Function of pair duets in the eastern whipbird: cooperative defense or sexual conflict?

Paired male and female eastern whipbirds, Psophodes olivaceus,sing precisely coordinated, male-led duets. Four broad explanationshave been proposed for the function of duets: 1) cooperativeresource defense, 2) prevention of partner usurpation, 3) defenseof an individual's own position within the partnership, or 4)mate identification and localization. These 4 hypotheses makedifferent predictions about how male and female residents shouldrespond to simulated intrusion by other pairs or individuals.We compared the behavioral and vocal responses of 20 pairs ofeastern whipbirds to simulated territorial intrusions by: 1)a solitary singing male, 2) a solitary singing female, and 3)a duetting pair. Males and females did not coordinate theirapproach to the playback speaker and showed sex-specific responsesto playback. Males did not respond differently to duetting versussolo singing intruders. By contrast, females approached moreclosely during solo female song than during solo male song orduet playback. Females also produced specific vocalizationsonly in response to duet and solo female playback. Both sexesapproached the speaker more closely and quickly during playbackof same-sex solo songs than opposite-sex solo songs. Finally,females answered more of their mate's songs during simulatedintrusion by a lone female than during simulated intrusion bya lone male. Our results suggest that duets in this speciesprimarily function to allow females to defend their exclusiveposition in a partnership. Mate defense by females is unusualin birds but may be promoted in eastern whipbirds by a female-biasedsex ratio and the need for exclusive access to male care. Thus,duets result from independent and conflicting strategies ofmate and territory defense in males and females.     

The function of duetting in magpie-larks: conflict, cooperation, or commitment?

Avian duetting is a poorly understood phenomenon despite many hypotheses as to its function. Contrary to the recent view that duetting functions for mate guarding and is a result of conflict between the sexes, Australian magpie-larks, Grallina cyanoleuca, do not use duetting as a paternity guard. I used a playback experiment to investigate the role of antiphonal duetting in territorial defence and pair bond maintenance, two traditional hypotheses about the function of duetting. The experiment showed that, like many nonduetting species, magpie-larks recognize neighbours on the basis of song. It also provided evidence of functional differences between duetting and solo singing which indicate that temporal coordination of song between partners is used to maintain the territory and pair bond. Duets were more threatening territorial signals than solo songs: males initiated more vocalizations in response to playback of duets than playback of solos. Simulated intrusion also caused males and females to approach the speaker together and coordinate more of their vocalizations to form duets. Females did not engage in sex-specific territorial defence, responding equally strongly to playback of male and female song, and maintaining both territory and pair bond by attempting to exclude intruders of either sex. Males initiated more vocalizations in response to playback of male than female song, and their likelihood of duetting appeared to be related more to threats to the pair bond, in particular desertion by their partner. Copyright 2000 The Association for the Study of Animal Behaviour.     

Song Functions in Nonduetting Gibbons: Evidence from Playback Experiments on Javan Gibbons (<Emphasis Type="Italic">Hylobates moloch</Emphasis>)

Territorial, pair-living primates usually perform long-distance calls as duets in which adult males and females coordinate their calls. Previous studies using playback experiments have shown that gibbon duets convey information about the status of the caller (location, familiarity, sex of the caller, and paired status) and gibbons use this information to respond to achieve several nonmutually exclusive functions, including intragroup contact, territorial defense, and pair-bond advertisement and strengthening. However, not all pair-living gibbons duet, and it is unclear whether the same results should be expected in nonduetting species. We conducted song playback experiments (N = 47 trials) to test hypotheses about song functions in nonduetting gibbons on two groups of wild Javan gibbons (Hylobates moloch) in the Gunung Halimun-Salak National Park, Indonesia. Javan gibbons initiated movement toward the speaker more quickly in response to songs broadcast in the center of the territory, stranger songs, and songs of unpaired individuals than to songs at the border, neighbor songs, and songs from paired individuals. These results suggest that Javan gibbons can localize songs, and that Javan gibbon songs transmit information about the identity and paired status of the caller. Our results imply that Javan gibbon solo songs are likely to function for territorial defense and pair-bond advertisement like duets in other primates.     

