workers of two colonies is affected by previous exposure to each other. We show that neighbors (i.e., colonies less that 2.6 m away) of either species are treated less aggressively than more distant colonies and that habituation may be a mechanism by which this discrimination is achieved. The variation in aggression among spatially distant colonies also suggests that additional genetic or environmental factors are involved in recognition. The function of the dear-enemy phenomenon in these ant species may be related to the greater risk to the resources of a colony presented by strange workers than workers from a neighboring colony. Received: 18 November 1999 / Received in revised form: 3 April 2000 / Accepted: 3 May 2000     

Territory ownership and familiarity status affect how much male root voles (Microtus oeconomus) invest in territory defence

Neighbour–stranger discrimination occurs when individuals respond with more aggression to strangers than to territorial neighbours—a phenomenon termed the “dear enemy phenomenon” (DEP). We investigated the DEP with male and female root voles (Microtus oeconomus Pallas 1776) using field dyadic arena tests conducted in enclosures where we could test for the effects of familiarity (familiar versus stranger), ownership (resident versus intruder status) and resource-holding potential (body mass) on territorial behaviours. The results showed that males put more effort into territorial defence than females, and males could discriminate between neighbours and strangers. In males, aggressiveness was influenced by a significant two-way interaction between treatment and ownership. Male residents were more aggressive towards stranger intruders than towards neighbour intruders, while male intruders were less aggressive towards stranger residents than towards neighbour residents. In females, neither treatment nor ownership status had a significant effect on aggressiveness. Familiar males performed more social behaviours but less non-social behaviours than stranger males. Furthermore, there was a clear dominance hierarchy between residents and intruders in stranger dyads, with the male territory holders dominating the intruder in pairwise interactions. To our knowledge, these results demonstrate for the first time DEP in a small mammal with a known pedigree and present the first evidence for “prior resident advantage” in voles. We argue that both ownership status and familiarity status affect how much an individual invests in territory defence. The benefits of neighbour–stranger discrimination for male root voles and the absence of neighbour–stranger discrimination in female root voles are discussed.     

Habituation in the green frog, Rana clamitans

Habituation and its role in the dear enemy effect was investigated in a population of green frogs, Rana clamitans. Green frogs have a prolonged breeding season, and males defend territories centered around suitable oviposition sites. We tested the prediction that male green frogs will habituate to broadcasts of synthetic conspecific stimuli. Our results indicate that male green frogs can discriminate familiar from unfamiliar stimuli. We suggest that habituation helps to mediate the territorial interactions between male green frogs. Strangers present a greater threat than familiar neighbors. By habituating to the advertisement vocalizations of their near neighbors, males avoid costly interactions with individuals that are not a major threat to their territories. Received: 20 July 1998 / Accepted after revision: 6 September 1998     

Tit for tat among neighboring hooded warblers

Summary The dear-enemy relationship of territorial songbirds could be mutually beneficial to neighbors, as males who recognize neighbors and reduce their responses to these neighbors would require less time and energy for territorial defense. In order for this relationship to be evolutionarily stable, this reduction in response to a neighbor must be conditional on reciprocal restraint by that neighbor. This study examined the possibility of such conditional responses in hooded warblers (Wilsonia citrina). Responses of territorial hooded warblers to playbacks of neighbors' songs from shared boundaries were measured before and after playbacks that simulated intrusions of those same neighbors (NNNN treatment) or strange birds (NSSN treatment) into the subjects' territories. Each male received both treatments separated by at least 8 days. Males increased their responses to playbacks of a neighbor's songs at the boundary after simulated intrusions of that same neighbor (NNNN) but did not increase their responses to such playbacks after simulated intrusions of strangers (NSSN). This increased response to a defecting neighbor suggests that the relationship between neighboring territorial hooded warblers is based on a conditional strategy like tit-for-tat.     

