A history of susceptibility to parasites and divergence in solitary, gregarious and agonistic behaviors of embiopterans.

Two species of Haploembia Ramburi (Oligotomidae: Embioptera), nonnative detritivores found in the western USA, display solitary tendencies, not typical for webspinners that usually share silk galleries. Reports from the 1960s based on native populations in Italy highlighted the impact of a gregarine that depressed male sterility and female survivorship in Haploembia solieri (Rambur). Sympatric asexual Haploembia tarsalis (Ross) lives a normal lifespan when parasitized, albeit suffering from reduced fecundity. Our goal is to characterize behavioral repertoires as individuals interact and to develop methods for future investigations focused on the impact of the little-known parasite. We quantified individual responses to conspecifics or other species in 10-min dyadic interactions and, in a 3-day trial, determined whether they aggregated when given dispersed resources. Replicated groups of four adult female H. solieri or H. tarsalis settled away from each other over the 3-day trials. In 10-min bouts of same or different species pairs, focal insects bolted back, retreated and attempted to escape when they encountered one another, especially when the opponent was H. tarsalis. Males courted conspecific females, but were dramatically repelled by H. tarsalis. Serving as a positive control, Oligotoma nigra (Hagen) (Oligotomidae) adult females paired with conspecifics displayed typical webspinner behaviors by sitting together, sharing silk. Haploembia solieri males did not respond negatively to O. nigra, not known to be parasitized by the gregarine, but did so when paired with H. tarsalis. Results align with the prediction that susceptibility to parasitism may have led to antisocial behaviors observed in two Haploembia species.

Redefining dominance calculation: Increased competition flattens the dominance hierarchy in dairy cows.

Dominance hierarchies are known for mitigating conflicts and guiding priority of access to limited resources in gregarious animals. The dominance hierarchy of dairy cows is typically investigated using agonistic interactions, often monitored at the feed bunk right after fresh feed delivery when competition is high, resulting in frequent interactions. Yet, the outcome of agonistic interactions during times of high competition may be more influenced by cows' high valuation of fresh feed than their intrinsic attributes, such that the dominance hierarchy constructed using agonistic interactions under high versus low competition times might differ. We tested how the structure of the dominance hierarchy changes in relation to different levels of competition in a dynamic group of 48 lactating dairy cows over 10 mo, with 6 cows exchanged every 16 d, for a total of 159 cows. Using a validated algorithm, we continuously detected the actor and reactor of replacement behaviors in 30 feed bins as cows competed for feed. We also calculated the percentage of occupied feed bins to characterize competition at the moment of each replacement. These data were combined to create hierarchies using Elo ratings, separately for 25 occupancy levels ranging from 13% to 100%. For each 1% rise in feeder occupancy, hierarchy steepness fell by 2.41 × 10-3 ± 9.71 × 10-5 (SE), and the percentage of dyads where both cows replaced each other rose by 0.13% ± 0.01%. At the highest feeder occupancy level in comparison to the lowest one, we observed 7.57% more dyads in which the dominant individual (those that won more interactions at the lowest feeder occupancy) started to lose proportionally more. The magnitude of decrease in the winning rate of the dominant individual in those dyads also got amplified by 1.06 × 10-3% ± 1.37 × 10-4% (SE) for each 1% increase in feeder occupancy. These findings illustrate how inferred hierarchies vary with competition, with high competition flattening the hierarchy due to increased success of subordinate animals. We suggest that during heightened competition, increased valuation of resources can affect competitive success more than the individual's intrinsic dominance attributes. We recommend against calculating dominance hierarchies based on agonistic interactions during periods of high competition alone, and more generally urge researchers to differentiate agonistic interactions based on context when constructing dominance hierarchies.

Production of sounds by squirrelfish during symbiotic relationships with cleaner wrasses.

