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Courtship by subordinate male Siamese fighting fish, Betta splendens: Their response to eavesdropping and na??ve females

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Abstract

In a social environment, a communication signal may provide information to individuals other than those interacting with the signaler. Eavesdropping is gathering information without be-ing directly involved in the communication interaction. Female Siamese ghting sh, Betta splendens, choose the winner of male-male aggressive interactions based upon information she extracts from eavesdropping.Naïve females, those that have not witnessed the interaction, show no consistent preference for either male. This suggests that losers would be more suc-cessful in courting a naïve female. We conducted two sets of trials: one set tested the losers' courting preference of eavesdropping females or naïve females whereas the other tested the winners' courting preference. We found that losers displayed gill cover erection, a courting behaviour, signi cantly more towards the naïve female than towards the eavesdropping fe-male whereas the winner showed no preference. These results suggest that male B. splendens can moderate their response to an audience in ways more complex than previously appreci-ated. Our data support the suggestion that communication can serve as a network that reaches beyond the immediate signaler and receiver. Understanding the complexity of communication networks will enable us to broaden our ideas about the mechanisms of sexual selection.

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... Second, males and females court each other using displays that are identical to those used in intrasexual contexts (Robertson & Sale 1974). Third, differences in male size and display behavior are known to be relevant to females (Doutrelant & McGregor 2000;Herb et al. 2003), but these traits often covary and few studies have tried to separate them. Fourth, both sexes are likely to be discriminating in their potential mates, as males provide all post-spawning parental care through the construction and maintenance of bubble nests. ...
... Female B. splendens have been shown to avoid males (Bronstein & Jones-Buxton 1996) and males may temper their aggressive behavior in the presence of females, perhaps to avoid deterring them (Doutrelant et al. 2001;Matos & McGregor 2002). Herb et al. (2003) found that females preferred winning males when they observed male-male interactions, presumably because these males would be better able to defend their future nests (cf. Jaroensutasinee & Jaroensutasinee 2001a). ...
... Jaroensutasinee & Jaroensutasinee 2001a). Naïve females in Herb et al.'s (2003) study showed no preference for winning or losing males, suggesting that there was nothing about their intrinsic behavior that was preferable to females. ...
Article
Courtship displays should be exaggerated enough to attract mates and yet tempered so as not to deter them. We tested this hypothesis in the fighting fish Betta splendens by studying courtship displays and body size and their relationships with male parental quality and female fecundity, as well as the effects of display behavior and body size on mate choice decisions and spawning success. Because of their high degree of parental investment, males are expected to be discriminating in their choice of mates. Males who displayed more frequently built larger nests, a measure of parental quality, but larger males did not. When females were paired with males with high display rates, however, the pair had fewer eggs in their nest, even when accounting for female body mass. In a mate choice test using computer-generated male stimuli that differed only in display behavior, females showed no preferences for displaying males vs. non-displaying males, or for males with higher display rates vs. lower display rates. In similar tests in which the computer-generated males differed only in size, females preferred larger males, but also preferred males that differed with respect to body size (negative assortative mating). Males preferred computer-generated females that performed courtship displays over non-displaying females, but showed no preferences for female body size. Neither a female's body size nor her display behavior was a significant predictor of her fecundity as estimated by the number of eggs released during spawning. Thus, our results suggest that female B. splendens must balance male parental quality (nest size) with the risk of potentially disruptive or dangerous behavior during spawning, and that females may minimize these risks through negative size-assortative mating. Female display behavior, while unrelated to fecundity in our study, may attract males because it indicates reproductive readiness or serves a species-recognition function.
... [26][27][28][29][30]), amphibians [31], fish (e.g. [32][33][34][35]) and even in insects [36]. The information an eavesdropper can gain is not always reliable, because one or both of the two interacting individuals might be cheating each other during interaction dependent on the presence or absence of an eavesdropper [37]. ...
... This so-called 'audience effect' or 'bystander effect' has been investigated intensively in several species [90][91][92][93] and especially in fish species (e.g. Siamese fighting fish Betta splendens [34,35], G. aculeatus [94]). In Poeciliids, the audience effect has been investigated in P. mexicana [95][96][97][98], P. reticulata [99], P. latipinna [100] and Xiphophorus birchmanni [101]. ...
Article
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In animals, including humans, the social environment can serve as a public information network in which individuals can gather public information about the quality of potential mates by observing conspecifics during sexual interactions. The observing individual itself is also a part of this information network. When recognized by the observed conspecifics as an audience, his/her presence could influence the sexual interaction between those individuals, because the observer might be considered as a potential mate or competitor. One of the most challenging questions in sexual selection to date is how the use of public information in the context of mate choice is linked to the fitness of individuals. Here, we could show that public information influences mate-choice behaviour in sailfin molly males, Poecilia latipinna, and influences the amount of sperm males transfer to a female partner. In the presence of an audience male, males spent less time with the previously preferred, larger of two females and significantly more time with the previously non-preferred, smaller female. When males could physically interact with a female and were faced with an audience male, three audience females or no audience, males transferred significantly more sperm to a female partner in the presence of an audience male than with female audience or no audience and spent less time courting his female partner. This is the first study showing that public information use turns into fitness investment, which is the crucial factor to understand the role of public information in the dynamic processes in sexual selection.
... This predictive failure of the dyad approach to assessing dominance relationships may result because interactions amongst many individuals may be influenced not only by intrinsic attributes of the individuals (size, weight, age, etc.) but also by extrinsic attributes relating to the social network in which the individuals exist. With regard to social fish, individuals are capable of ascertaining the fighting abilities of other individuals that they observe in contests (McGregor, 1993;Johnsson and Akerman, 1998;Oliveria et al., 1998;Herb et al., 2003) and some of these bystanders even undergo hormonal changes when observing such contests ( Oliveira et al., 2001) or develop preferences for interacting with those who have not seen them lose an encounter ( Herb et al., 2003). As with these studies, our results-wherein hierarchies determined by dyads were the same as hierarchies seen in group settings only 50% of the time-indicate that dyadic relationships are not an effective predictive tool for determining a dominance hierarchy in a group of four to five fish. ...
... This predictive failure of the dyad approach to assessing dominance relationships may result because interactions amongst many individuals may be influenced not only by intrinsic attributes of the individuals (size, weight, age, etc.) but also by extrinsic attributes relating to the social network in which the individuals exist. With regard to social fish, individuals are capable of ascertaining the fighting abilities of other individuals that they observe in contests (McGregor, 1993;Johnsson and Akerman, 1998;Oliveria et al., 1998;Herb et al., 2003) and some of these bystanders even undergo hormonal changes when observing such contests ( Oliveira et al., 2001) or develop preferences for interacting with those who have not seen them lose an encounter ( Herb et al., 2003). As with these studies, our results-wherein hierarchies determined by dyads were the same as hierarchies seen in group settings only 50% of the time-indicate that dyadic relationships are not an effective predictive tool for determining a dominance hierarchy in a group of four to five fish. ...
Article
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Unlike other balistids, grey triggerfish Balistes capriscus occur in social groups in subtropical reef assemblages and have been noted to cooperate in capturing large crustacean prey. The objective of this study were to determine the structure of dominance hierarchies of these social groups and the factors that influence hierarchies of wild-caught grey triggerfish in a naturalistic setting. From observations of four groups of triggerfish (n = 19 fish) in both dyad and group (4 – 5 fish) settings, we provide a description of triggerfish behaviors and coloration patterns and an explanation of the social context in which suites of behaviors are used by dominant, middle-ranking, and subordinate fish. Sixteen behaviors and nine coloration patterns were noted for grey triggerfish. Grey triggerfish groups form linear hierarchies in both dyads and groups as measured by Landau’s Index of Linearity (h = 1.0 for Groups 1, 3, and 4 and h = 0.95 for Group 2 in dyads; h = 1.0 for all groups in group settings). Dyadic hierarchies, however, were not necessarily good predictors of the hierarchies found in larger group settings, as they only predicted two of the four group hierarchies. Sex played no role in influencing status or behavior. Size had the greatest influence on dominance status, with larger fish being more dominant than smaller fish. An individual’s dominance ranking influenced both body coloration and posture. These results suggest that color patterns and body postures may also be used by observers as an indicator of an individual’s social status in groups [Current Zoology 56 (1): 18–35 2010].