When to be neighbourly: differential agonistic responses in the lizard Platysaurus broadleyi

Differential treatment of neighbours and strangers (the dear enemy phenomenon) can reduce the costly effects of territory defence. Individual recognition, and by extension the dear enemy phenomenon, was examined in the cordylid lizard Platysaurus broadleyi. I had no knowledge of familiarity between individuals and therefore classed intruders as neighbours or non-neighbours and tested for differences in agonistic response consistent with the dear enemy phenomenon. In the dear enemy phenomenon, levels of aggression are inversely related to degree of familiarity such that residents are more tolerant of immediate neighbours than distant territory holders. A manipulative field experiment in conjunction with field observations of known territory holders revealed that resident males allowed neighbours to approach more closely than non-neighbours before challenging them, residents were more aggressive towards non-neighbours than neighbours, contests between neighbours were significantly longer than between neighbours and non-neighbours and contests between residents and non-neighbours were likely to result in a win for the resident, while neighbours frequently drew contests. These results suggest a level of recognition consistent with the dear enemy phenomenon. I also manipulated front leg colour to test the hypothesis that it alone could serve as a cue for individual recognition. I found no support for this hypothesis and suggest that multiple cues operate in individual recognition. Received: 4 March 1999 / Received in revised form: 11 April 1999 / Accepted: 19 April 1999     

Northern harriers on feeding territories respond more aggressively to neighbors than to floaters

Summary Territory owners often respond less aggressively towards intruding neighbors than towards intruding floaters, an observation termed the dear enemy phenomenon. Comparisons of territory owners' responses to intruding neighbors versus their responses to intruding floaters usually have been made for owners of multi-purpose and/or breeding territories. Here, I describe responses of female northern harriers Circus cyaneus (owners) on winter feeding territories towards three types of intruders (female neighbors, female floaters, and male floaters) and show that the dear enemy phenomenon does not occur. Owners' responses towards neighbors were more intense (mostly flights rather than calls) than responses towards female floaters, which in turn were more intense than responses towards male floaters. The greater intensity of owners' responses towards neighbors compared to owners' responses towards male and female floaters may be related to differences in the threat posed by each of the three intruder types in terms of fighting ability (RHP) and potential losses from intrusion. Hence, whether owners respond more aggressively towards neighbors or floaters, and whether the dear enemy phenomenon is observed, may depend upon the relative magnitude of threat presented by neighbors and floaters to owners in terms of fighting ability and potential losses from intrusion.     

Effects of kinship on territorial conflicts among groups of lions,<Emphasis Type="Italic"> Panthera leo</Emphasis>

Inclusive fitness theory predicts that cost of tolerant behaviour during competitive interactions is lower for relatives than for nonrelatives. Many studies have examined the effect of relatedness on behaviour within social groups. In contrast, kin selection acting among groups has received less attention. The genetic structure of African lion (Panthera leo) populations creates a strong possibility that kin selection among groups modifies behaviour during group conflicts. We used playback experiments and genetic data to investigate the importance of relatedness during simulated territorial disputes in lions. However, we found no effect of relatedness on territorial behaviour. Degree of relatedness did not affect the decision to approach simulated intrusions, nor did it affect the behaviour during approaches. The decision to approach was instead affected by position within the territory and consecutive playback number (a measure of habituation). For playbacks that did elicit an approach, the speed of response was not detectably affected by relatedness, but was affected by odds (the ratio of residents to intruders), number of intruders, number of bouts, presence of cubs, position within the territory, temperature and playback number. Although responses were unaffected by relatedness, it remains possible that other aspects of behaviour during natural encounters among prides are affected by kin selection.Communicated by L. Sterck     

Neighbours, strangers and male-male aggression as a determinant of lek size

Interactions between males on leks may play an influential role in lek formation and the regulation of lek size. In this paper I present the results of a playback experiment that simulated de novo settlement at sites adjacent to currently existing display territories of the ochre-bellied flycatcher, Mionectes oleagineus. In the study population, males displayed both solitarily and at small leks. A large proportion of males held no display territory at all. A stranger's song was played to both solitary and lekking males from 10 m outside their territorial boundaries. In separate playbacks, lekking males were also played neighbour's song. Both lekking and solitary territorial males reacted to the playback by decreasing their song rate, approaching the playback speaker and, on occasion, attacking the model. Solitarily displaying males responded more aggressively to playback of stranger's song than did lek males. Lek males were able to distinguish between their neighbour's and a stranger's song and did so irrespective of whether it was played from the neighbour's territory or from outside the lek. In addition to distinguishing between neighbours and strangers, lek males modified their responses to these different playbacks depending on where the playback originated. These results suggest that male-male interactions can be influential in structuring leks. In M. oleagineus, interactions between males are aggressive and act to limit rather than augment lek size. Received: 6 March 1996 / Accepted after revision: 9 December 1996     