Examples of symbiotic relationships often include cleaning mutualisms, typically involving interactions between cleaner fish and other fish, called the clients. While these cleaners can cooperate by removing ectoparasites from their clients, they can also deceive by feeding on client mucus, a behavior usually referred to as "cheating behavior" that often leads to a discernible jolt from the client fish. Despite extensive studies of these interactions, most research has focused on the visual aspects of the communication. In this study, we aimed to explore the role of acoustic communication in the mutualistic relationship between cleaner fishes and nine holocentrid client species across four regions of the Indo-Pacific Ocean: French Polynesia, Guam, Seychelles, and the Philippines. Video cameras coupled with hydrophones were positioned at various locations on reefs housing Holocentridae fish to observe their acoustic behaviors during interactions. Our results indicate that all nine species of holocentrids can use acoustic signals to communicate to cleaner fish their refusal of the symbiotic interaction or their desire to terminate the cooperation. These sounds were predominantly observed during agonistic behavior and seem to support visual cues from the client. This study provides a novel example of acoustic communication during a symbiotic relationship in teleosts. Interestingly, these vocalizations often lacked a distinct pattern or structure. This contrasts with numerous other interspecific communication systems where clear and distinguishable signals are essential. This absence of a clear acoustic pattern may be because they are used in interspecific interactions to support visual behavior with no selective pressure for developing specific calls required in conspecific recognition. The different sound types produced could also be correlated with the severity of the client response. There is a need for further research into the effects of acoustic behaviors on the quality and dynamics of these mutualistic interactions.

Modelling animal contests based on spatio-temporal dynamics.

We present a general theoretical model for the spatio-temporal dynamics of animal contests. Inspired by interactions between physical particles, the model is formulated in terms of effective interaction potentials, which map typical elements of contest behaviour into empirically verifiable rules of contestant motion. This allows us to simulate the observable dynamics of contests in various realistic scenarios, notably in dyadic contests over a localized resource. Assessment strategies previously formulated in game-theoretic models, as well as the effects of fighting costs, can be described as variations in our model's parameters. Furthermore, the trends of contest duration associated with these assessment strategies can be derived and understood within the model. Detailed description of the contestants' motion enables the exploration of spatio-temporal properties of asymmetric contests, such as the emergence of chase dynamics. Overall, our framework aims to bridge the growing gap between empirical capabilities and theory in this widespread aspect of animal behaviour.

Effects of Positive Fighting Experience and Its Subsequent Deprivation on the Expression Profile of Mouse Hippocampal Genes Associated with Neurogenesis.

The hippocampus is known as the brain region implicated in visuospatial processes and processes associated with learning and short- and long-term memory. An important functional characteristic of the hippocampus is lifelong neurogenesis. A decrease or increase in adult hippocampal neurogenesis is associated with a wide range of neurological diseases. We have previously shown that in adult male mice with a chronic positive fighting experience in daily agonistic interactions, there is an increase in the proliferation of progenitor neurons and the production of young neurons in the dentate gyrus (in hippocampus), and these neurogenesis parameters remain modified during 2 weeks of deprivation of further fights. The aim of the present work was to identify hippocampal genes associated with neurogenesis and involved in the formation of behavioral features in mice with the chronic experience of wins in aggressive confrontations, as well as during the subsequent 2-week deprivation of agonistic interactions. Hippocampal gene expression profiles were compared among three groups of adult male mice: chronically winning for 20 days in the agonistic interactions, chronically victorious for 20 days followed by the 2-week deprivation of fights, and intact (control) mice. Neurogenesis-associated genes were identified whose transcription levels changed during the social confrontations and in the subsequent period of deprivation of fights. In the experimental males, some of these genes are associated with behavioral traits, including abnormal aggression-related behavior, an abnormal anxiety-related response, and others. Two genes encoding transcription factors (Nr1d1 and Fmr1) were likely to contribute the most to the between-group differences. It can be concluded that the chronic experience of wins in agonistic interactions alters hippocampal levels of transcription of multiple genes in adult male mice. The transcriptome changes get reversed only partially after the 2-week period of deprivation of fights. The identified differentially expressed genes associated with neurogenesis and involved in the control of a behavior/neurological phenotype can be used in further studies to identify targets for therapeutic correction of the neurological disturbances that develop in winners under the conditions of chronic social confrontations.

Acoustic signal dominance in the multimodal communication of a nocturnal mammal.