... There is widespread evidence for audience effects, in competitive as well as cooperative contexts (for competitive: [6][7][8]; for cooperative: [9,10]). To date, many studies on various taxa have focused on the modulation of the communication system in response to audience effects (chickens, Gallus gallus domesticus: [11]; house sparrow, Passer domesticus: [12]; marmots, Marmota flaviventris: [13]; and various primates species: chimpanzees, Pan troglodytes: [14][15][16]; brown capuchins, Cebus paella: [17]; vervet monkeys: [18]) but only a few have studied a modification of behaviour (siamese fish, Beta splendens: [19]; cichlids, Astatotilapia burtoni: [20] and primates: rhesus macaques, Macaca mulatta: [21,22]; orangutans, Pongo pygmaeus: [23]). However, experimental studies manipulating the audience composition to test such effects remain scarce [10,[19][20][21]24]. ...
... To date, many studies on various taxa have focused on the modulation of the communication system in response to audience effects (chickens, Gallus gallus domesticus: [11]; house sparrow, Passer domesticus: [12]; marmots, Marmota flaviventris: [13]; and various primates species: chimpanzees, Pan troglodytes: [14][15][16]; brown capuchins, Cebus paella: [17]; vervet monkeys: [18]) but only a few have studied a modification of behaviour (siamese fish, Beta splendens: [19]; cichlids, Astatotilapia burtoni: [20] and primates: rhesus macaques, Macaca mulatta: [21,22]; orangutans, Pongo pygmaeus: [23]). However, experimental studies manipulating the audience composition to test such effects remain scarce [10,[19][20][21]24]. ...
Article
Group living promotes opportunities for both cooperation and competition. Selection on the ability to cope with such opposing social opportunities has been proposed as a driving force in the evolution of large brains in primates and other social species. However,we still knowlittle about the degree of complexity involved in such social strategies. Here, we report advanced social strategies in wild vervet monkeys. Building on recent experimental evidence that subordinate females trade grooming for tolerance from higher-ranking individuals during foraging activities,we showthat the audience composition strongly affects this trade. First, tolerance was lower if the audience contained individuals that outranked the subordinate partner, independently of audience size and kinship relationships. Second, we found a significant interaction between previous grooming and relative rank of bystanders: dominant subjects valued recent grooming by subordinates while intermediate ranked subjects valued the option to aggress subordinate partners in the presence of a dominant audience. Aggressors were also more likely to emit coalition recruitment calls if the audience contained individuals that outranked the subordinate partner. In conclusion, vervet monkeys include both recent grooming and knowledge about third-party relationships to make complex decisions when trading grooming for tolerance, leading to a finely balanced trade-off between reciprocation and opportunities to reinforce rank relationships. © 2017 The Author(s) Published by the Royal Society. All rights reserved.
... Among the recent experimental studies are those investigating 'eavesdropping' and audience effects: how observation of a contest in a pair affects third parties or how being observed by third parties affects the participants in a pair contest (e.g. McGregor, 1993;Johnson & Akerman, 1998;Oliveira et al., 1998;Herb et al., 2003). This research indicates, for example, that observers in Siamese ghting sh behave differently towards animals that they have seen either win or lose a contest than they behave towards animals that have either won or lost but have not been observed to do so (Oliveira et al., 1998). ...
... In particular, they are slower to approach and display to observed winners than to observed losers and equally quick to approach and display to unobserved winners and losers. Also in Siamese ghting sh individuals that observe contests experience hormonal changes in comparison to non-observers (Oliveira et al., 2001), and male Siamese ghting sh that have lost contests 'prefer' females that have not observed them losing over those females that have observed them losing (Herb et al., 2003). ...
Article
We performed experiments with cichlid fish to test whether several basic aspects of dominance were the same in isolated pairs as in pairs within a social group of three or four. We found that the social context, whether a pair was isolated or within a group, strongly affected the basic properties of dominance relationships. In particular, the stability of relationships over time, the replication of relationships in successive meetings, and the extent of the loser effect were all significantly less in socially embedded pairs than in isolated pairs. We found no significant winner effect in either isolated or socially embedded pairs. These findings call into question many current approaches to dominance that do not consider social context as an important factor in dominance behavior. These findings also cast serious doubt on the validity of empirical and theoretical approaches based on dyadic interactions. Among these approaches are game theoretic models for the evolution of aggressive behavior, experimental designs evaluating how asymmetries in attributes influence the outcome of dominance
... First, this finding is similar to that of studies on winner ⁄ loser effects where males are more aggressive after winning a fight than after losing. Second, females may prefer males they have seen win fights and males quickly escalate their aggression levels to both obtain a mating and expel a rival male (Doutrelant & McGregor 2000;Herb et al. 2003). A final possibility is that males increased biting because the dummy could not reciprocate the aggression and, therefore, appeared subordinate. ...
... The social interactions of Betta have been studied in the contexts of dominance hierarchies (Goldstein, 1975;Rhoad et al., 1975;Lobb and McCain, 1976;Cain and Baenninger, 1980), mate choice (Doutrelant and McGregor, 2000;Herb et al., 2003), and communication networks (Oliveira et al., 1998;Doutrelant and McGregor, 2000;Doutrelant et al., 2001). The social partner preferences of Betta, however, have been largely overlooked. ...
Article
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While the social interactions of Betta splendens have been studied in the contexts of dominance hierarchies, mate choice and communication networks, the social partner preferences of Betta have been largely overlooked. In this study, we presented male and female Betta with a single male, a single female, and a group of three females in dichotomous choice tests in order to better understand basic social interactions in this largely nonsocial species. The highly territorial Betta preferred associating with conspecifics in nearly every configuration we tested, with exceptions noted when single females were given the choice between a lone male and an empty chamber, and when males were presented with a single female and an empty chamber. Also, in most tests, the fish chose to spend more time with the larger group of females. The motivation for this preference certainly varied from reproductive to anti-predator. While such behavior might not suggest true shoaling behavior, it does demonstrate a subtle degree of sociality.
... As with functional reference little is known, however, about the cognitive mechanisms that underpin audience effects of call production (Zuberbühler 2008). The fact that fish and domestic chickens adjust general social behavior according to the audience composition (Grosenick et al. 2007;Herb et al. 2003;Marler et al. 1986) means that complex cognitive tools may not be required for audience effects to take place: it might just be that the presence of certain individuals is an external stimulus that automatically triggers certain behaviors. Primates, however, seem to modulate their social communication as a function of very subtle social cues, such as the attention of others (Liebal et al. 2004) and their capacity to help (Slocombe and Zuberbühler 2007), indicating that the psychological mechanisms involved in audience effect in primates might be more complex that in other animals (Zuberbühler 2008). ...
Article
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Language is a uniquely human trait, and questions of how and why it evolved have been intriguing scientists for years. Nonhuman primates (primates) are our closest living relatives, and their behavior can be used to estimate the capacities of our extinct ancestors. As humans and many primate species rely on vocalizations as their primary mode of communication, the vocal behavior of primates has been an obvious target for studies investigating the evolutionary roots of human speech and language. By studying the similarities and differences between human and primate vocalizations, comparative research has the potential to clarify the evolutionary processes that shaped human speech and language. This review examines some of the seminal and recent studies that contribute to our knowledge regarding the link between primate calls and human language and speech. We focus on three main aspects of primate vocal behavior: functional reference, call combinations, and vocal learning. Studies in these areas indicate that despite important differences, primate vocal communication exhibits some key features characterizing human language. They also indicate, however, that some critical aspects of speech, such as vocal plasticity, are not shared with our primate cousins. We conclude that comparative research on primate vocal behavior is a very promising tool for deepening our understanding of the evolution of human speech and language, but much is still to be done as many aspects of monkey and ape vocalizations remain largely unexplored.
