TY - JOUR
AU - Dabelsteen,, Torben
AB - Abstract Territorial contests often occur in the presence of conspecifics not directly involved in the interaction. Actors may alter their behavior in the presence of this audience, an “audience effect,” and audiences themselves may alter their behavior as a result of observing an interaction, a “bystander effect.” Previous work has documented these effects by looking at each in isolation, but to our knowledge, none has investigated their interaction; something that is more likely to represent a realistic scenario for species where individuals aggregate spatially. We therefore have a somewhat limited understanding of the extent and direction of these potentially complex indirect social effects on behavior. Here, we examined how audience and bystander effects work in tandem to modify resident male aggressive behavior towards intruders in European fiddler crabs, Afruca tangeri. We found that male crabs with an audience showed greater aggressive behavior towards an intruder compared with males without an audience, but only if they had acted as a bystander to an aggressive signaling interaction prior to the intrusion. Indeed, bystanding during aggressive interactions elevated aggressive responses to intruders maximally if there was an audience present. Our results suggest that bystanding had a priming effect on territory-holding males, potentially by providing information on the immediate level of competition in the local neighborhood, and that same-sex audiences only matter if males have been primed. This study highlights the fundamental importance of considering broader interaction networks in studying real-world dyadic interactions and of including nonvertebrate taxonomic groups in these studies. BACKGROUND Considering communication in the context of a wider network has revealed behaviors and effects such as eavesdropping, audience effects, and bystander effects that would not be observable from a dyadic approach (McGregor 2005). Territorial contests are typically settled through pair-wise interactions within a network of multiple individuals, rather than in isolation (McGregor and Dabelsteen 1996). Audience effects occur when animals modify their behavior due to the presence of other individuals not involved in the interaction (Zuberbühler 2008) and these effects have been shown to alter the intensity of agonistic displays in a number of species (Fitzsimmons and Bertram 2013; Cruz and Oliveira 2015; Setoguchi et al. 2015; Montroy et al. 2016; dos Santos et al. 2017). In the context of territorial defense, the audience effect has been demonstrated to be dependent not only on the sex of the audience, but also on the territorial status and familiarity of individuals in the wider network (Dzieweczynski et al. 2005; Bertucci et al. 2014). Observed differences in behavior in the presence of an audience suggest that individuals can 1) assess attributes of their audience and 2) adjust their behavior as a strategy to counter costs (or strengthen benefits) that can come with eavesdropping, or more generally, gathering of social information by audiences (Earley and Dugatkin 2002). In the bystander effect, the audience members themselves are influenced by observing an interaction (Oliveira et al. 2001; Earley and Dugatkin 2002; Peake et al. 2006). The observation acts directly on the motivational system (Oliveira et al. 2001; Hirschenhauser and Oliveira 2006) and prepares individuals for what may happen next in their social environment (Antunes and Oliveira 2009). Individuals can thus be primed to augment their levels of aggression in interactions following bystanding. For example, Clotfelter and Paolino (Clotfelter and Paolino 2003) found increased aggressiveness by Siamese fighting fish, Betta splendens, towards a novel male conspecific after the observation of an aggressive interaction. However, reverse priming, a reduction in aggressive behavior, has also been found, for example, in a study of crayfish, Orconectes rusticus (Zulandt et al. 2008), which to our knowledge is also the only invertebrate species in which bystander effects have been investigated. Although audience and bystander effects are frequently documented, it is not clear from the literature how bystander and audience effects may interact. This represents a potential key gap in our understanding of the strength or importance of these effects in in situ contexts where they are likely to happen concurrently, particularly in species where individuals live in aggregated communities. A study in Siamese fighting fish, suggests