Extended and cumulative effects of experimentally induced intergroup conflict in a cooperatively breeding mammal

Conflict between rival groups is rife in nature. While recent work has begun exploring the behavioural consequences of this intergroup conflict, studies have primarily considered just the 1–2 h immediately after single interactions with rivals or their cues. Using a habituated population of wild dwarf mongooses ( Helogale parvula ), we conducted week-long manipulations to investigate longer-term impacts of intergroup conflict. Compared to a single presentation of control herbivore faeces, one rival-group faecal presentation (simulating a territorial intrusion) resulted in more within-group grooming the following day, beyond the likely period of conflict-induced stress. Repeated presentations of outsider cues led to further changes in baseline behaviour by the end of the week: compared to control weeks, mongooses spent less time foraging and foraged closer to their groupmates, even when there had been no recent simulated intrusion. Moreover, there was more baseline territorial scent-marking and a higher likelihood of group fissioning in intrusion weeks. Consequently, individuals gained less body mass at the end of weeks with repeated simulated intrusions. Our experimental findings provide evidence for longer-term, extended and cumulative, effects of an elevated intergroup threat, which may lead to fitness consequences and underpin this powerful selective pressure.

Male White-shouldered Fairywrens (Malurus alboscapulatus) elevate testosterone when courting females but not during territorial challenges

Testosterone mediates suites of physical and behavioral traits across vertebrates, and circulation varies considerably across and within taxa. However, an understanding of the causal factors of variation in circulating testosterone has proven difficult despite decades of research. According to the challenge hypothesis, agonistic interactions between males immediately prior to the breeding season produce the highest levels of testosterone measured during this period. While many studies have provided support for this hypothesis, most species do not respond to male-male competition by elevating testosterone. As a result, a recent revision of the hypothesis (challenge hypothesis 2.0) places male-female interactions as the primary cause of rapid elevations in testosterone circulation in male vertebrates. Here, we offer a test of both iterations of the challenge hypothesis in a tropical bird species. We first illustrate that male White-shouldered Fairywrens (Malurus alboscapulatus) differ by subspecies in plasma testosterone concentrations. Then we use a social network approach to find that males of the subspecies with higher testosterone are characterized by greater social interaction scores, including more time aggregating to perform sexual displays. Next, we use a controlled experiment to test whether males respond to simulated territorial intrusion or courtship interaction contexts by elevating testosterone. Males sampled during courtship had greater plasma testosterone both relative to flushed controls and males sampled during simulated intrusion. Ultimately, our results are consistent with challenge hypothesis 2.0, as males rapidly elevated testosterone following interactions with females, but not during territorial challenges.

Testing the maintenance of natural responses to survival-relevant calls in the conservation breeding population of a critically endangered corvid (Corvus hawaiiensis)

Vocal communication serves an important role in driving animals' social interactions and ultimately their survival. However, natural vocal behavior can erode in human care. Determining if animals in conservation breeding programs exhibit and respond appropriately to species-specific vocalizations is therefore important for ensuring their survival post-release. We tested whether endangered 'alala (Corvus hawaiiensis), which are extinct in nature, have retained their natural responses to survival-relevant vocal calls. We conducted our studies on breeding populations derived from a small number of founding 'alala maintained in human care since their extinction in the wild in 2002. We presented pairs of 'alala with alarm, territorial intrusion, and two types of control playback calls (a non-threatening territorial maintenance call and a novel heterospecific call). 'Alala were significantly more likely to approach the speaker following alarm call playback than other call types, and were more likely to respond to territorial intrusion calls with the same aggressive territorial calls. Males were more likely to make these aggressive calls than females, mirroring their roles in territory defense. We also found individual consistency in the level of vocal behavior response across all call types, indicating that some individuals are more vocal than others. These results are encouraging, showing that 'alala exhibit relevant, species-specific behaviors despite generations under human care. It does illustrate, however, that all individuals do not respond uniformly, so vocal response may be an important factor to consider in determining the release suitability of individuals.

Territorial Responses of Nuthatches Sitta europaea—Evaluation of a Robot Model in a Simulated Territorial Intrusion

In birds, aggressive behavior can be elicited in the field with a simulated territory intrusion (STI). Here, we compared four different STI methods in nuthatches in the non-breeding season: playback only, playback combined with an inactive wooden model mounted on a robot device, playback and an active model mounted on a robot device, and playback with the robot device only. In the control treatment, birds were not exposed to STI. Experiments were carried out in 12 territories. Behavioral observations included latency to approach, latency to start calling, time spent in the same tree, number of flyovers, minimum distance, number of individuals, number of flights into the tree, and number of calls. There was no significant influence of stimulus presentation. Nuthatches responded equally to all four experimental manipulations, but order of presentation had a strong influence.