... Due to the fact that female B. splendens eavesdrop on male contests, Herb et al. (2003) tested mate-choice decisions of male fighting fish after losing or winning a contest. They conducted two sets of trials: in one set-up they tested the losers' courting preference for eavesdropping females or naïve females (females which had no opportunity to observe the contestants). ...
Chapter
IntroductionSexual imprintingLearning after reaching maturityEavesdroppingMate-choice copyingSocial mate preferences overriding genetic preferencesCultural evolution through mate-choice copyingDoes mate-choice copying support the evolution of a novel male trait?Is mate-choice copying an adaptive mate-choice strategy?OutlookConclusions References
... Due to the fact that female B. splendens eavesdrop on male contests, Herb et al. (2003) tested mate-choice decisions of male fighting fish after losing or winning a contest. They conducted two sets of trials: in one set-up they tested the losers' courting preference for eavesdropping females or naïve females (females which had no opportunity to observe the contestants). ...
Chapter
Full-text available
IntroductionSexual imprintingLearning after reaching maturityEavesdroppingMate-choice copyingCultural evolution through mate-choice copyingDoes mate-choice copying support the evolution of a novel male trait?Is mate-choice copying an adaptive mate-choice strategy?OutlookConclusions Acknowledgements
... For example, an audience may influence the intensity of aggressive male–male interactions (Zajonc, 1965; Doutrelant et al., 2001; Matos & McGregor, 2002; Matos et al., 2003; Dzieweczynski et al., 2005 Dzieweczynski et al., , 2006). Some studies reported that individuals may base their mate choice on information extracted from the observation of interactions among others (eavesdropping; Doutrelant & McGregor, 2000; Herb et al., 2003), but the question of whether, and to what extent, the presence of an audience influences mate choice decisions remained little investigated. Here, we report that an audience affects the strength of expression of a male mating preference . ...
Article
Audience effects occur when an observing (by-standing) animal influences the behaviour of an observed individual. A recent study (Plath, M., Blum, D., Schlupp, I. & Tiedemann, R., Anim. Behav. 75, 21-29 (2008)) has demonstrated an effect of a visual audience male on male mating preferences in the surface form of a livebearing fish, the Atlantic molly (Poecilia mexicana). Surface dwelling P. mexicana are highly aggressive; hence, males dedicating simultaneous attention to mate choice and aggressive interactions may explain this audience effect. Here we examined the effect of an audience on male mate choice in the cave form of that species, which — unlike other cavefishes — have maintained eyes and still respond to visual cues under experimental conditions. Cave mollies were especially interesting to study, because they have reduced aggressive behaviour. We gave males an opportunity to choose between two females, and we repeated the tests with an audience male present. The focal males tended to divide their attentions more equally between the two females when an audience male was presented. The observed effect did not statistically differ between surface and cave dwelling P. mexicana, suggesting that (1) the response to a visual audience is maintained in this cavefish and (2) the described audience effect is largely independent of aggressive interactions among males. Generally, its adaptive significance may be linked to the avoidance of sperm competition when males sharing the same (innate) preferences compete for mates. Moreover, males may conceal their preference to prevent other males from copying their mate choice.
... In each fish replica context, the dummy displayed its body axis orthogonal to the central longitudinal line of the tank, to exhibit the lateral colour pattern of its body. In addition, the fish replica was oscillating on its longitudinal axis with an angle of 30°in amplitude and with a frequency of 0.5 Hz to emulate the decreased locomotor activity of real B. splendens behaving individually in a tank (e.g. a fish used as control, having no visual contact with the conspecific), as well as a female starting mate evaluation or eavesdropping (Doutrelant et al., 2001;Herb et al., 2003;Cantalupo et al., 1996;Clotfelter et al., 2006). The fish replica autonomously yawed 180°every 5 min to invert the head-tail orientation, to avoid positional bias. ...
Article
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Among territorial animals, several species are characterized by males showing the same initial behaviours towards both sexes, leading to significant chances of injuries against conspecifics. In this study, we investigated how visual stimuli exhibited by a female-mimicking robotic replica can be exploited by highly territorial Betta splendens males to discriminate males from females. In addition, we tested the effect of light stimuli, mimicking the colour pattern of a reproductive female, on the consistence of courtship displays in B. splendens males. The intensity of male behaviours used in both courtship and not-physical agonistic interactions (e.g. fin spreading and gill flaring) was not importantly modulated by different stimuli. Conversely, behavioural displays used specifically in male–female interactions significantly increased when the robotic replica colour pattern mimicked a reproductive female. Furthermore, male courtship behaviours obtained in response to the robotic replica exhibiting light stimuli were comparable with responses towards authentic conspecific females. Our biomimetic approach to establish animal–robot individual interaction can represent an advanced strategy for trait-based ecology investigation, a rapidly developing research field.
... Female mate choice can be affected by external factors such as the physical and social signalling environment (Herb et al. 2003;Matos et al. 2003;Gordon & Uetz 2011;Clark et al. 2012) and previous experience (Tudor & Morris 2009;Rutledge et al. 2010;Bailey 2011;Wong et al. 2011). Female mate choice can also be affected by internal factors such as genetics (Tregenza & Wedell 2000;Chenoweth & Blows 2006;Horth 2007) and female condition (Cotton et al. 2006a). ...
Article
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Many male signallers convey information to female receivers in multimodal courtship displays. While much is known about how males vary in terms of signalling, variation in female detection of these multimodal signals is relatively unexplored. We suggest that there is a critical, albeit underdeveloped, link between multimodal sensory reception and individual variation in mate choice. This review addresses the potential effects of developmental and conditional factors (e.g. nutrient availability, hormone profiles and age) on female multimodal processing, and illustrates that differences in the (1) source of individual variation and (2) the number of sensory processing modes affected by this variation can influence the receiver's mate choice patterns. Based on these two factors, we outline novel predictions of preference functions and choosiness in a redundant multimodal signalling context. Moreover, we explore the theoretical implications of individual variation in multimodal signal perception in relation to sensory drive, honest signalling, assortative mating and intrasexual selection. We propose that understanding the role of variation in sensory processing and its relation to mate choice can help us better identify the factors that influence sender and receiver fitness, and subsequently the rate and direction of signal evolution. (c) 2012 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.
... Social eavesdropping has been described in a wide variety of animal taxa as an attempt to exploit information in signals of conspecifics in varied social contexts. These contexts may include assessment of potential mates by observing preferences of others, i.e., mate choice copying (Auld and Godin 2015;Gierszewski et al. 2018), assessment of potential mates based on observing their performance in competitive interactions (Doutrelant and McGregor 2000;Ophir and Galef 2003;Herb et al. 2003;Crockford et al. 2007;Loranger and Bertram 2016), assessment of potential rivals through observing contest outcome (Peake et al. 2001;Matessi et al. 2005;Garcia et al. 2019), and a means of mate competition via detection and location of females (Kiflawi and Gray 2000;Peake 2005;Balsby and Dabelsteen 2005;Crockford et al. 2007;Hauber and Zuk 2010;Milner et al. 2010;Smith et al. 2011;Webster and Laland 2013;Stoffer et al. 2016). ...