that simply viewing an unfamiliar male prior to a contest with another male can prime males for increased aggression during a fight, irrespective of whether the audience is present or not during the fight itself (Matos et al. 2003). It also highlights the complexity of bystander and audience effects and the need to manipulate these effects in tandem in order to deepen our understanding of the significance of dynamics in the social environment for the expression of behavior. Here, we investigate the interplay between audience and bystander effects on the territorial behavior of male European fiddler crabs, Afruca tangeri. Fiddler crabs are very unlikely to be found in isolation (Pope 2005), and therefore have a high likelihood of engaging in social interactions with multiple receivers within signal range. Males actively defend territories around their burrows (Hemmi and Zeil 2003) with their one greatly enlarged claw, used for, among other things, the production of highly conspicuous visual agonistic and courtship signals and for fighting (Wolfrath 1993; Oliveira and Custódio 1998). Male crabs without a burrow will wander through the population and challenge burrow holders in attempts to acquire a burrow (Jordao and Oliveira 2005). Burrow-holding males must therefore invest in an agonistic response in order to retain residency (Oliveira and Custódio 1998). Recent work in another fiddler crab species suggests that same-sex audiences do not matter for expressed levels of aggression (dos Santos et al. 2017). Given the ubiquity of males in the immediate social environment (i.e., males are highly likely to have male neighbors), this is perhaps not surprising. We propose instead that males will fine-tune their responsiveness to their social environment according to current social information, such as the immediate level of competition in the neighborhood. If males observe a territorial dispute in their neighbor’s territory, they should be primed for a possible challenge of their own territory and therefore will be more responsive or sensitive to their immediate social environment. In this case, bystanding to an aggressive interaction between a neighboring male and a stranger (intruding male), should augment the aggressive response exhibited by males towards intruders appearing close in time. This should be particularly so when the social stimuli are increased by the presence of a male audience because this audience can gather information on the interaction outcome (e.g., the competitive ability of his male neighbor) or, in the least, can act as an indicator of a more competitive social environment (i.e., more males are present) compared to when no male audience is present. In this investigation, we used an in situ experimental manipulation of the social environment that burrow-holding male crabs experienced to test such effects during territorial confrontations in the home environment. METHODS The study was carried out in the Parque Natural da Ria Formosa, Portugal (N370927, E073244) from May to July 2012 in order to quantify audience and bystander effects in situ in males of a free roaming population of European fiddler crabs. To test focal male crabs at their home burrow, an arena made of bamboo and sand-colored fabric was placed around 2 neighboring males and their burrows (Figure 1). Neighboring male pairs were selected according to 3 criteria: 1) they were matched in claw size (estimated by visual comparison); 2) they were exhibiting courtship behavior (courtship waving) indicating that they were actively defending a burrow; and 3) they were within 1 m of one another (measured using a measuring tape stretched in a straight line from the center of one burrow to the other). Stimulus crabs captured from other areas of the mudflat were tethered with 10 cm of clear monofilament line to bamboo posts inserted into the substrate within the arena to simulate wandering male crabs (intruders). This method of tethering stimulus crabs is relatively standard and has been used successfully in this and several other fiddler crab species to elicit both courtship and territorial behavior as observed under natural conditions (Pope 2005; Detto et al. 2006; Reaney 2007; How et al. 2008; Detto and Backwell 2009; Detto et al. 2010; Milner et al. 2010; Booksmythe et al. 2010). Intruders were matched in claw size to focal crabs using visual comparison. Focal crabs were exposed to two 5-min phases: a “bystanding phase” (neighbor interacts with a simulated intruder) and a subsequent “interaction phase” (focal male interacts with a