Dominance, Status Signals and Coloration in Male Mandrills (Mandrillus sphinx)

Where individuals contest access to a resource, escalated physical fighting presents a risk to all involved. The requirement for mechanisms of conflict management has led to the evolution of a variety of decision rules and signals that act to reduce the frequency of aggression during competitive encounters. We examined strategies of conflict management in male mandrills (Mandrillus sphinx) living in two semi‐free‐ranging groups in Gabon. Adult male mandrills are large (31 kg), with long canines, making the costs of conflict potentially very high. We found that males formed dominance hierarchies, but that male–male relationships were characterized by avoidance, appeasement and ignoring. Fights were rare, but could result in death. Examination of the relationship between dominance and signaling showed that males use facial and gestural signals to communicate dominance and subordinance, avoiding escalated conflict. Male mandrills also possess rank‐dependent red coloration on the face, rump and genitalia, and we examined the hypothesis that this coloration acts as a ‘badge of status’, communicating male fighting ability to other males. If this is the case, then similarity in color should lead to higher dyadic rates of aggression, while males that differ markedly should resolve encounters quickly, with the paler individual retreating. Indeed, appeasement (the ‘grin’ display), threats, fights and tense ‘stand‐off’ encounters were significantly more frequent between similarly colored males, while clear submission was more frequent where color differences were large. We conclude that male mandrills employ both formal behavioral indicators of dominance and of subordination, and may also use relative brightness of red coloration to facilitate the assessment of individual differences in fighting ability, thereby regulating the degree of costly, escalated conflict between well‐armed males.     

Repeated Recent Aggressive Encounters Do Not Affect Behavioral Consistency in Male Siamese Fighting Fish

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.     

Morphological and Behavioural Traits Affecting the Intensity and Outcome of Male Contests in Gallotia galloti galloti (Family Lacertidae)

This study describes the sequence of behaviour during aggressive encounters between male ‘tizón’ lizards (G. galloti galloti) and assesses the effect of morphological and behavioural traits on the outcome and intensity of staged aggressive encounters between males. Aggressive encounters ranged from only throat extension to escalated fights with biting and rolling over. Winners were heavier, had longer heads, and performed tongue-flicking, throat extension and biting at a higher rate than losers. The rate of aggressive behaviour increased with decreasing difference in snout-to-vent length, head length and head width of the contestants. The results are in agreement with some predictions of the sequential assessment game model in that probability of victory increased with the difference in fighting ability and that the rate of aggressive behaviour was higher in contests between animals of similar size.     

Self assessment in insects: honeybee queens know their own strength

Contests mediate access to reproductive opportunities in almost all species of animals. An important aspect of the evolution of contests is the reduction of the costs incurred during intra-specific encounters to a minimum. However, escalated fights are commonly lethal in some species like the honeybee, Apis mellifera. By experimentally reducing honeybee queens' fighting abilities, we demonstrate that they refrain from engaging in lethal contests that typically characterize their reproductive dominance behavior and coexist peacefully within a colony. This suggests that weak queens exploit an alternative reproductive strategy and provides an explanation for rare occurrences of queen cohabitation in nature. Our results further indicate that self-assessment, but not mutual assessment of fighting ability occurs prior to and during the agonistic encounters.     

Is the parallel walk between competing male fallow deer, Dama dama, a lateral display of individual quality?