Defending as a unit: sex- and context-specific territorial defence in a duetting bird

Behaviours such as territorial defence represent functionally integrated traits that underlie multiple behavioural variables such as physical and acoustic responses. Characterizing the multivariate structure of such traits is fundamental to understand their evolution. In bird species that form stable pair bonds and are territorial year-round, both sexes are expected to defend their territory; however, the role that each sex plays in defending their shared territory remains largely unknown. Evidence for the sex-roles during territorial defence is mixed and sex- and context-specific characterizations of territorial defence embracing the multivariate nature of the trait are currently lacking. Here we investigated sex- and context-specific variation in a hypothesised latent variable called “territorial defence” and tested whether duets were part of territorial defence in a wild population of rufous hornero (Furnarius rufus). To do so, we combined a simulated territorial intrusion approach during nest building and provisioning contexts with a structural equation modelling approach. Our results showed that, in males and females, the six measured behavioural variables were linked by a single latent trait, territorial defence, in both contexts. Flights over the decoy and duet songs were equally good proxies of territorial defence. Although males were defending more the territory than females, pair members showed a positive correlation in their behaviour. The structural equation modelling framework enabled us to capture a complex correlation pattern among behavioural variables, expanding upon a classic body of research on territorial defence. Thus, the combination of classical behavioural approaches with sophisticated statistical analyses brings new exciting possibilities to the field of behavioural ecology.Significance statementTerritorial defence is a key behaviour in territorial species as it plays a major role in an individual’s reproductive success and survival. Additionally, territorial defence has been proposed as one possible evolutionary driver of duetting behaviour, one of the most fascinating vocal behaviours in birds. As behaviours are evolutionary characters, they must be studied in a multivariate framework. In this study we focused on characterizing territorial defence during a simulated territorial intrusion in an integrative manner using a classical territorial intrusion framework. We did so in male and female rufous horneros (Aves: Furnaridae) across two breeding contexts, while simultaneously testing theoretical predictions about the role of duetting behaviour as key part of territorial defence. Overall, our study provides for the first time a sex- and context-comparison of the multivariate, latent variable “territorial defence” in duetting birds, while highlighting the potential of combining field behavioural approaches with structural equation modelling.

Great tits in noisy territories and avoiding overlapping respond stronger to territorial intruders

Animals often communicate with each other in noisy environments where interference from the ambient noise and other signallers may reduce the effectiveness of signals. Signalling behaviours may also evolve to interfere with signals of their opponents, e.g. by temporally overlapping them with their own, such as the song overlapping behaviour that is seen in some songbirds during aggressive interactions. Song overlapping has been proposed to be a signal of aggressive intent, but few studies directly examined the association between song overlapping and aggressive behaviours of the overlapping bird (the predictive criterion). In the present paper we examined the question of whether song overlapping is correlated with aggressive behaviours displayed during a simulated territorial intrusion in a population of great tits (Parus major) living in an urban-rural gradient. We also examined whether aggressive behaviours are correlated with the ambient noise levels. We found that overlapping was associated negatively with aggressive behaviours males displayed against a simulated intruder. These results fail to support the predictive criterion for song overlapping, raising the question whether overlapping is in fact a signal of aggressive intent. Ambient noise levels were associated positively with aggressive behaviours but did not correlate with song rate, song duration or song overlapping. Great tits in noisy urban habitats may display higher levels of aggressive behaviours due to either interference of noise in aggressive communication or another indirect effect of noise.

Different Messages are Transmitted by Individual Duet Contributions and Complete Duets in a Species with Highly Overlapped Duets

Background: Duet function hypotheses have been mostly studied in bird species that produce duets with male and female solo songs. However, in order to understand if patterns of duet function are similar across all duetting species, it is highly necessary to test the duet function hypotheses in species that produce duets with vocalizations other than solo songs. Objective: We studied the responses of territorial pairs to each sex’s individual duet contribution and complete duets in a species that produces duets with a vocalization other than male and female solo songs. Methods: We conducted a playback experiment where we presented duet contributions of each sex to three populations of White-eared Ground-sparrows (Melozone leucotis) in Costa Rica, during this species’ breeding season in 2016. Results: The responses to complete duets were stronger than those to each sex’s duet contribution, suggesting that complete duets and each sex’s duet contribution have different functions. Complete duets are used to protect resources from intruders (supporting the resource defense hypothesis), and to prevent the partner from being usurped by intruders (supporting the mate-guarding hypothesis). Males used solo songs in response to female duet contributions, and this may work to attract intruder females (increasing the probability of extra-pair copulation). Males also use solo songs in response to male duet contributions, which may work as a signal to repel intruder males and guard their female. In this case, where mate attraction occurs with a completely different type of vocalization than used for duetting, we found a clear pattern of a double agenda for males when a territorial intrusion occurs. Conclusions: This study provides strong support for the dual function hypothesis in duets and reveals conflicting selective pressures between pair members relative to each hypothesis.