Article
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Multimodal courtship signals may compensate for environmental interference or loss of signals in some sensory modes but may also increase detection by eavesdroppers. Studies on the wolf spider Schizocosa ocreata (Hentz) have demonstrated that males eavesdrop on visual courtship cues of other males and subsequently initiate courtship. Since S. ocreata males use multimodal courtship signals, we examined responses of males to playback of signals in different sensory modes (visual, vibration, multimodal) to test their relative importance for eavesdropping on courting male rivals. We used a recently developed technique to present male wolf spiders with video and/or vibratory stimuli: (1) a multimodal courting male stimulus, with synchronous visual and vibratory cues; (2) a visual-only courting male stimulus (minus the vibratory cues); (3) vibratory cues only (minus the visual cues); and (4) a control (visual background, no courtship). In single-presentation (no-choice) tests, males displayed more courtship bouts and longer durations of courtship bouts to the vibratory stimulus compared with either the visual or multimodal stimuli. However, in two-choice tests where isolated vibratory cues were paired against visual or multimodal stimuli, test males responded with more courtship bouts and longer durations of courtship bouts to the multimodal and visual stimuli. Results of these experiments suggest that male wolf spiders may vary eavesdropping courtship behaviors to compensate for missing sensory information concerning the location and other distinguishing characteristics of the rival male and the whereabouts of the female. Significance statement Social eavesdropping is used to exploit information in signals of conspecifics, e.g., as a means of mate competition. Studies on Schizocosa ocreata wolf spiders have shown that males eavesdrop on visual courtship displays of other males and subsequently initiate multimodal courtship. We used video/vibration playback to examine responses of males to signals in different sensory modes (visual, vibration, both) and determine their relative importance for eavesdropping on courting male rivals. Results suggest that depending on sensory modes of their rivals’ signals, males may vary their own courtship displays to compensate for missing information. For example, when signals are presented individually, eavesdropping male response patterns are distinctly different from when presented a choice between modes. Males show more displays with isolated vibratory signals, but given a choice, males more often increase their tapping when multimodal cues were present. Ultimately, eavesdroppers that adjust behaviors in response to available sensory cues would be able to “level the playing field” with rivals and potentially increase the probability of gaining attention of females.
... First, this finding is similar to that of studies on winner ⁄ loser effects where males are more aggressive after winning a fight than after losing. Second, females may prefer males they have seen win fights and males quickly escalate their aggression levels to both obtain a mating and expel a rival male (Doutrelant & McGregor 2000;Herb et al. 2003). A final possibility is that males increased biting because the dummy could not reciprocate the aggression and, therefore, appeared subordinate. ...
Article
Consistent individual differences in behavior suggest that individuals respond in a predictable and repeatable manner in a specific situation while differing from other individuals. Male Siamese fighting fish exhibit consistent individual differences in decision‐making strategies when they encounter a receptive female and a rival male simultaneously. However, whether these differences are altered by recent experience is unknown. We examined the influence of repeated aggressive encounters on behavioral consistency and decision‐making. Males were presented with paired female–male dummies prior to any aggressive experiences to obtain a baseline measure. Next, males either won or lost three consecutive contests against rivals and then received the paired female–male dummies after each of these encounters. Overall levels of highly aggressive behaviors were affected by contest outcome, while levels of female‐directed were not. Not surprisingly, winning a fight led to an increase in male‐directed bites, an overtly aggressive behavior that only occurs after fights have escalated. Fighting a male before encountering the dummies caused males to perform more tail beats to the dummy male, perhaps as a result of increased motivation. Males exhibited similar levels of repeatability and used the same strategies when faced with conflicting stimuli regardless of fighting experience. Thus, while winning or losing a fight impacts overall aggression, it does not influence behavioral consistency. This study demonstrates that consistent individual differences and decision‐making strategies may be resistant to recent aggressive experiences, even over a period of days.
... Signaling may be affected by other animals directly interacting with the sender (Doutrelant et al. 2001) or the presence of animals not directly involved in the interaction (Doutrelant and McGregor 2000). The audience gathers information from conspecifics (Herb et al. 2003) in order to evaluate their potential to defend resources (McGregor and Peake 2000). Nevertheless, the audience presence may affect the type of signals sent (Semple et al. 2009), and thus involves additional costs and/or benefits for the signalers (Matos and McGregor 2002). ...
Article
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Animal communication occurring in wide networks can involve signals sent to several receivers. The animals composing the audience may affect how individuals display during an interaction and may change the message that is sent. In this study, we investigated whether the presence of a conspecific affected the intensity of agonistic interaction between male fiddler crabs, Uca maracoani. Pairs of males of the same size were observed when in the presence of a male, a female or no crab as audience. We found that if there is a female audience, males became more aggressive and interacted the most. Also, the female audience leads to an increase in incidence of male foaming, possibly indicating predisposition for mating. If the audience was a male or if there was no audience, there was no significant difference in interaction. These results indicate that the presence of an audience affects the way male fiddler crabs interact and the type of displays they show. Therefore, the context seems to guide the fiddler crab behavior in terms of how they perform in order to send information about themselves to conspecifics.
... The audience effect is a wide-spread phenomenon and has been studied in various contexts (e.g., feeding [77], food caching [78], predator detection [79], and mate choice [80]). The audience effect has been studied in various taxa (insects [81,82], birds [83][84][85][86][87][88], mammals [78,79,[89][90][91][92]), especially in fish: e.g., in the fighting fish Betta splendens [93][94][95], in the three-spine stickleback G. aculeatus [96], in the guppy P. reticulata [64,80], in the sailfin molly P. latipinna [97,98], and in the Atlantic molly P. mexicana [99][100][101]. ...
Article
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Animals often use public information for mate-choice decisions by observing conspecifics as they choose their mates and then copying this witnessed decision. When the copier, however, is detected by the choosing individual, the latter often alters its behavior and spends more time with the previously non-preferred mate. This behavioral change is called the audience effect. The deception hypothesis states that the choosing individual changes its behavior to distract the audience from the preferred mate. The deception hypothesis, however, only applies if the audience indeed copies the pretended mate choice of the observed individual. So far, this necessary prerequisite has never been tested. We investigated in Atlantic molly males and females whether, first, focal fish show an audience effect, i.e., alter their mate choices in the presence of an audience fish, and second, whether audience fish copy the mate choice of the focal fish they had just witnessed. We found evidence that male and female Atlantic mollies copy the pretended mate choice of same-sex focal fish. Therefore, a necessary requirement of the deception hypothesis is fulfilled. Our results show that public information use in the context of mate choice can be costly.
... However, existing stud- ies investigating the link between personalities, or non-sexual behaviour in general, and mate choice are rare and deliver divergent results. Some studies found a general preference for [17][18][19] or against [20,21] certain behavioural traits among females of a species. Other studies found females to differ in their mating preference, depending on their own behavioural type, leading to positive assortment [22][23][24] or dis-assortment [25]. ...
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Although personality traits can largely affect individual fitness we know little about the evolutionary forces generating and maintaining personality variation. Here, we investigated the hypothesis that personality variation in aggression is sexually selected in the monogamous, bi-parental cichlid Pelvicachromis pulcher. In this species, breeding pairs form territories and they aggressively defend their territory and offspring against con- and heterospecific intruders. In our mate choice study, we followed up two alternative hypotheses. We either expected females to show a directional preference for a high level and high consistency of aggression (potentially indicating mate choice for male parental quality). Alternatively, we expected females to choose males for (dis-)similarity in the level/consistency of aggression (potentially indicating mate choice for compatibility). Individual level and consistency of aggression were assessed for males and females using mirror tests. After eavesdropping on aggressive behaviour of two males (differing in level and consistency of aggression) females were then allowed to choose between the two males. Males, but not females, showed personality variation in aggression. Further, females generally preferred consistent over inconsistent males independent of their level of aggression. We did not detect a general preference for the level of male aggression. However, we found an above average preference for consistent high-aggression males; whereas female preference for inconsistent high-aggression did not deviate from random choice. Our results suggest behavioural consistency of aggression in male rainbow kribs is selected for via female mate choice. Further, our study underlines the importance of considering both the level and the consistency of a behavioural trait in studies of animal behaviour.
... Such social eavesdropping can be beneficial for the bystander as it provides a relatively low-cost, low-risk alternative to gathering the same socio-sexual information acquired through direct interaction with the individual being observed (Dabelsteen 2005). For the individual being eavesdropped upon, however, the opposite may be true (Dabelsteen 2005 witnessed a male-male aggressive interaction, subsequently preferred as a mate the winner of the interaction over the loser, whereas females that did not witness the aggressive interaction showed no bias towards either the winner or loser male (Herb et al. 2003). In this case, the presence of a female audience was beneficial for the winner of the fight, but detrimental for the loser. ...