Multimodal communication in animals is common, and is particularly well studied in signals that include both visual and auditory components. Multimodal signals that combine acoustic and olfactory components are less well known. Multimodal communication plays a crucial role in agonistic interactions in many mammals, but relatively little is known about this type of communication in nocturnal mammals. Here, we used male Great Himalayan leaf-nosed bats Hipposideros armiger to investigate multimodal signal function in acoustic and olfactory aggressive displays. We monitored the physiological responses (heart rate [HR]) when H. armiger was presented with 1 of 3 stimuli: territorial calls, forehead gland odors, and bimodal signals (calls + odors). Results showed that H. armiger rapidly increased their HR when exposed to any of the 3 stimuli. However, the duration of elevated HR and magnitude of change in HR increased significantly more when acoustic stimuli were presented alone compared with the presentation of olfactory stimuli alone. In contrast, the duration of elevated HR and magnitude of change in HR were significantly higher with bimodal stimuli than with olfactory stimuli alone, but no significant differences were found between the HR response to acoustic and bimodal stimuli. Our previous work showed that acoustic and chemical signals provided different types of information; here we describe experiments investigating the responses to those signals. These results suggest that olfactory and acoustic signals are non-redundant signal components, and that the acoustic component is the dominant modality in male H. armiger, at least as it related to HR. This study provides the first evidence that acoustic signals dominate over olfactory signals during agonistic interactions in a nocturnal mammal.

Aggressive signaling among competing species of birds.

Aggressive interactions help individuals to gain access to and defend resources, but they can be costly, leading to increased predation risk, injury, or death. Signals involving sounds and color can allow birds to avoid the costs of intraspecific aggressive encounters, but we know less about agonistic signaling between species, where fights can be frequent and just as costly. Here, we review photographic and video evidence of aggressive interactions among species of birds (N = 337 interactions documenting the aggressive signals of 164 different bird species from 120 genera, 50 families, and 24 orders) to document how individuals signal in aggressive encounters among species, and explore whether these visual signals are similar to those used in aggressive encounters with conspecifics. Despite the diversity of birds examined, most aggressively signaling birds displayed weapons (bills, talons, wings) used in fighting and placed these weapons closest to their heterospecific opponent when signaling. Most species oriented their bodies and heads forward with their bills pointing towards their heterospecific opponent, often highlighting their face, throat, mouth, and bill. Many birds also opened their wings and/or tails, increasing their apparent size in displays, consistent with the importance of body size in determining behavioral dominance among species. Aggressive postures were often similar across species and taxonomic families. Exceptions included Accipitridae and Falconidae, which often highlighted their talons in the air, Columbidae, which often highlighted their underwings from the side, and Trochilidae, which often hovered upright in the air and pointed their fanned tail downward. Most species highlighted bright carotenoid-based colors in their signals, but highlighted colors varied across species and often involved multiple colors in combination (e.g., black, white, and carotenoid-based colors). Finally, birds tended to use the same visual signals in aggressive encounters with heterospecifics that they use in aggressive encounters with conspecifics, suggesting that selection from aggressive interactions may act on the same signaling traits regardless of competitor identity.

Dominance style and intersexual hierarchy in wild bonobos from Wamba.

Dominance hierarchies vary between species and possess particular characteristics depending on the distribution and abundance of food resources that affect the competitive regime. Bonobos have been described as having female intersexual dominance, based mainly on female coalitionary support against males, and more egalitarian hierarchies than chimpanzees. In this study, we tested whether female intersexual dominance is dependent on female coalitions or whether it still arises when only dyadic interactions are considered. We also examined the role of food abundance in shaping dominance style in a wild population of bonobos in Wamba, Democratic Republic of Congo. We found partial support concerning our first prediction in which we expected a male dominance over females when only dyadic agonistic interactions were considered because females were not systematically dominant over males, finding instead an intersexual codominance pattern. We failed to find support for our second prediction that hierarchies become more despotic under low fruit abundance, in fact, we found the opposite pattern. We discuss that codominance based on dyadic interactions in this group may arise as a consequence of male deference rather than females winning conflicts against males and that more despotic hierarchies during high fruit season may arise as a consequence of competition for high-quality resources or variation in party size.

Acoustic and visual adaptations to predation risk: a predator affects communication in vocal female fish.