simulated intruder) in 1 of 4 treatments (3 control and 1 experimental): (1) null control—no neighbor–intruder interaction in the “bystanding phase” and no audience in the “interaction phase” (n = 11); (2) audience control—no neighbor–intruder interaction in the “bystanding phase” and an audience in the “interaction phase” (n = 10); (3) bystander control—neighbor–intruder interaction in the “bystanding phase” and no audience in the “interaction phase” (n = 11); and (4) bystander and audience (experimental)—neighbor–intruder interaction in the “bystanding phase” and audience in the “interaction phase” (n = 11) (Figure 1). Figure 1 View largeDownload slide Overview of experimental arenas and 4 treatments (3 control and 1 experimental) used to investigate bystander and audience effects in male European fiddler crabs (see text for details). Grey circles denote a burrow entrance. Figure 1 View largeDownload slide Overview of experimental arenas and 4 treatments (3 control and 1 experimental) used to investigate bystander and audience effects in male European fiddler crabs (see text for details). Grey circles denote a burrow entrance. In the “bystanding phase” of treatments (3) and (4), a stimulus male was tethered 40 cm from the neighbor’s burrow (minimum 90 cm from the focal male’s burrow) and left for 5 min after both males (focal and neighbor) had emerged from their burrows. In the “bystanding phase” of treatments (1) and (2), the arena was approached (and stimulus male placement simulated) and the crabs were then left for 5 min after both males had emerged from their burrow. In the interaction phase, a stimulus male was tethered 40 cm from the focal male’s burrow to simulate a wandering male. In treatments (2) and (4), the neighboring male was allowed to emerge from his burrow and act as an audience, whereas in treatments (1) and (3), the neighboring male’s burrow was blocked to prevent him from emerging during the trial period. Phases began when the focal or both males had surfaced (carapace and major claw visible), as applicable, and lasted for 5 min. Following a trial, burrows were marked with a small flag and within a semilunar tidal cycle these areas were avoided for further testing to ensure that males were not reused over the course of the study. We quantified aggression as a behavioral state by measuring the duration of time that focal males were engaged in aggressive behaviors toward the simulated intruder in the 5-min interaction phase. The observed aggressive behavior, following previously published methods (see Wolfrath 1993; Oliveira et al. 1998; Burford et al. 2000), included both noncontact aggression (threat displays) and that involving physical contact (pushing, grappling, and tossing) and the intensity of enactment represents the focal male’s willingness to escalate the contest (Wolfrath 1993; Oliveira et al. 1998). We compared the time focal males spent performing aggressive behavior among treatments with an analysis of variance with treatment as a fixed effect (SPSS v. 22). We included interburrow distance between the focal and neighbor as a covariate in the model as this varied across focal individuals. Post hoc analyses were carried out on the marginal means using a least significant difference adjustment for multiple comparisons. All behaviors were scored from video recordings (Panasonic HDC-SD800 camcorder) of the trials by a single observer (M.K.M.) naive to the trial condition. The study was carried out under permit (ICNF, Portugal) following ethical approval from the first author’s home institution. All captured crabs were kept singly in shaded containers filled with sea water and mud prior to testing and released back to their area of capture at the end of a trial. Crabs were marked with nontoxic paint to ensure that they were not used more than once in case of recapture. RESULTS There was an overall effect of the treatment that focal males experienced on the amount of aggressive behavior they displayed toward a simulated intruder (F3,38 = 11.797, P < 0.0001, Figure 2). Our post hoc analysis revealed that there was an effect of having a neighboring male audience present during a territorial intrusion on the level of aggression expressed by a burrow-holding male, but only if burrow-holding males had previously acted as bystander (Treatment 4) to a neighboring aggressive interaction (Table 1; Figure 2). In short, males in Treatment 4 behaved more aggressively than in any other treatment. Males in 2 of the control treatments (Treatments 1 and 2) did not differ from one another in behavior, but