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Males in polygamous mating systems may inadvertently transmit information regarding their mating preferences to bystanding sexual competitors, thereby permitting bystanders to use this information to enhance their own mating success by copying the mate choice of signallers. If males are at risk of having their mate choice copied and consequently face a higher risk of sexual competition, then selection should favour males that reduce conspicuous mating behaviours in the presence of an audience of sexual competitors. In the current study, we used the Trinidadian guppy (Poecilia reticulata), a species that exhibits alternative male mating tactics, to test this sexual competition avoidance hypothesis and the predictions that males would decrease their overall mating effort and exhibit fewer conspicuous courtship displays relative to more inconspicuous sneak mating attempts in the presence of either one or two sexual rivals compared to the absence of any audience. Male guppies significantly reduced their overall mating effort in the presence of increasing numbers of rival audience males. This was reflected in similar monotonic decreases in the frequencies of courtship displays and sneak mating attempts and in the proportional use of courtship displays (relative to sneak mating attempts) across treatments. These findings are consistent with the sexual competition avoidance hypothesis. Our novel results contribute to an increasing body of knowledge showing that the social environment can influence the mating effort and mating decisions of individuals and thus have important implications for sexual selection and evolution.
... However, it has recently been argued that agonistic behavior fits to the context of the broader social environment instead of dyadic interactions (Snekser et al 2006) and phenomenon known as communication networks (i.e. the selective environment of networks of signaling and receiving individuals (McGregor 1993)) has been successfully discussed in B. splendens in terms of eavesdropping and its subsequent effects on intra and/or intersexual interactions (eg. Oliveira et al 1998;Doutrelant et al 2001;Herb et al 2003;Matessi et al 2010). Generally, it has been accepted that female B. splendens chooses the winner of male-male aggressive interactions based upon information she obtains from eavesdropping (Snekser et al 2006). ...
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There has been a growing trend towards examining interactions in the context of communication networks consisting of many individuals rather than isolated pairs of signaling animals. An important question in state-dependent behavior is how multiple influences on state are integrated to determine current behavior. The goal of the following study was to investigate whether the availability of female, both in terms of density and accessibility, affects male-male interactions in male Siamese fighting fish, Betta splendens. For this purpose, direct male-male fighting trials were performed considering 12 different scenarios. The findings indicated that it is the physical isolation and/or presence of an escorting male in the community, not female density, which modifies male-male interactions. These results suggest that the resource holding power (RHP) in a specimen can be innately recognized by other opponents in bettas.
... For example, Marler and colleagues (1986) observed that male domestic chickens (Gallus gallus) preferably gave food calls when they discovered foods in presence of hens. Another interesting case of audience effect was reported by Herb et al. (2003). In Siamese fighting fish (Betta splendens), males who have lost a fight with another male spend less time displaying to a female who was a bystander of the contest compared to naïve females. ...
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While allogrooming (i.e. the visual examination, searching and manipulation of the skin and hair of another individual-hereafter grooming) is one of the most studied behaviours in the field of primatology and is assumed to be an important component of sociality in many non-human primate species, its potential contagion among group mates in group-living species is poorly studied. Grooming is known to provide many social benefits like tolerance from dominant individuals or tolerance over food. Moreover, grooming has also been shown to be associated with positive emotional state through the anxiolytic and relaxation effects this behaviour provides to both givers and receivers of grooming. In group-living species, social interactions generally occur in the presence of other group members. Many studies provided evidence that the behaviour of the individuals in a group impacts the behaviour of neighbouring bystanders. The purpose of this study is to determine whether observing grooming interactions involving group mates impacts the behaviour, mainly grooming, and the emotional state of surrounding bystanders. To do so, observational research on social behaviour was carried out in twenty semi-free living adult female Barbary macaques (Macaca sylvanus) at Trentham Monkey Forest (Stoke-on-Trent, UK). Grooming and Post-Grooming Observations (G-PGOs) and Matched Controls (MCs) were collected. During these observations, the time to the next grooming interaction involving the focal individual after the observation of group mates involved in a grooming interaction, as well as the affiliative, aggressive and self-directed behaviours expressed by the focal individuals and the proximity to others were recorded. Results provided evidence that grooming is contagious among group mates. A reduced anxiety level was also observed during G-PGOs compared to corresponding MCs, suggesting that the bystanders of grooming interactions experienced the same emotional state as individuals involved in grooming interaction. A lower proximity to others and a higher rate of affiliative behaviours were observed during G-PGOs compared to MCs, suggesting an increase of tolerance to others and a more general contagion of affiliation. All together, these results provide evidence that the contagion of grooming and the positive emotional state associated to this behaviour may contribute to maintain group cohesion by strengthening social bonds, and may encourage affiliative interactions among group mates. Such simple psychological phenomenon may have many consequences on the general interactions and exchanges between individuals and may underpin within group cooperation and prosociality. These findings may also help to improve the welfare of captive non-human primates living in social groups. ii
... Male Atlantic mollies redirect their attentions to non-preferred females when they know another male is eavesdropping. (Herb et al. 2003;Plath et al. 2008) ...
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More than 1.5 billion fish are traded internationally each year. Fish are the most numerous type of pet and are kept in approximately 10% of Western households. Several studies indicate that most people do not consider fish welfare to be important, although scientific evidence shows that fish are behaviourally complex and feel pain. The aim of this thesis is to develop pathways for improving the welfare of pet fish. The welfare of captive fish is influenced by factors contributing to fish health (such as water quality, stress and behavioural needs) as well as factors contributing to the owner’s provision of care (such as knowledge, attitudes, social norms and media coverage). The relationships between these factors were explored using three methods: a survey of fish owners; an intervention using a short film to improve owner attitudes and behaviour; and the development of preference and motivational testing for determining the value of enrichment for fish. The survey identified fish owners’ perceptions of the main welfare issues affecting pet fish and helped model the attitudes that underlie aquarist behaviour. Fish owners responding to the survey (n = 534) reported that disease and old age were the most common causes of death for their pet fish, although it is likely that many of them underestimated the role of water quality in fish health. The majority of respondents (73%) reported that they are knowledgeable about fish care and actively share and seek information about their fish. However, more than a quarter of respondents (27%) admitted that they had limited knowledge of fish care and rarely sought information or social support for their hobby. Almost all respondents provided structural enrichment such as gravel and shelters for their fish, but less knowledgeable owners were more likely to provide artificial plants than real ones. Providing fish owners (n = 195) with a short film encouraging them to clean their aquaria weekly did increase the frequency of tank cleaning, but only if they already intended to do so. There was no measurable change in attitudes after watching the film, but using a positively framed film appeared to increase recall of the key message compared to the negatively framed film.
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The impact of social environment on mating success is especially pronounced in species where both intraspecific and interspecific selection influence reproduction, such as the Siamese fighting fish. Males alter male–male interactions when either a male or female audience is present, but how males change their behavior toward a female when a rival male is present is unknown. This study addresses whether males alter their behavior toward a female in a way that would prevent a rival male from interrupting courtship. The behavior of male Siamese fighting fish toward a dummy female was examined under various degrees of visual cover, both in the presence and absence of a rival male, to investigate whether males use concealment provided by the structural environment to their advantage. While males did not use barriers to conceal courtship as hypothesized, males altered their behavior by increasing courtship and monitoring their nest when a rival was visible. This increase in courtship is in contrast to most studies on courtship in the presence of a rival that find a reduction in courtship behavior. Males spent more time opercular gill flaring when no barriers were present, suggesting that males may be trying to court the female and communicate to the rival simultaneously. There was also a trend for aggression toward the female and the rival to decrease as screen length increased. Thus, males compensate for the presence of a rival by adjusting their courtship and aggressive behaviors, which could have important implications for courtship success.