Duets defend mates in a suboscine passerine, the warbling antbird (Hypocnemis cantator)

Despite the widespread occurrence of avian duets, their adaptivesignificance is poorly understood. It is generally assumed thatthey function in the joint defense of territories, but no studyhas successfully distinguished between this hypothesis, whichinvokes cooperation between the sexes, and mate defense, whichinvokes conflict. Further, most duetting studies have focusedon oscine passerines, the songs of which are learnt and relativelycomplex. We therefore tested the mate defense hypothesis inthe warbling antbird (Hypocnemis cantator), an Amazonian suboscinethat produces simple sex-specific songs and duets. Acousticanalysis of songs showed (1) that solos were often producedby males, but rarely by females; (2) that duets consisted ofa male song and a female reply; and (3) that, although femalesong was invariable, a swift reply resulted in males producingshorter songs with fewer notes. These results suggest that duetting,and the structure of duets, is chiefly a product of female behavior,a scenario more suggestive of conflict than cooperation. Toinvestigate this idea we carried out playback experiments, whichshowed that (4) the response to solo songs was sex specific(i.e., male solos elicited a strong response from paired males,and female solos elicited a strong response from paired females);(5) males and females responded to same-sex solos more stronglythan to duets; and that (6) females answered their partner'ssongs more often, and more rapidly, in response to female solosthan male solos or duets. Although it can be argued that sex-specificresponses to solo song result from intrasexual territorial defense,we cannot use the same reasoning to explain (5) or (6). Instead,these observations imply that solitary intruders were more threateningthan paired intruders, and thus that the perceived threat wasto the partnership rather than the territory. Taken together,findings (1) to (6) suggest that females adjust their vocalbehavior in relation to the level of perceived threat to thepartnership, and duet with males in order to repel same-sexrivals. This study therefore strengthens support for the matedefense hypothesis, and suggests that conflict—ratherthan cooperation—may have played a major role in the evolutionand maintenance of avian duets.     

Duet function in the yellow-naped amazon, Amazona auropalliata: evidence from playbacks of duets and solos

The question of why animals participate in duets is an intriguing one, as many such displays appear to be more costly to produce than individual signals. Mated pairs of yellow-naped amazons, Amazona auropalliata, give duets on their nesting territories. We investigated the function of those duets with a playback experiment. We tested two hypotheses for the function of those duets: the joint territory defense hypothesis and the mate-guarding hypothesis, by presenting territorial pairs with three types of playback treatments: duets, male solos, and female solos. The joint territory defense hypothesis suggests that individuals engage in duets because they appear more threatening than solos and are thus more effective for the establishment, maintenance and/or defense of territories. It predicts that pairs will be coordinated in their response (pair members approach speakers and vocalize together) and will either respond more strongly (more calls and/or more movement) to duet treatments than to solo treatments, or respond equally to all treatments. Alternatively, the mate-guarding hypothesis suggests that individuals participate in duets because they allow them to acoustically guard their mate, and predicts uncoordinated responses by pairs, with weak responses to duet treatments and stronger responses by individuals to solos produced by the same sex. Yellow-naped amazon pairs responded to all treatments in an equivalently aggressive and coordinated manner by rapidly approaching speakers and vocalizing more. These responses generally support the joint territory defense hypothesis and further suggest that all intruders are viewed as a threat by resident pairs.     

Functions of duetting in the tropical boubou, Laniarius aethiopicus: territorial defence and mutual mate guarding

Duetting, found in many animal taxa, is still a poorly understood form of signalling behaviour despite numerous hypotheses to explain its function. One contentious issue is whether duetting is a cooperative endeavour or signals conflict between the sexes. We studied the function of duetting in the tropical boubou, using interactive playback experiments. We staged encounters by presenting paired birds with four variants of each of four duet types (neighbour and stranger solos and neighbour and stranger duets). Multiple lines of evidence suggest that both joint territorial defence and mutual mate guarding are important functions of duetting, and that both cooperation and conflict between males and females have shaped duetting behaviour. Of a repertoire of 12 duet types, eight were used for joint territorial defence. Two of these eight duet types were probably also used for mutual mate guarding, suggesting that individual duets can have multiple functions. We found an unusual response for duetting birds in that females often synchronized notes with male solo playback to form precise duets. In turn, males attempted to jam the song of rivals when females joined the song of male solo playback, providing further evidence for the importance of acoustic mate guarding. Song jamming by males has not previously been described for duetting species. Finally, we suggest that the repertoire of duet types is used to fine-tune territorial encounters in a network environment of multiple interactants.     