males in the null control (no bystanding and no audience; Treatment 1), were less aggressive than males in the bystander control (bystanding, no audience) (Treatment 3). Table 1 Results of the post hoc analysis of differences among treatments in burrow-holding male aggressive responses to simulated intrusions by conspecific male fiddler crabs 95% CI for Difference Treatment A Treatment B Mean difference Lower bound Upper bound P 1 - Null control 2 - Audience control −31.04 −82.41 20.33 0.229 3 - Bystander −53.73 −103.39 −4.07 0.035 4 - Bystander and audience −143.18 −194.50 −91.85 <0.001 2 - Audience control 3 - Bystander −22.69 −73.91 28.53 0.375 4 - Bystander and audience −112.14 −163.35 −60.92 <0.001 3 - Bystander 4 - Bystander and audience −89.45 −140.48 −38.42 0.001 95% CI for Difference Treatment A Treatment B Mean difference Lower bound Upper bound P 1 - Null control 2 - Audience control −31.04 −82.41 20.33 0.229 3 - Bystander −53.73 −103.39 −4.07 0.035 4 - Bystander and audience −143.18 −194.50 −91.85 <0.001 2 - Audience control 3 - Bystander −22.69 −73.91 28.53 0.375 4 - Bystander and audience −112.14 −163.35 −60.92 <0.001 3 - Bystander 4 - Bystander and audience −89.45 −140.48 −38.42 0.001 View Large Table 1 Results of the post hoc analysis of differences among treatments in burrow-holding male aggressive responses to simulated intrusions by conspecific male fiddler crabs 95% CI for Difference Treatment A Treatment B Mean difference Lower bound Upper bound P 1 - Null control 2 - Audience control −31.04 −82.41 20.33 0.229 3 - Bystander −53.73 −103.39 −4.07 0.035 4 - Bystander and audience −143.18 −194.50 −91.85 <0.001 2 - Audience control 3 - Bystander −22.69 −73.91 28.53 0.375 4 - Bystander and audience −112.14 −163.35 −60.92 <0.001 3 - Bystander 4 - Bystander and audience −89.45 −140.48 −38.42 0.001 95% CI for Difference Treatment A Treatment B Mean difference Lower bound Upper bound P 1 - Null control 2 - Audience control −31.04 −82.41 20.33 0.229 3 - Bystander −53.73 −103.39 −4.07 0.035 4 - Bystander and audience −143.18 −194.50 −91.85 <0.001 2 - Audience control 3 - Bystander −22.69 −73.91 28.53 0.375 4 - Bystander and audience −112.14 −163.35 −60.92 <0.001 3 - Bystander 4 - Bystander and audience −89.45 −140.48 −38.42 0.001 View Large Figure 2 View largeDownload slide Average time that male European fiddler crabs spent engaging in aggressive behavior toward experimentally introduced male intruders in 4 social treatments (see Figure 1) designed to test for bystander and audience effects (Estimated Marginal Mean [EMM] ±SE; matching letters indicate treatments where P > 0.05 for tested differences). Figure 2 View largeDownload slide Average time that male European fiddler crabs spent engaging in aggressive behavior toward experimentally introduced male intruders in 4 social treatments (see Figure 1) designed to test for bystander and audience effects (Estimated Marginal Mean [EMM] ±SE; matching letters indicate treatments where P > 0.05 for tested differences). DISCUSSION Gleaning information from the social environment is essential for an individual to respond to this environment appropriately (Danchin et al. 2004; Seppanen et al. 2007; Valone 2007). Individuals living in systems that are likely to be high in social noise will be under particular pressure to either filter out information to avoid an inappropriate response or to use specific cues to alert them to a situation where an active response may be required (e.g., Naguib et al. 2004; Fitzsimmons et al. 2008). This latter priming effect may be particularly important in predicting that, for example, a territorial challenge is likely to occur. The anticipation of being challenged is likely to affect not only how individuals respond to a rival conspecific during a direct interaction, but also how they respond to the presence of other conspecifics in the immediate environment. These individuals that are not part of the interaction may themselves be gleaning social information (i.e., they are conspecific audiences). In this study, we found evidence that a neighboring male audience moderated male European fiddler crab behavior towards an intruder, but only if males had acted as bystander to an aggressive interaction between a male neighbor and a same-sex intruder immediately prior to the encounter. This suggests that focal males that had viewed an aggressive interaction between his neighbor and an intruder were somehow primed for heightened responsiveness to having a male audience present during an interaction between himself and a territorial