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Unlike other balistids, grey triggerfish Balistes capriscus occur in social groups in subtropical reef assemblages and have been noted to cooperate in capturing large crustacean prey. The objective of this study were to determine the structure of dominance hierarchies of these social groups and the factors that influence hierarchies of wild-caught grey triggerfish in a naturalistic setting. From observations of four groups of triggerfish (n = 19 fish) in both dyad and group (4 – 5 fish) settings, we provide a description of triggerfish behaviors and coloration patterns and an explanation of the social context in which suites of behaviors are used by dominant, middle-ranking, and subordinate fish. Sixteen behaviors and nine coloration patterns were noted for grey triggerfish. Grey triggerfish groups form linear hierarchies in both dyads and groups as measured by Landau's Index of Linearity (h = 1.0 for Groups 1, 3, and 4 and h = 0.95 for Group 2 in dyads; h = 1.0 for all groups in group settings). Dyadic hierarchies, however, were not necessarily good predictors of the hierarchies found in larger group settings, as they only predicted two of the four group hierarchies. Sex played no role in influencing status or behavior. Size had the greatest influence on dominance status, with larger fish being more dominant than smaller fish. An individual's dominance ranking influenced both body coloration and posture. These results suggest that color patterns and body postures may also be used by observers as an indicator of an individual's social status in groups [Current Zoology 56 (1): – 2010].
Article
Males of several species have been shown to alter their mate preference in the presence of an eavesdropping rival. This evasive tactic has been interpreted as an attempt by the courting male to drive the attention of the rival away from the preferred female. The fitness return of this deceptive strategy will depend on the costs of cheating for the actor (the displayer) and the benefits for the rival (the bystander) of copying the choice of the displayer. We developed a two-person nonzero sum game between two males that compete for mating with one of two receptive females. Males could assess female quality with a varying level of uncertainty, which was modelled using a Bayesian statistical decision theory approach. We explored the actor and bystander payoffs under different levels of uncertainty in mate assessment and difference in quality between females. We found that when being eavesdropped on is costly (i.e. when females differ largely in quality), males are expected to cheat to reduce the amount of public information that is available to the unintended audience. However, under these circumstances, the value of the public information is low and the bystander is not expected to copy the choice of the actor. Our model suggests that deceptive male choice may evolve only under relatively restricted conditions and suggest that other explanations, such as, for example, a reduction in the risk of precopulatory male-male competition may be more likely. Future theoretical and empirical work will be necessary to test alternative interpretations of the audience effects in male mate choice.
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Male Siamese fighting fish, Betta splendens, are known for their ferociousness when defending their territories against male or female conspecific intruders. The aim of this study was to investigate whether this species exhibits the dear enemy phenomenon, where territorial males would be less aggressive toward neighbors with already established territories than toward complete stranger males. In experiment 1, a male Betta was placed in an aquarium. A second male was placed in a glass jar that was fitted in one corner of the aquarium. After 24 h, a cover was placed around the jar, and then removed after 1 h. Alternatively, the jar male was replaced with a different male of a different color while the jar was covered. The frequency and duration of opercular expansion by the aquarium male were recorded. In experiment 2, two male Bettas were placed, each in a half of a large aquarium that was equally divided by a clear, perforated Plexiglas divider. After 24 h, one male (intruder) was transferred into the other male’s (resident) half. In another variation of the experiment, the intruder was a stranger male Betta that the resident male had never encountered before. The opercular expansion and duration were scored for the resident male. The results indicated that male Siamese fish reacted similarly to familiar and stranger males. This lack of dear enemy effect in Siamese fish could be due to an inability to discriminate between neighboring males and non-neighboring males. Alternatively, territorial males could be equally aggressive to all intruders because all intruders represent equal danger. Key wordsSiamese fighting fish-dear enemy-conspecific discrimination
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Social cognition, in particular the derivation of social information from observation of interactions between members of a social group, has been widely investigated in primates, but it has received little attention in other social mammals, although it has been anecdotally reported in the domestic dog, Canis familiaris. We recorded the behaviour of dogs (‘spectators’) that had observed controlled interactions between a human and a dog (the ‘demonstrator’) competing for an object, and that were subsequently allowed to interact freely with both participants. When the competitions were playful, as indicated by signals performed by the human, the spectator was more likely to approach the winner first and/or more rapidly, suggesting that winners of games are perceived as desirable social partners. When the human did not perform play signals, changing the social context from play to contest over a resource, spectators were slower to approach either of the participants, suggesting that participants in contests were less desirable as social partners than participants in games. If the dog was prevented from seeing the game, it still reacted differently to the winner and the loser, but its behaviour was not the same as after games that it had seen. We conclude that spectator dogs gain information from the players' subsequent behaviour as well as from direct observation of the game.
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In the common goby, Pomatoschistus microps, males compete aggressively for nest sites when these are in short supply. I investigated whether prior information about the opponent influenced aggression levels during nest competition. Males that, through a one-way mirror, had been observing two other males compete for a nest were placed together with either the winner or the loser. The new male pairs also received nest sites to compete for, and their behaviour was videorecorded. As a control, naïve males, which had not been watching the earlier contest, were also recorded when competing with previous winners or losers. Both bystanders and naïve males acquired the nest more often when competing with a loser than with a winner. The male that got the nest was generally more aggressive than his opponent. However, in trials with bystanders and losers, the male obtaining the nest was less aggressive than in contests between bystanders and winners and in contests between naïve males and losers. Furthermore, when bystanders competed with losers, they were less aggressive than were naïve males competing with losers. There was no difference in aggression between bystanders and naïve males competing with winners. Information about the opponent did not increase the probability of obtaining the nest, but the benefit, when applicable, might instead have consisted of a decrease in aggression, and thus a lowered energetic cost of nest competition.
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The processes that underlie the formation of the dominance hierarchy in a group are since long under debate. Models of self-organisation suggest that dominance hierarchies develop by the self-reinforcing effects of winning and losing fights (the so-called winner-loser effect), but according to 'the prior attribute hypothesis', dominance hierarchies develop from pre-existing individual differences, such as in body mass. In the present paper, we investigate the relevance of each of these two theories for the degree of female dominance over males. We investigate this in a correlative study in which we compare female dominance between groups of 22 species throughout the primate order. In our study female dominance may range from 0 (no female dominance) to 1 (complete female dominance). As regards 'the prior attribute hypothesis', we expected a negative correlation between female dominance over males and species-specific sexual dimorphism in body mass. However, to our surprise we found none (we use the method of independent contrasts). Instead, we confirm the self-organisation hypothesis: our model based on the winner-loser effect predicts that female dominance over males increases with the percentage of males in the group. We confirm this pattern at several levels in empirical data (among groups of a single species and between species of the same genus and of different ones). Since the winner-loser effect has been shown to work in many taxa including humans, these results may have broad implications.
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This paper studied the feeding rhythms of 5-, 8-, and 12-day old paradise fish (Macropodus opercularis) larvae under natural illumination, successive illumination, and successive dark. Under natural illumination, all the test larvae performed a typical daytime feeding rhythm, and fed actively during 12:00-16:00. However, under successive illumination and successive dark, the larvae did not show any identifiable feeding rhythm. Under successive illumination, the larvae fed actively all the day, and their feeding amount during 20:00-next 4:00 was significantly greater than that of the larvae under natural illumination. Conversely, the larvae under successive dark had a lower feeding intensity, particularly during 8:00-16:00, compared with the larvae under natural illumination. In conclusion, M. opercularis larvae performed a typical daytime feeding rhythm, and light condition greatly affected their feeding activity during their early life stages.
Book
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Gull chicks beg for food from their parents. Peacocks spread their tails to attract potential mates. Meerkats alert family members of the approach of predators. But are these--and other animals--sometimes dishonest? That's what William Searcy and Stephen Nowicki ask inThe Evolution of Animal Communication. They take on the fascinating yet perplexing question of the dependability of animal signaling systems. The book probes such phenomena as the begging of nesting birds, alarm calls in squirrels and primates, carotenoid coloration in fish and birds, the calls of frogs and toads, and weapon displays in crustaceans. Do these signals convey accurate information about the signaler, its future behavior, or its environment? Or do they mislead receivers in a way that benefits the signaler? For example, is the begging chick really hungry as its cries indicate or is it lobbying to get more food than its brothers and sisters? Searcy and Nowicki take on these and other questions by developing clear definitions of key issues, by reviewing the most relevant empirical data and game theory models available, and by asking how well theory matches data. They find that animal communication is largely reliable--but that this basic reliability also allows the clever deceiver to flourish. Well researched and clearly written, their book provides new insight into animal communication, behavior, and evolution.