Territorial counter-singing in male sac-winged bats (<Emphasis Type="Italic">Saccopteryx bilineata</Emphasis>): low-frequency songs trigger a stronger response

In many animal species, individuals compete for resources but avoid escalated conflicts by threat displays, i.e. a mutual signalling behaviour that enables the opponents to predict the outcome of the conflict without the necessity of actual fighting. For example, territory holders may use acoustic signals to communicate not only their own identity and the borders of their territory but also their competitive quality, fighting ability and motivation. Here, we show that male sac-winged bats, Saccopteryx bilineata, adjust their vocal territorial displays according to the fundamental frequency of territorial songs of their opponents. In playback experiments with territorial males, low-frequency stimuli elicited a higher territorial song rate and length than high-frequency stimuli. Male S. bilineata that sing more often and with lower fundamental frequencies have been shown to sire more offspring than their competitors. Fundamental frequency of territorial songs, hence, may reveal male quality and, consequently, the resulting threat posed to competing males. We argue that this is reflected in the increased response of competitors to low-frequency territorial songs shown here. Such competitive signalling behaviour has been shown in a few mammal species like red deer and baboons but, thus far, not in bats. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Territorial vocal behavior in hybrid smooth froglets, Geocrinia laevis complex (Anura: Myobatrachidae)

Males of the parapatically distributed myobatrachid frogs Geocrinia laevis and G. victoriana have highly divergent advertisement calls. Furthermore, the two species differ strongly in the complexity of their vocal repertoires, with males of G. victoriana possessing, and those of G. laevis lacking, a distinct territorial vocalization (encounter call). We investigated the territorial vocal behavior of males in a persistent natural hybrid population. Most hybrid males possessed encounter calls functionally equivalent to those of G. victoriana, that were produced following exposure to playback of recorded advertisement calls presented at >110 dB peak sound pressure level. The territorial acoustic responses were not associated with an index of hybridity derived from the structure of the advertisement call, suggesting genetic and functional decoupling of the two components of the vocal repertoire; i.e., advertisement calls and encounter calls. This decoupling may be the result of sexual selection favouring those hybrids with pronounced territorial behavior and the associated vocalization, regardless of the structure of their advertisement calls.Communicated by A. Mathis     

Advertisement-call modification,male competition,and female preference in the bird-voiced treefrog <Emphasis Type="Italic">Hyla avivoca</Emphasis>

Senders and receivers influence dynamic characteristics of the signals used for mate attraction over different time scales. On a moment-to-moment basis, interactions among senders competing for a mate influence dynamic characteristics, whereas the preferences of receivers of the opposite gender exert an influence over evolutionary time. We observed and recorded the calling patterns of the bird-voiced treefrog Hyla avivoca to assess how the dynamic characters of calls vary during interactions among groups of males in a chorus. This question was also addressed using playback experiments with males. Playback experiments with females showed how changes in dynamic call properties are likely to affect male mating success. Frogs calling in pairs, groups, or in response to playbacks produced longer calls than did isolated males. During call overlap, males often increased the duration of the silent interval (gaps) between the pulses of their calls so that the pulses of the calls of two neighbors interdigitated. This change resulted in increased variability of pulse rate, a traditionally static acoustic property; however, males also produced high proportions of non-overlapped calls in which variability in pulse rate was low and had species-typical values. Females preferred long calls to short- and average-duration calls, and non-overlapped calls to overlapped calls. Given a choice between pairs of overlapped calls, females preferred pairs in which the proportion of overlap was low and pairs in which the pulses of such calls interdigitated completely. The observed patterns of vocal competition thus reflect the preferences of conspecific females, which have influenced the evolution of the calling behavior of H. avivoca. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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.