Predation is an important ecological constraint that influences communication in animals. Fish respond to predators by adjusting their visual signaling behavior, but the responses in calling behavior in the presence of a visually detected predator are largely unknown. We hypothesize that fish will reduce visual and acoustic signaling including sound levels and avoid escalating fights in the presence of a predator. To test this we investigated dyadic contests in female croaking gouramis (Trichopsis vittata, Osphronemidae) in the presence and absence of a predator (Astronotus ocellatus, Cichlidae) in an adjoining tank. Agonistic behavior in T. vittata consists of lateral (visual) displays, antiparallel circling, and production of croaking sounds and may escalate to frontal displays. We analyzed the number and duration of lateral display bouts, the number, duration, sound pressure level, and dominant frequency of croaking sounds as well as contest outcomes. The number and duration of lateral displays decreased significantly in predator when compared with no-predator trials. Total number of sounds per contest dropped in parallel but no significant changes were observed in sound characteristics. In the presence of a predator, dyadic contests were decided or terminated during lateral displays and never escalated to frontal displays. The gouramis showed approaching behavior toward the predator between lateral displays. This is the first study supporting the hypothesis that predators reduce visual and acoustic signaling in a vocal fish. Sound properties, in contrast, did not change. Decreased signaling and the lack of escalating contests reduce the fish's conspicuousness and thus predation threat.

How feedback and feed-forward mechanisms link determinants of social dominance.

In many animal societies, individuals differ consistently in their ability to win agonistic interactions, resulting in dominance hierarchies. These differences arise due to a range of factors that can influence individuals' abilities to win agonistic interactions, spanning from genetically driven traits through to individuals' recent interaction history. Yet, despite a century of study since Schjelderup-Ebbe's seminal paper on social dominance, we still lack a general understanding of how these different factors work together to determine individuals' positions in hierarchies. Here, we first outline five widely studied factors that can influence interaction outcomes: intrinsic attributes, resource value asymmetry, winner-loser effects, dyadic interaction-outcome history and third-party support. A review of the evidence shows that a variety of factors are likely important to interaction outcomes, and thereby individuals' positions in dominance hierarchies, in diverse species. We propose that such factors are unlikely to determine dominance outcomes independently, but rather form part of feedback loops whereby the outcomes of previous agonistic interactions (e.g. access to food) impact factors that might be important in subsequent interactions (e.g. body condition). We provide a conceptual framework that illustrates the multitude potential routes through which such feedbacks can occur, and how the factors that determine the outcomes of dominance interactions are highly intertwined and thus rarely act independently of one another. Further, we generalise our framework to include multi-generational feed-forward mechanisms: how interaction outcomes in one generation can influence the factors determining interaction outcomes in the next generation via a range of parental effects. This general framework describes how interaction outcomes and the factors determining them are linked within generations via feedback loops, and between generations via feed-forward mechanisms. We then highlight methodological approaches that will facilitate the study of feedback loops and dominance dynamics. Lastly, we discuss how our framework could shape future research, including: how feedbacks generate variation in the factors discussed, and how this might be studied experimentally; how the relative importance of different feedback mechanisms varies across timescales; the role of social structure in modulating the effect of feedbacks on hierarchy structure and stability; and the routes of parental influence on the dominance status of offspring. Ultimately, by considering dominance interactions as part of a dynamic feedback system that also feeds forward into subsequent generations, we will understand better the factors that structure dominance hierarchies in animal groups.

Asymmetric competition over space use and territory between native brown trout (Salmo trutta) and invasive brook trout (Salvelinus fontinalis).

Interference competition over food and territory can shape population structure and habitat use within and between species. The introduction of invasive species often leads to novel competitive interactions over shared resources and invaders can eventually exclude the native species from preferred habitats. Invasive brook trout (Salvelinus fontinalis) introduced to northern Europe have excluded native brown trout (Salmo trutta) from numerous headwater streams. The fact that invasive brook trout can displace the more aggressive brown trout is puzzling. However, the earlier spawning and hatching of brook trout, compared to brown trout, may lead to unequal competition due to size advantage and prior resident status of brook trout at the fry stage. In this study, we examine the effect of competition between brown trout and brook trout using the natural size distribution of the two species. In two consecutive experiments, we first measured space use and feeding of a fry (age 0+) in the presence of a juvenile (age 1+). In experiment 2, we assessed territorial interactions between the species at the fry stage (age 0+) and if smaller brown trout could compensate the disadvantage by manipulating residence duration. Fry of brook trout feed sooner and spend more time close to the larger individual than brown trout fry. We also found that brook trout fry won most territorial contests against brown trout, and that increased residence duration led to longer and more aggressive interactions. The results suggest that smaller brown trout are displaced to suboptimal habitats in the presence of a larger brook trout. Therefore, the later emergence from gravel beds resulting in the naturally occurring size disadvantage of brown trout at the fry stage may lead to unequal territorial interactions that could explain why brown trout are displaced from preferred habitats in sympatry with brook trout.