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We report on a field study in which determinants of female breeding dispersal (i.e. the shift in the mean home range coordinates between successive breeding events) was investigated. Offspring were released in full sib groups (or half sib ones if there was within-clutch multiple paternity) at a separation distance from the females that varied between 'families'. This allowed for analysis of 'offspring nearness' effects on maternal dispersal. When a female's offspring were released more closely to her, she responded with greater dispersal. Furthermore, when the data set was truncated at 100 m maternal-offspring separation distance at offspring release (because perception at longer distances is likely to be unrealistic), maternal dispersal resulted in greater separation distance between female and offspring in the following year. A corresponding analysis for juveniles revealed no effect of maternal nearness on offspring dispersal but identified a significant effect of clutch size, to our surprise with dispersal declining with increasing clutch size. We discuss this result in a context of the 'public information hypothesis' (reinterpreted for juveniles in a nonsocial foraging species), suggesting that conspecific abundance perhaps acts as an indicator of local habitat quality. Thus, our analysis suggests a microgeographic structuring of the adult female population driven by genetic factors, either through inbreeding avoidance, or from simply avoiding individuals with a similar genotype regardless of their pedigree relatedness, while a nongenetic factor seems more important in their offspring.
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Animal communication takes place in a complex environment that is constantly modulated by particular social conditions. The majority of examples of signalling behaviours modulated by social context involve the presence of an individual of a particular sex or one that simply represents general competition. However, the identity of the individuals and the social relationships among individuals could also significantly modulate acoustic behaviour. In this study, I examined whether the presence of another male competitor modulates the post-interaction vocal response of a male subject to an oestrous female stimulus in golden hamsters, Mesocricetus auratus. I found that the presence of a potential competitor during an interaction with a female across a wire-mesh barrier significantly decreased the duration, tempo and energy of ‘post-female calls’ over time (experiment 1). Moreover, the call duration and energy of one-note simple calls changed over time depending on the identity of the stimulus male. Males that experienced social conflict and lost a fight maintained call duration and increased the energy of their calls over time, but only if the social interaction with the female was in the presence of another familiar neutral male and not in the presence of a familiar winner male (experiment 2). When the winner male stimulus was present, the duration and energy of the calls produced by the loser decreased with time. Individual recognition between familiar conspecifics with different shared experiences likely modulated the motivational state of the male subject and his vocal response after a social interaction with the female. This study provides new evidence that social complexity (competition and individual recognition) can induce dynamic changes of spectrotemporal features of hamster sexual ultrasonic vocalizations.
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The evolution of signals has mainly been considered in the context of an emitter-receiver dyadic interaction. However, communication usually occurs in the presence of individuals (an audience) that are not directly involved in the communication interaction, and it is more realistic to assume that signal evolution occurs in a network. Several types of information could be available to an audience, and, therefore, the presence of an audience could have effects on the behavior of the communicating animals and on signal evolution. We investigated whether the presence of an audience of conspecifics affected intrasexual aggressive communication in male fighting fish. We found that if the audience was a female, males increased the intensity of conspicuous displays that can be used in communication with both males and females and decreased highly aggressive displays that are solely directed to males. If the audience was a male of similar size, there was no significant change in the way in which males displayed. These results suggest that the presence of an audience could be one reason that many long-range and conspicuous signals are often shaped to transmit information to both males and females.
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Male singing behaviour correlates with extra-pair success in several passerine birds. Singing interactions during territorial contests provide relative information on the males involved. Such information may be important in female extra-pair behaviour and eavesdropping on singing interactions among males may allow females to make such relative assessments. We used interactive playback to instigate singing contests with male great tits during the peak fertile period of their mate in an attempt to alter females' assessment of mates' quality relative to neighbours (potential extra-pair partners). We escalated a contest to one male (by overlapping his songs) and then subsequently de-escalated a contest (by alternating) to a neighbour. Intrusions onto neighbouring territories by females mated to either treatment male were then monitored. Females mated to escalation treatment males were more likely to intrude following playbacks than females mated to de-escalation treatment males. Although the absolute song output of males did not differ between treatments, males produced more song relative to playback in de-escalation treatments and relative song output was positively correlated with female intrusions. Therefore, female great tits eavesdrop on singing interactions and change their visitation rates to neighbouring territories according to their mate's singing performance relative to neighbours.
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Male song reflects the quality of the singer in many animals and plays a role in female choice of social and copulation partners. Eavesdropping on male-male vocal interactions is a means by which females can compare different males' singing behavior directly and make immediate comparisons between
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Aggressive contests probably occur in networking environments where information about fighting ability is conveyed both to an opponent and to individuals peripheral to the fight itself, the bystanders. Our primary aim was to investigate the relative influences of eavesdropping and prior social experience on the dynamics of aggressive contests in Xiphophorus helleri. A bystander's ability to witness an encounter was manipulated using clear, one-way mirror, and opaque partitions. After watching (or not watching) the initial contest, the bystander encountered either the winner or loser of the bout. Treatment comparisons of bystander-winner or bystander-loser contest dynamics indicated the presence or absence of winner, loser, or eavesdropping effects. Winner and loser effects had negligible influences on bystander contest dynamics. Eavesdropping significantly reduced the bystander's propensity to initiate aggression, escalate, and win against seen winners regardless of whether the watched bout had escalated or not. Though eavesdropping had relatively little effect on bystander-loser contest dynamics, bystanders were less prone to initiate aggression and win against losers that had escalated in the witnessed bout. Thus, bystanders appear to preferentially retain and utilize information gained about potentially dangerous opponents (winners or persistent losers). Our data lend clear support for the importance of eavesdropping in visually based aggressive signalling systems.
Article
Recent models of signaling have assumed that the expenditure required to ensure detection of a display is negligible and have concentrated instead on the costs that may be necessary to maintain honesty. Such models predict that individuals who share the same interests are likely to communicate using "conspiratorial whispers," signals that are cheap and inconspicuous. Here, I present a game-theoretical model of signal detection (in a noisy environment, in the presence of potential eavesdroppers), which demonstrates that the idea of conspiratorial whispers is far too simplistic. It is true that in "cooperative" signaling systems (where signalers attempt to elicit responses that are beneficial for receivers), signal cost is not required to maintain honesty. However, some level of expenditure is still needed to ensure that a signal is reliably detected. Moreover, there exists a conflict of interest between signalers and receivers over the division of this expenditure. To predict the stable level of display in such cases, one needs to know how this conflict of interest will be resolved. The model reveals that the outcome may range from a whisper to a conspicuous and costly (though still conspiratorial) display. The more closely related the receiver is to the signaler, the greater the level of signal exaggeration that is expected-the opposite prediction to that of honest signaling models.
Article
Recent models of signaling have assumed that the expenditure required to ensure detection of a display is negligible and have concentrated instead on the costs that may be necessary to maintain honesty. Such models predict that individuals who share the same interests are likely to communicate using 'conspiratorial whispers,' signals that are cheap and inconspicuous. Here, I present a game-theoretical model of signal detection (in a noisy environment, in the presence of potential eavesdroppers), which demonstrates that the idea of conspiratorial whispers is far too simplistic. It is true that in 'cooperative' signaling systems (where signalers attempt to elicit responses that are beneficial for receivers), signal cost is not required to maintain honesty. However, some level of expenditure is still needed to ensure that a signal is reliably detected. Moreover, there exists a conflict of interest between signalers and receivers over the division of this expenditure. To predict the stable level of display in such cases, one needs to know how this conflict of interest will be resolved. The model reveals that the outcome may range from a whisper to a conspicuous and costly (though still conspiratorial) display. The more closely related the receiver is to the signaler, the greater the level of signal exaggeration that is expected-the opposite prediction to that of honest signaling models.