Sex Differences in Territorial Behavior Exhibited by the Spotted Hyena (Hyaenidae, Crocuta crocuta)

Spotted hyenas ( Crocuta crocuta ) are gregarious carnivores that defend group territories against encroachment by neighboring conspecifics. Here we monitored the behavior of members of one clan of free-ranging spotted hyenas during border patrols, 'wars' with neighboring clans, and other interactions with alien intruders, to document differences between the sexes in territorial behavior in this species. We also examined the possibility that the probability or rate of attack on alien hyenas encountered within the clan's territory would vary with the sex of the intruders. Initiation and leadership of most cooperative territorial behaviors were by adult female clan members, although border patrols were occasionally conducted by groups composed exclusively of resident males. The vast majority of alien intruders into the territory of the study clan were males. Resident females were more likely to attack intruding females than intruding males, but hourly rates of aggression directed by females towards aliens did not vary with intruder sex. Resident males were more likely than resident females to attack alien males, and resident males directed significantly higher hourly rates of aggression towards intruding males than females. Although female leadership in most cooperative territorial behaviors distinguishes spotted hyenas from many mammalian carnivores, other sex differences in the territorial behavior of spotted hyenas resemble those documented in other gregarious predators. Sex differences observed in hyena territoriality are consistent with the hypothesis that male and female clan members derive different selective benefits from advertisement and defense of group territories.     

Acoustic Monitoring Reveals Congruent Patterns of Territorial Singing Behaviour in Male and Female Tropical Wrens

Many animals defend territories against conspecific individuals using acoustic signals. In birds, male vocalizations are known to play a critical role in territory defence. Territorial acoustic signals in females have been poorly studied, perhaps because female song is uncommon in north‐temperate ecosystems. In this study, we compare male vs. female territorial singing behaviour in Neotropical rufous‐and‐white wrens Thryothorus rufalbus, a species where both sexes produce solo songs and often coordinate their songs in vocal duets. We recorded free‐living birds in Costa Rica using an eight‐microphone Acoustic Location System capable of passively triangulating the position of animals based on their vocalizations. We recorded 17 pairs of birds for 2–4 consecutive mornings and calculated the territory of each individual as a 95% fixed kernel estimate around their song posts. We compared territories calculated around male vs. female song posts, including separate analyses of solo vs. duet song posts. These spatial analyses of singing behaviour reveal that males and females use similarly sized territories with more than 60% overlap between breeding partners. Territories calculated based on solo vs. duet song posts were of similar size and similar degrees of overlap. Solos and duets were performed at similar distances from the nest for both sexes. Overall, male and female rufous‐and‐white wrens exhibit very similar spatial territorial singing behaviour, demonstrating congruent patterns of male and female territoriality.     

Partners coordinate territorial defense against simulated intruders in a duetting ovenbird

Duets in breeding pairs may reflect a situation of conflict, whereby an individual answers its partner's song as a form of unilateral acoustic mate guarding or, alternatively, it may reflect cooperation, when individuals share in territory defense or safeguard the partnership. The degree of coordination between the sexes when responding to solo versus paired intruders may elucidate the function of songs in duets. We examined this issue in a study with rufous horneros (Furnarius rufus), a duetting, socially monogamous Neotropical species with low levels of extrapair paternity. We exposed social pairs during the nonbreeding season to playbacks of duets, male solos, female solos, and control heterospecific songs. Partners approached all conspecific stimuli together and responded by singing quickly, at higher rates and by coordinating ~80% of their songs into duets. For both sexes, most response variables (seven of nine) did not vary across conspecific treatments. These results suggest that partners duet and coordinate behaviors to cooperatively defend common territories. However, females spent more time in territorial vigilance, and partners were highly coordinated (correlated responses) in response to duets and female solos in comparison with male solos. This indicates that female intrusions (paired or solo) might be more threatening than male intrusions in the nonbreeding season, especially for territorial females, and that females are less cooperative with their partners in territory defense against male intruders.     