Pelt Biting as a Practical Indicator of Social and Environment Stress in Farmed Red Deer.

Agonistic behavioural interactions play a decisive role in the competition for food, space, mating opportunities, and establishing social rank. We used pelt biting (number of bites on an animal's body) as a proxy for assessing the intensity of agonistic animal interactions and how it responded to social, population, and heat stress factors. We modelled a 14-year time series of pelt biting records and observational data of agonistic interactions on a population of captive Iberian red deer (Cervus elaphus). We found that (i) the higher the social rank of deer, the lower the number of pelt bites received; (ii) increasing heat stress conditions caused deer to suffer more pelt bites; (iii) males received more bites than females; (iv) the heavier the deer, the lower the number of bites on their bodies; (v) the bigger the group, the more bites exhibited on its members; (vi) deer 5-6 years old suffered greater rate of pelt biting than younger or older deer; and (vii) hinds that gave birth earlier in the parturition period suffered less pelt biting than those that gave birth around the peak of the parturition season (p < 0.01 for all effects). Pelt biting is useful to predict management situations in which deer welfare could be at stake.

Male-male behavioral interactions drive social-dominance-mediated differences in ejaculate traits.

Higher social status is expected to result in fitness benefits as it secures access to potential mates. In promiscuous species, male reproductive success is also determined by an individual's ability to compete for fertilization after mating by producing high-quality ejaculates. However, the complex relationship between a male's investment in social status and ejaculates remains unclear. Here, we examine how male social status influences ejaculate quality under a range of social contexts in the pygmy halfbeak Dermogenys collettei, a small, group-living, internally fertilizing freshwater fish. We show that male social status influences ejaculate traits, both in the presence and absence of females. Dominant males produced faster swimming and more viable sperm, two key determinants of ejaculate quality, but only under conditions with frequent male-male behavioral interactions. When male-male interactions were experimentally reduced through the addition of a refuge, differences in ejaculate traits of dominant and subordinate males disappeared. Furthermore, dominant males were in a better condition, growing faster, and possessing larger livers, highlighting a possible condition dependence of competitive traits. Contrary to expectations, female presence or absence did not affect sperm swimming speed or testes mass. Together, these results suggest a positive relationship between social status and ejaculate quality in halfbeaks and highlight that the strength of behavioral interactions between males is a key driver of social-status-dependent differences in ejaculate traits.

Gene Expression Changes in the Ventral Tegmental Area of Male Mice with Alternative Social Behavior Experience in Chronic Agonistic Interactions.

Daily agonistic interactions of mice are an effective experimental approach to elucidate the molecular mechanisms underlying the excitation of the brain neurons and the formation of alternative social behavior patterns. An RNA-Seq analysis was used to compare the ventral tegmental area (VTA) transcriptome profiles for three groups of male C57BL/6J mice: winners, a group of chronically winning mice, losers, a group of chronically defeated mice, and controls. The data obtained show that both winners and defeated mice experience stress, which however, has a more drastic effect on defeated animals causing more significant changes in the levels of gene transcription. Four genes (Nrgn, Ercc2, Otx2, and Six3) changed their VTA expression profiles in opposite directions in winners and defeated mice. It was first shown that Nrgn (neurogranin) expression was highly correlated with the expression of the genes involved in dopamine synthesis and transport (Th, Ddc, Slc6a3, and Drd2) in the VTA of defeated mice but not in winners. The obtained network of 31 coregulated genes, encoding proteins associated with nervous system development (including 24 genes associated with the generation of neurons), may be potentially useful for studying their role in the VTA dopaminergic neurons maturation under the influence of social stress.

Time of Grain Supplementation and Social Dominance Modify Feeding Behavior of Heifers in Rotational Grazing Systems.