During competitive encounters protagonists are expected to use signals of individual quality particularly if there is a risk of injury or death. Lateral presentation of body profile, by which information regarding phenotypic characteristics associated with individual quality are displayed, may represent such a strategy. During aggressive interactions, male fallow deer frequently engage in parallel walking which is assumed to represent a mutual display of quality, as mediated by exposure of the maximal profile of the body or antlers. We examined the context and role of the parallel walk during competitive encounters to investigate whether there was evidence that dyads of competing males were assessing differences in phenotypic characteristics. There was no evidence to support the hypotheses that the parallel walk is a lateral display of body size or weaponry or that its use is associated with a reduced level of escalated or risky behaviours during fighting. Total time spent fighting was not shorter when a parallel walk was present than when there was no parallel walk. The parallel walk was highly associated with fighting and it was more likely to be initiated by the subsequent loser. Furthermore, parallel walking frequently followed bouts of fighting and as such may represent a strategy that permits an animal the opportunity to decide whether to continue fighting. Parallel walking was also associated with a failure to resolve contests in favour of one animal indicating that it may be a means of withdrawing from further fighting without incurring a loss in dominance status. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Heritable variation for territorial success in two Drosophila melanogaster populations

Male fruitflies from stocks from two localities were artificially selected for defence of a food area against intruding males. Both stocks showed a rapid replicable response to selection over a few generations, indicating considerable genetic variation for territorial success in the base populations. Crosses between lines indicate directional dominance for increased territorial success and no maternal or paternal effects. Selected males escalated relatively more frequently against territory residents than control males, and won relatively more escalated encounters. There was no correlated response in body weight. In the presence of territories, selected males had a higher mating success with inseminated females than control males, but did not differ in mating speed. Indirect selection for territorial success was carried out by allowing flies to mate in the presence of different types of resources. After 21–26 generations, males from lines held in cages with discrete resources where territories could be established had greater territorial success than males from lines held in cages with one large resource. This genetic divergence in response to resource type is consistent with the heritable variation in territorial success and mating success of territorial males.     

Mating Behavior of Daphnia: Impacts of Predation Risk,Food Quantity,and Reproductive Phase of Females

High predation risk and food depletion lead to sexual reproduction in cyclically parthenogenetic Daphnia. Mating, the core of sexual reproduction, also occurs under these conditions. Assessment of the environmental conditions and alteration of mating efforts may aid in determining the success of sexual reproduction. Here, we evaluated the impacts of predation risk, food quantity, and reproductive phase of females on the mating behavior of Daphnia obtusa males including contact frequency and duration using video analysis. Mating–related behavior involved male–female contact (mating) as well as male–male contact (fighting). Mating frequency increased while unnecessary fighting decreased in the presence of predation risk. In addition, low food concentration reduced fighting between males. Males attempted to attach to sexual females more than asexual females, and fighting occurred more frequently in the presence of sexual females. Duration of mating was relatively long; however, males separated shortly after contact in terms of fighting behavior. Thus, assessment of environmental factors and primary sexing of mates were performed before actual contact, possibly mechanically, and precise sex discrimination was conducted after contact. These results suggest that mating in Daphnia is not a random process but rather a balance between predation risk and energetic cost that results in changes in mating and fighting strategies.     

Acoustic Dueling in Tarbush Grasshoppers: Settlement of Territorial Contests via Alternation of Reliable Signals

A field study of disputes over ownership of mating territories by male tarbush grasshoppers (Ligurotettix planum) revealed that most contests were settled ritualistically via the antiphonal exchange of acoustic signals. Males used a special aggressive signal, referred to as the “shuck” call, in these encounters. Individuals that never produced shuck calls invariably departed the contested sites, and a playback experiment showed that shuck calls elicited higher response levels from males than sexual advertisement calls. Approximately 1/5 of all territorial contests escalated to biting, grappling, or kicking. The outcomes of territorial contests that were settled via purely acoustic encounters could not be predicted by the size of the participants. Prior residency was a predictor of winning in some contests, but often the competing males did not possess clear intruder/resident roles. However, a signal parameter that was a combined measure of the rate of shuck calls and their mean length predicted the winner of most acoustic encounters. Contests that escalated, though, were generally distinguished by participants that displayed comparable measures of all signal parameters save call length. A tradeoff occurred between call rate and length in most individuals. This, as well as the finding that call length did predict winning in escalated contests, implies that these signal parameters may reflect an individual's strength or motivation. Contests in which the signal parameters of the males were similar tended to be prolonged and escalated. This relationship suggests that individuals assess the signals of their rivals and depart if they cannot match their call rate and length. That both call rate and length are assessed may enhance the reliability with which signals can be used to predict a rival's fighting ability in the event of escalation. Various simple mechanisms for signal assessment are proposed. A second playback experiment suggested that the insects heard less effectively while they called. Timing mechanisms that preclude the overlap of calls during contests occur, and they may have evolved because of the importance of precise assessment of the signals of rivals. Such mechanisms circumvent the potential impairment in perception noted above and also generate a conspicuous feature of L. planum contests — mutual alternation of signals.     