intruder. The observed increase in aggressiveness was likely also a function of a general priming for an aggressive response as a result of the bystanding, since we did detect greater aggressiveness when males without an audience had acted as bystander compared with males in our null control (no audience and no bystanding). The prevalence of the use of public information (Danchin et al. 2004) suggests that it could be beneficial for individuals to employ strategies to manage the perceptions of unintended receivers. Fighting could communicate the focal male’s motivation to defend his territory (Detto et al. 2010) or ability to do so (Peake et al. 2001) to his neighbor. However, we did not see an effect of a neighboring male audience unless the focal male had previously acted as a bystander to an aggressive interaction between his neighbor and an intruder. Bio-regulatory mediators of the adjustment of aggressive behavior are likely to be affected in males that anticipate a territorial challenge based on cues in their social environment; facilitating an adjustment to an increased competitive environment with the appropriate behavioral response (Antunes and Oliveira 2009). It could be that the simple presence of a neighbor is not enough of a cue of the extent of the competitive environment, given the ubiquity of territorial neighbors, and their familiarity, in a male’s visual field (Hemmi and Zeil 2003; Pope 2005; Detto et al. 2010). Instead, if a territorial challenge has been observed immediately prior to an intrusion, effects on bio-regulatory mechanisms (Oliveira et al. 2001) could drive a response directly or at least provide a cue as to the probability of an escalation of aggression that could lead to a territorial takeover (Oliveira 2009). The bystander effect detected in this study is consistent with work in vertebrate species demonstrating increased levels of aggression (e.g., Clotfelter and Paolino 2003), but inconsistent with the reverse priming demonstrated in another crustacean species (Zulandt et al. 2008). In our study, burrow-holding males will have invested energy into establishing territory boundaries with their neighbors (Detto et al. 2010) and as such, intruders are a potential threat to the territory more generally if they were to take over a neighbor’s burrow. Zulandt and colleagues (Zulandt et al. 2008) suggest that the bystander effect could be resource dependent such that when resources are abundant, and consequently their value reduced, an individual will avoid escalation if the observation of a fight indicates that there is an increased likelihood of being challenged within their network. In the present study, males were at their home burrow and as a consequence were defending a valuable resource; an added ecological realism that may have contributed to the direction of the effect. Our study demonstrates that there is likely to be a complex interaction between audience and bystander effects in systems where individuals are part of a network of conspecific interactants. The results suggest that selection has enhanced the use of public information to increase the appropriateness of responses to social stimuli. We investigated the effects of the presence of a fiddler crab’s nearest neighbor, an individual that the focal is likely to have interacted with before and to be familiar with, however, laboratory work on audience effects has demonstrated that attributes of an audience can influence physiological and behavioral effects on individuals, for example, sex and territorial status (Dzieweczynski et al. 2005). Future in situ work investigating behavioral reactions when the audience is a non-neighbor or a female would therefore be particularly enlightening. FUNDING The work was supported by the Danish Council for Independent Research/Natural Sciences (10-084844 and DFF – 1323-00105) to T.D. and S.K.D. and a Leverhulme Trust Early Careers Fellowship to S.K.D. (ECF/2010/0672). Acknowledgments We give special thanks to E. Vžesniauskaitė for assistance in the field. Data accessibility: Analyses reported in this article can be reproduced using the data provided by Darden and colleagues (2018). 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TI - Territorial defense in a network: audiences only matter to male fiddler crabs primed for confrontation
JF - Behavioral Ecology
DO - 10.1093/beheco/ary169
DA - 2019-04-05
UR - https://www.deepdyve.com/lp/oxford-university-press/territorial-defense-in-a-network-audiences-only-matter-to-male-fiddler-uKm0sEXIP2
SP - 336
VL - 30
IS - 2
DP - DeepDyve
ER -