Article
Territorial systems are characterized by the relative longevity and stability of interactions between neighbouring individuals. Two abilities of signal receivers that can be seen as adaptations increasing the efficiency of territory defence will be discussed: identifying neighbouring individuals and ranging (i.e. determining the distance to) signallers. The costs involved in such discriminations will also be outlined. Although signalling has been traditionally considered as occurring between two individuals (signaller and receiver), long-range signals will be received by many individuals. In territorial systems a group of neighbours could be considered as a communication network: consisting as its simplest of a signaller and a number of receivers. The scope for low cost, low-risk information gathering in such networks by eavesdropping will be discussed with particular reference to territorial songbirds and electric fish.
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Communication in the natural environment often involves more than a simple sender-receiver dyad because signals may be detected by more than one individual (i.e. communication occurs in networks). The presence of individuals other than those involved in the signalling interaction has been shown to change signallers' behaviour. Previous experiments have shown that intra-sexual communication of male fighting fish (Betta splendens) is affected by the presence of a female but not by a male conspecific. However the experimental design did not allow the effect of the sex of the audience to be compared. We used an experimental design that allowed direct investigation of the effect of the sex of an audience on male-male fighting fish interactions. Our results show that the sex of a conspecific audience influences male-male aggressive displays. When a male audience was present subjects attempted significantly more bites and spent less time near the opponent than with a female audience. The results of this experiment support the view that the presence and sex of an audience is important in determining how individuals should display during an interaction.
Article
I. We recorded the responses of male Betta splendens when presented with various stimuli behind a transparent partition. The stimuli comprised i) live male and female B. splendens; ii) lifelike (motionless) models of male and female B. splendens; and iii) six stylised model B. splendens. The stylised models consisted of three pairs, one member of which had long male-like fins and raised opercula, and the other with short female-like fins and lowered opercula. The first pair had
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Despite the fact that most communication occurs in the context of networks of several individuals, the consequences of considering communication as a network on individuals' capacity for gathering information on congeners has been little investigated. Eavesdropping is the behaviour of a receiver extracting information from an interaction in which it is taking no part. Due to the fact that signals used in aggressive interactions are assumed to be reliable, eavesdropping could be an effective way of evaluating the quality of potential mates. We conducted two experiments designed to discover if female fighting fish (Betta splendens) monitor aggressive interactions between two males and if information gained by eavesdropping is used in the initial stages of subsequent mate choice. We found that females that had seen the interaction visited the winner first more often and spent significantly more time near, looking at and displaying to the winner of the interaction. By contrast females that had not seen the interaction visited the loser first more often and did not behave significantly differently to winner and loser. Overall these results are consistent with the idea that in the initial stages of mate choice females eavesdrop, i.e. use information gathered from male-male displays.
Article
Reproductive and agonistic behaviors in Siamese fighting fish were investigated in eight experiments, and some consequences and determinants of these sequences were isolated. First, fights and the formation of dominance-subordinancy relations were studied. Second, it was determined that large body size as well as males' prior residency in a tank produced an agonistic advantage; the magnitude of this advantage was positively related to the duration of residency. Third, the prior-residency effect in Bettas was determined by males' familiarity with visual and/or tactile cues in their home tanks. Fourth, dominant males had greater access to living space and were more likely to display at a mirror, build nests, and approach females than were subordinates. Finally, it was discovered that chemical cues associated with presumedly inert plastic tank dividers influence Bettas' social behavior.
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Zahavi's handicap theory, formalized by Grafen, suggests that 'cheaters' must be at a disadvantage if a communication system such as ritualized aggression is to evolve (Grafen 1991, In: Behavioural Ecology: An Evolutionary Approach (Ed. by J. R. Krebs & N. B. Davies), pp. 5-31. Oxford: Blackwell Scientific). To determine whether cheating is disadvantageous in Betta splendens, we held a series of live interactions, after inducing hyper-aggression by socially isolating and then briefly 'priming' the fish. Primed isolates, which were no stronger than their rivals, 'cheated' by escalating rapidly to tailbeating and biting. These cheaters, however, usually lost fights to non-isolated opponents. Unprimed isolates, i.e. socially isolated fish that were not primed, were not initially hyper-aggressive and thus did not cheat. They lost fewer fights than the cheaters. Results suggested that cheaters lost because they exhausted themselves by their hyper-aggressiveness, allowing their non-hyper-aggressive opponents to win. This result is consistent with the Zahavi-Grafen model of how an 'honest' level of ritualized aggression can be stabilized in a population. Copyright 1998 The Association for the Study of Animal Behaviour.
Article
During vocal interactions birds can time their song output so that their songs overlap those of a conspecific. Such overlapping is usually interpreted as a directed signal of arousal or as a signal of readiness to escalate contests. Inevitably, these interactions can often be heard by other conspecifics. To investigate if these conspecifics would perceive and use information derived from listening to others' interactions we conducted playback experiments in the field on male territorial nightingales, Luscinia megarhynchos. We tested whether asymmetric interactions, in which one bird overlaps the song of another individual, influences the behaviour of additional, passive conspecific receivers. To test the influence of song overlapping on a third individual, each subject received one playback treatment in which two intruders were simulated by a dual speaker playback design. Songs broadcast from one loudspeaker overlapped those from the other loudspeaker. Subjects responded more intensely at the side of the overlapper than at the side of the loudspeaker from which the song that was overlapped was broadcast. These differences in response persisted in a second test in which songs of only one of the formerly interacting rivals were played. The results suggest that, even if song overlapping is a signal directed towards the singer whose song is overlapped, this information is perceived and used by additional receivers. Such information on rivals' behaviour during an interaction might help an individual decide which strategies to adopt in possible future interactions with these conspecifics.Copyright 1997 The Association for the Study of Animal Behaviour
Article
Females of Photuris versicolor prey on males of other species by mimicking the flash responses of the prey's own females. They adjust their responses according to the male pattern, and attract males of four species with distinctively different flashed responses. The capabilities of the firefly brain are more complex than previously suspected. The mimicry is quite effective, and females seldom answered more than ten males without catching one.
Signalling in territorialsystems: a context for individual identi?ca-tion, ranging and eavesdropping Communication networks: social environments for receiving and signallingbehaviour
  • P K Mcgregor
  • R Soc
  • Lond
McGregor, P.K. (1993). Signalling in territorialsystems: a context for individual identi?ca-tion, ranging and eavesdropping.— R. Soc. Lond. B 340, p. 237-244. — — & Peake, T.M. (2000). Communication networks: social environments for receiving and signallingbehaviour.— Acta Ethol. 2, p. 71-81
Agnostic and reproductive interactions in Betta splendens Predictors of dominance in male Betta splendensOn thepredictability,sensitization,and habituationofaggressioninmale Bettas (Betta splendens)
  • P M Bronstein
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Bronstein, P.M. (1984). Agnostic and reproductive interactions in Betta splendens. — J. Comp. Psych. 98, p. 421-431. — — (1985). Predictors of dominance in male Betta splendens. — J. Comp. Psych. 99, p. 47-55. —— (1994).On thepredictability,sensitization,and habituationofaggressioninmale Bettas (Betta splendens). — J. Comp. Psych. 108, p. 45-57
Predictors of dominance in male Betta splendens
— — (1985). Predictors of dominance in male Betta splendens. — J. Comp. Psych. 99, p. 47-55.
On the predictability,sensitization,and habituationof aggression in male Bettas (Betta splendens)
(1994). On the predictability,sensitization,and habituationof aggression in male Bettas (Betta splendens).-J. Comp. Psych. 108, p. 45-57.
Principles of animal communication.-Sinauer Associates
  • J W Bradbury
  • S L Vehrencamp
Bradbury, J.W. & Vehrencamp, S.L. (1993). Principles of animal communication.-Sinauer Associates, Sunderland, MA.