Social hierarchy affects the access of animals to feed resources. On daily rotational pasture systems, supplementation time may influence feeding behavior. This trial was designed to test the effect of grain delivery time on the feeding behavior of heifers. Heifers divided into two groups according to breed (n = 15 Braford and n = 19 Jersey) were tested in a crossover design with two treatments: INITIAL-supplement at 8 am (entry time to a fresh paddock), and MIDDLE-supplement at 4 pm (middle time of paddock use). Animals entered a new paddock every morning, and grain supplement at 2 kg/animal/day was offered at the fence line (1 m/animal). Then, ingestive and other behaviors were registered by direct visual observation through scan sampling at 2-min intervals for 1 h after grain supply. Agonistic interactions were recorded continuously (instigator-victim) to build a social matrix whereby each heifer was defined as dominant, intermediate, or subordinate. Weekly pasture samples were collected according to the order that animals left the feeding area, using the hand-plucking technique, to determine crude protein and fiber content. Heifers spent more time grazing on the INITIAL treatment (p < 0.0001) but exhibited more behaviors on the MIDDLE treatment (p < 0.0001). Dominant heifers spent more time eating grain (p = 0.0008), whereas subordinate heifers spent more time grazing along the paddock (p = 0.0067), but not along the fence (p = 0.0008). The crude protein content of pasture samples was higher for the INITIAL treatment (p < 0.0001). Behavioral interaction occurred with respect to the order of leaving the feeding area, social rank, and crude protein consumed (p = 0.04). Subordinate heifers consistently grazed more and ate less grain supplement than dominant and intermediate heifers. However, when grain supplement was offered at the time animals entered the paddock, more grazing activity took place during supplement feeding, and subordinate heifers could select a high-protein diet. In the INITIAL treatment, this means that subordinate animals could benefit from the better pasture available, keeping a distance from dominant heifers, reducing agonistic interactions and likely improving their welfare.

Social network analysis in pigs: impacts of significant dyads on general network and centrality parameters.

In general, one animal is considered dominant over another animal if it has won more fights than its opponent. Whether this difference in won and lost fights is significant is neglected in most studies. Thus, the present study evaluates the impact of two different calculation methods for dyadic interactions with a significant asymmetric outcome on the results of social network analysis regarding agonistic interactions of pigs in three different mixing events (weaned piglets, fattening pigs and gilts). Directly after mixing, all animals were video recorded for 17 (fattening pigs, gilts) and 28 h (weaned piglets), documenting agonistic interactions. Two calculation methods for significant dyads, that is, dyadic interactions with a clear dominant subordinate relationship in which one animal has won significantly more fights than its encounter, were proposed: pen individual limits were calculated by a sign test considering the differences of won and lost fights of all dyadic interactions in each pen; dyad individual limits were determined by a one-sided sign test for each individual dyad. For all data sets (ALL, including all dyadic interactions; PEN or DYAD, including only significant dyads according to pen or dyad individual limits), networks were built based on the information of initiator and receiver with the pigs as nodes and the edges between them illustrating attacks. General network parameters describing the whole network structure and centrality parameters describing the position of each animal in the network were calculated. Both pen and dyad individual limits revealed only a small percentage of significant dyads for weaned piglets (12.4% or 8.8%), fattening pigs (4.2% or 0.6%) and gilts (3.6% or 0.4%). The comparison between the data sets revealed only high Spearman's rank correlation coefficients (rS) for the density, that is, percentage of possible edges that were actually present in the network, whereas the centrality parameters showed only moderate rS values (0.37 to 0.75). Thus, the rank order of the animals changed due to the exclusion of insignificant dyads, which shows that the results obtained from social network analysis are clearly influenced if insignificant dyads are excluded from the analyses. Due to the fact that the pen individual limits consider the overall level of agonistic interactions within each pen, this calculation method should be preferred over the dyad individual limits. Otherwise, too many animals in the group became isolated nodes with zero centrality for which no statement about their position within the network can be made.

Inter-group aggressive interaction patterns indicate male mate defense and female cooperation across bonobo groups at Wamba, Democratic Republic of the Congo.

OBJECTIVES: Although conflicts between groups over valuable resources are common in the animal kingdom, an individual's strategy toward out-group individuals may differ according to the benefits and costs received from inter-group interactions. Groups of bonobos encounter each other frequently and may mingle and range together from a few hours to a few days. During these inter-group associations, individuals across groups exhibit both aggressive and affiliative interactions. This study aimed to examine the strategies that bonobos employ with other groups, by comparing the patterns of within- and inter-group aggression. MATERIALS AND METHODS: We observed the aggressive interactions within a group of wild bonobos and between the group and three neighboring groups in Wamba, Luo Scientific Reserve, DR Congo. RESULTS: Bonobos increased the level of cooperation to attack out-group individuals more than they do to attack within-group individuals. Additionally, they reduced the aggression between within-group members during inter-group associations, compared to that when not associated with other groups. Males selectively and cooperatively attacked out-group males. Inter-group aggression among females was rare. Furthermore, females sometimes formed coalitions with out-group individuals to attack a common target. DISCUSSION: Our results support the hypothesis that inter-group competition occurs in bonobos, with males across groups competing over mates. Females across groups were tolerant and even cooperative with each other. Regardless of the ideal male strategy, female tolerant and cooperative relationships across groups and female within-group superiority over males could preserve tolerant inter-group relationships in bonobos.