The Role of Female Behavior in Maintaining Monogamy of a Coral-Reef Filefish

The corallivorous filefish Oxymonacanthus longirostris is a predominantly monogamous species without parental care. Polygyny reduces female fitness because monogamous females benefit from male assistance with feeding–territory defense that secures both a feeding area and time to feed. I studied the strategy of females to prevent polygyny, with the help of field observations and manipulative field experiments. Monogamous females always behaved aggressively towards intruding floater females. When a caged male or female was placed within the territory of a monogamous pair, the resident female was more likely to pay attention to the caged female than to the caged male. Experimentally transplanted floater females succeeded in settling as secondary females in the monogamous territory more frequently when the resident female was caged. These results suggest that monogamous females behave aggressively towards floater females to defend their mating status; such aggressive behavior is successful in preventing polygyny. Furthermore, the removal experiment showed that polygynous (especially secondary) females moved to territories of widowed males located near them more frequently than monogamous females did. This outcome indicates that polygynous females can change their mating status and escape polygyny by moving to another territory if the opportunity occurs. Thus, female behavior can play an important role in the maintenance of monogamy and in lowering the level of polygyny within populations of this species.     

Experimental test of female black howler monkey (Alouatta Pigra) responses to loud calls from potentially infanticidal males: effects of numeric odds, vulnerable offspring, and companion behavior

During group defense, the contribution of female black howler monkeys (Alouatta pigra) may help deter male intruders; however, their involvement during natural intergroup encounters is facultative. Experimental playback trials simulating potentially infanticidal males were used to examine whether a female's reproductive investment and/or her group's relative fighting ability would influence her participation in loud call displays. Female howlers never responded to recordings without alpha male accompaniment, but their response patterns did not simply mimic his. For example, unlike alpha males, females with small, vulnerable offspring were no more likely than females without infants to participate in howling displays during playback trials. Further, although females without any small infants in their group got closer to speakers than females living in groups with small infants, males did not respond in the same way. To artificially simulate different "numeric odds" scenarios, recordings of one or three howling males (simulated intruders) were broadcast to females living in groups with 1-3 resident males. Responses were consistent with the hypothesis that females assessed intergroup fighting ability. As in alpha males, the weakest female responses occurred when the numeric odds were against their group. However, whereas alpha males participated most when numeric odds were in their favor, females had the most intense approach responses when the number of defending and intruding males was equal. Females appeared to use a cost-effective strategy, reserving their assistance for when their participation could have the greatest impact.     

Acoustic Competition in Male Midwife Toads Alytes obstetricans and Alytes cisternasii: Response to Neighbor Size and Calling Rate. Implications for Female Choice

Effects of neighbor on male calling behavior was studied through playback experiments of synthetic calls to males of two species of midwife toads. The responses of resident males were scored considering two temporal parameters (call duration and calling rate) and one spectral parameter (dominant frequency). The sounds used for the playback tests included two levels of fundamental frequency (correlated with male size) and two levels of call repetition rate. In both species, resident males only changed their calling rate in the presence of an intruder, and the response was different for synthetic calls with two levels of dominant frequencies and with two calling rates. Resident size was not significantly correlated with the magnitude of the change in the calling rate. On the other hand, resident calling rate was significantly and positively correlated with the magnitude of the increase in calling rate of the stimulus. The maximum relative increase in calling rate was observed in A. cisternasii. In phonotaxis tests, females are preferentially attracted to calls emitted at a higher rate confirming the importance of changes in calling rate for female attraction.     

Male and female rufous‐and‐white wrens do not match song types with same‐sex rivals during simulated territorial intrusions

In birds with song repertoires, song‐type matching occurs when an individual responds to another individual's song by producing the same song type. Song‐type matching has been described in multiple bird species and a growing body of evidence suggests that song‐type matching may serve as a conventional signal of aggression, particularly in male birds in the temperate zone. Few studies have investigated song‐type matching in tropical birds or female birds, in spite of the fact that avian biodiversity is highest in the tropics, that female song is widespread in the tropics, and that female song is the ancestral state among songbirds. In this study of rufous‐and‐white wrens Thryophilus rufalbus, a resident neotropical songbird where both sexes sing, we presented territorial males and females with playback that simulated a territorial rival producing shared and unshared songs. In response, both males and females sang matched song types at levels statistically equal to levels expected by chance. Furthermore, males and females exhibited similar levels of aggression and similar vocal behaviours in response to playback of both shared and unshared songs. These results indicate that rufous‐and‐white wrens do not use song‐type matching in territorial conflicts as a conventional signal of aggression. We discuss alternative hypotheses for the function of song‐type sharing in tropical birds. In particular, we point out that shared songs may play an important role in intra‐pair communication, especially for birds where males and females combine their songs in vocal duets, and this may supersede the function of song‐type matching in some tropical birds.     