What do we need to know to recognize a contest?

Animals frequently use agonistic contests as a way to solve disputes over indivisible resources. Such agonistic contests often represent interactions between an owner and a non-owner of a resource. However, some behaviors adopted by rivals during agonistic interactions are similar to behaviors adopted in other types of interactions. Thus, the possibility exists that some interactions between individuals can be misinterpreted as actual agonistic contests. Herein, we synthesize information from prior studies that present interactions that may be confounded with actual agonistic contests. We also point potential problems when different types of confoundment occur and provide suggestions of how to distinguish between agonistic contests and alternative interactions. For this, we made a distinction between completely non-agonistic interactions and quasi-agonistic interactions (i.e., interactions in which at least one rival is not motivated to fight). We also show potential biases in the understanding of how rivals decide who is the winner of a contest for studies that consider non- or quasi-agonistic interactions as actual agonistic contests.

A practical guide for inferring reliable dominance hierarchies and estimating their uncertainty.

Many animal social structures are organized hierarchically, with some individuals monopolizing resources. Dominance hierarchies have received great attention from behavioural and evolutionary ecologists. There are many methods for inferring hierarchies from social interactions. Yet, there are no clear guidelines about how many observed dominance interactions (i.e. sampling effort) are necessary for inferring reliable dominance hierarchies, nor are there any established tools for quantifying their uncertainty. We simulate interactions (winners and losers) in scenarios of varying steepness (the probability that a dominant defeats a subordinate based on their difference in rank). Using these data, we (1) quantify how the number of interactions recorded and the steepness of the hierarchy affect the performance of five methods for inferring hierarchies, (2) propose an amendment that improves the performance of a popular method, and (3) suggest two easy procedures to measure uncertainty and steepness in the inferred hierarchy. We find that the ratio of interactions to individuals required to infer reliable hierarchies is surprisingly low, but depends on the steepness of the hierarchy and the method used. We show that David's score and our novel randomized Elo-rating are the best methods when hierarchies are not extremely steep, where the original Elo-rating, the I&SI and the recently described ADAGIO perform less well. In addition, we show that two simple methods can be used to estimate uncertainty at the individual and group level, and that the randomized Elo-rating repeatability provides researchers with a standardized measure valid for comparing the steepness of different hierarchies. We provide several worked examples to guide researchers interested in studying dominance hierarchies. Methods for inferring dominance hierarchies are relatively robust. We recommend that a ratio of observed interactions to individuals of at least 10 (for steep hierarchies), and ideally 20 serves as a good benchmark. Our simple procedures for estimating uncertainty in the observed data will facilitate evaluating whether sufficient data have been collected, while plotting the shape of the hierarchy will provide new insights into the social structure of the study organism.

Appraisal of unimodal cues during agonistic interactions in Maylandia zebra.

Communication is essential during social interactions including animal conflicts and it is often a complex process involving multiple sensory channels or modalities. To better understand how different modalities interact during communication, it is fundamental to study the behavioural responses to both the composite multimodal signal and each unimodal component with adequate experimental protocols. Here we test how an African cichlid, which communicates with multiple senses, responds to different sensory stimuli in a social relevant scenario. We tested Maylandia zebra males with isolated chemical (urine or holding water coming both from dominant males), visual (real opponent or video playback) and acoustic (agonistic sounds) cues during agonistic interactions. We showed that (1) these fish relied mostly on the visual modality, showing increased aggressiveness in response to the sight of a real contestant but no responses to urine or agonistic sounds presented separately, (2) video playback in our study did not appear appropriate to test the visual modality and needs more technical prospecting, (3) holding water provoked territorial behaviours and seems to be promising for the investigation into the role of the chemical channel in this species. Our findings suggest that unimodal signals are non-redundant but how different sensory modalities interplay during communication remains largely unknown in fish.