The Responses of Duetting Antbirds to Stereo Duet Playback Provide Support for the Joint Territory Defence Hypothesis

In multiple animal taxa, including many birds and primates, members of mated pairs produce coordinated acoustic displays known as duets. By observing the behaviour of territorial animals as they respond to playback‐simulated duets of rivals, we can gain insight into the behavioural significance of vocal duets. Playback experiments, however, have been conducted across a very narrow range of duetting animals. Furthermore, many studies have been conducted with single‐speaker playback, whereas stereo‐speaker playback offers more spatially realistic simulation of duets. Moreover, by evaluating the reactions of animals to separate loudspeakers broadcasting male and female duet contributions, we can study the interactions of both males and females with same‐sex vs. opposite‐sex rivals. We used a paired experimental design to broadcast duet stimuli through a single‐speaker and a stereo‐speaker apparatus to 30 pairs of duetting barred antshrikes Thamnophilus doliatus in Costa Rica. Our goals were (1) to evaluate whether territorial antbirds respond more aggressively to male vs. female duet components and (2) to assess aggressive responses of antbirds towards single‐speaker vs. stereo‐speaker playback. Neither males nor females differentiated between the loudspeaker simulating the male vs. female duet contribution during stereo‐speaker playback trials. Barred antshrikes displayed significantly stronger responses to stereo‐speaker playback compared with single‐speaker playback. Males displayed stronger playback responses than females with closer, quicker and more vocal responses. These results provide evidence for a joint resource defence function of antbird duets given that pairs responded together with equivalent intensity to male and female simulated intruders. This is the first study to show that although duetting is an aggressive territorial signal, birds do not necessarily respond to sex‐specific components of duets. Our results support the idea that spatially realistic stereo presentation of duet stimuli is critical for experimental duet research.     

Characteristics of the Signals of Male <Emphasis Type="Italic">Anoeconeossa bundoorensis</Emphasis> Taylor and Burckhardt (Hemiptera: Aphalaridae) Associated with Female Responsiveness

Sexual selection requires intra-specific variation in the characteristics mediating mate choice. In species reliant on substrate-borne vibrational signalling (SBVS), differences in the attractiveness of individual signallers’ calls can influence relative mating success since they can indicate the quality of the sender. We used laser vibrometry and playback experiments to study duet signalling in a psyllid (Anoeconeossa bundoorensis Taylor and Burckhardt), in particular to identify characteristics linked to female responsiveness. Signals were sex-specific with the syllables (or calls) of the smaller males attaining higher frequencies than the syllables of the larger females. Male syllables build over time with more energy in the second half of the call while those of females have a more uniform energy distribution. Male syllables vary in the timing of the halfway point of energy production. We used playback to examine species recognition and female responsiveness to the first and second halves of male calls. We demonstrated that females responded only to calls of male A. bundoorensis and that they were more likely to respond to the first half of male calls. Based on the results of our second playback experiment, we found that a gradation in energy production is more important for eliciting a female reply than a uniform distribution of energy. Our findings show that the rate of energy production facilitates differentiation between syllables which could be used to indicate the quality of individual males as mates.     

The function of Barbary macaque copulation calls.

In a wide variety of animal species, females produce vocalizations specific to mating contexts. It has been proposed that these copulation calls function to incite males to compete for access to the calling female. Two separate advantages of inciting male-male competition in this way have been put forward. The first suggests that as a result of calling, females are only mated by the highest ranking male in the vicinity (indirect mate choice hypothesis). The second proposes that copulation calling results in a female being mated by many males, thus promoting competition at the level of sperm (sperm competition hypothesis). In this paper, I give results from the first experimental study to test these hypotheses. Playback was used to examine the function of copulation calls of female Barbary macaques (Macaca sylvanus) in Gibraltar. Although rank did not affect lone males'' likelihood of approaching copulation calls, when playbacks were given to pairs of males only the higher ranking individual approached. Moreover, females were mated significantly sooner after playback of their copulation call than after playback of a control stimulus. These results suggest that the copulation calls of female Barbary macaques play a key role in affecting patterns of male reproductive behaviour, not only providing an indirect mechanism of female choice, but also promoting sperm competition by reducing the interval between copulations. Potential fitness benefits of inciting male-male competition at these two levels are discussed.     

Relative importance of size versus territorial residency in intraspecific aggression by symbiotic male water mites (Acari: Unionicolidae)

Body size plays an important role in the outcome of aggressive encounters between males of the symbiotic freshwater miteUnionicola formosa. Since intruding males displace resident males as often as resident males successfully defend their territory (mantle cavity of a mussel), resident males do not hold a competitive advantage during male-male aggressive encounters. Larger male size probably enhances a male's success when competing for receptive females.     

The adaptive significance of non-contact mate guarding by males of the dragonfly,Nannophya pygmaea Rambur (Odonata: Libellulidae)

InNannophya pygmaea, ovipositing females were frequently disturbed by conspecific males. Disturbed females often copulated with one of these males or flew away from the pool. Females which flew away from the pool due to male disturbance often returned later the same day and mated with different males. A territorial male would guard his ovipositing mate by hovering above her, presumably trying to prevent her from moving out of his territory. A non-territorial male would also guard his mate in a similar way, both at a vacant water area which was not occupied by any territorial males, or within the territory of a resident male. In addition, both territorial and non-territorial males chased intruding males in an attempt to prevent their mates from being stolen. Territorial males defended their mates better than non-territorial males. Both males and females often mated more than once in the course of a single day. Some territorial males copulated with a new female while another mate oviposited in their territories. This observation supported the “multiple mating hypothesis” proposed by Alcock (1979) and Uéda (1979) but other evidence suggested that this is an inadequate explanation for the non-contact guarding ofN. pygmaea.     

Significance of temporal and spectral acoustic cues for sexual recognition in Xenopus laevis

As in many anurans, males of the totally aquatic species, Xenopus laevis, advertise their sexual receptivity using vocalizations. Unusually for anurans, X. laevis females also advertise producing a fertility call that results in courtship duets between partners. Although all X. laevis calls consist of repetitive click trains, male and female calls exhibit sex-specific acoustic features that might convey sexual identity. We tested the significance of the carrier frequency and the temporal pattern of calls using underwater playback experiments in which modified calls were used to evoke vocal responses in males. Since males respond differently to male and female calls, the modification of a key component of sexual identity in calls should change the male's response. We found that a female-like slow call rhythm triggers more vocal activity than a male-like fast rhythm. A call containing both a female-like temporal pattern and a female-like carrier frequency elicits higher levels of courtship display than either feature alone. In contrast, a male-like temporal pattern is sufficient to trigger typical male-male encounter vocalizations regardless of spectral cues. Thus, our evidence supports a role for temporal acoustic cues in sexual identity recognition and for spectral acoustic cues in conveying female attractiveness in X. laevis.     

Female sooty mangabeys (Cercocebus torquatus atys) respond differently to males depending on the male's residence status-preliminary data

I tested the hypothesis that free-ranging female mangabeys (Cercocebus torquatus atys) differentiate between males depending on their residence status. Adult males in this species employ two group-membership strategies: they are either full-time residents or part-time residents (i.e., present for several weeks at a time, then gone for several weeks, then present again, etc.). In a playback experiment I tested whether adult females with young infants responded differently to calls of adult males that were group members during the previous mating season as compared to males that belonged to neighboring groups during the mating season or were complete strangers. Males whose calls were played were present residents, part-time residents (absent for at least 3 months), neighbors, or complete strangers. Because infanticide is a potential risk in mangabeys, I predicted that females would respond differently to males that had resided in the group when their infant was conceived than they would to those that had not been present. The females showed a clear difference in their reaction to the playbacks of vocalizations from resident and part-time resident males as opposed to nonresidents.