Always Gamble on an Empty Stomach: Hunger Is Associated with Advantageous Decision Making

Three experimental studies examined the counterintuitive hypothesis that hunger improves strategic decision making, arguing that people in a hot state are better able to make favorable decisions involving uncertain outcomes. Studies 1 and 2 demonstrated that participants with more hunger or greater appetite made more advantageous choices in the Iowa Gambling Task compared to sated participants or participants with a smaller appetite. Study 3 revealed that hungry participants were better able to appreciate future big rewards in a delay discounting task; and that, in spite of their perception of increased rewarding value of both food and monetary objects, hungry participants were not more inclined to take risks to get the object of their desire. Together, these studies for the first time provide evidence that hot states improve decision making under uncertain conditions, challenging the conventional conception of the detrimental role of impulsivity in decision making.     

Costs increase as ritualized fighting progresses within and between phases in the sierra dome spider, Neriene litigiosa

Magnitudes and patterns of energy expenditure in animal contests are seldom measured, but can be critical for predicting contest dynamics and understanding the evolution of ritualized fighting behaviour. In the sierra dome spider, males compete for sexual access to females and their webs. They show three distinct phases of fighting behaviour, escalating from ritualized noncontact display (phase 1) to cooperative wrestling (phase 2), and finally to unritualized, potentially fatal fighting (phase 3). Using CO₂ respirometry, we estimated energetic costs of male-male combat in terms of mean and maximum metabolic rates and the rate of increase in energy expenditure. We also investigated the energetic consequences of age and body mass, and compared fighting metabolism to metabolism during courtship. All three phases involved mean energy expenditures well above resting metabolic rate (3.5×, 7.4× and 11.5×). Both mean and maximum energy expenditure became substantially greater as fights escalated through successive phases. The rates of increase in energy use during phases 2 and 3 were much higher than in phase 1. In addition, age and body mass affected contest energetics. These results are consistent with a basic prediction of evolutionarily stable strategy contest models, that sequences of agonistic behaviours should be organized into phases of escalating energetic costs. Finally, higher energetic costs of escalated fighting compared to courtship provide a rationale for first-male sperm precedence in this spider species.     

Direct benefits of social dominance in juvenile crayfish

Crayfish are known for their innate aggressiveness and willingness to quickly establish dominance relationships among group members. Consequently, the formation of dominance hierarchies and the analysis of behavioral patterns displayed during agonistic encounters have mostly been tested in environments that provide no immediate resources besides space. We tested the hypothesis that social hierarchy formation in crayfish serves to determine access to future resources. Individuals within groups of three juvenile crayfish were allowed to form a social hierarchy in a featureless environment before a single food resource was presented. Higher dominance indices were significantly correlated with increased access to the food. The highest ranked crayfish spent more time in contact with the food than did medium-ranked and lowest ranked crayfish, and crayfish of medium rank spent more time in contact with the resource than did lowest ranked animals. The highest ranked crayfish consolidated their dominant status in the presence of food, indicated by a complete absence of any submissive behaviors during that period. The results of these experiments show that the disposition of crayfish to engage in fighting and formation of a dominance hierarchy in a featureless environment serves to determine future access to an emerging resource, thereby entailing greater benefits for animals of higher social rank.     

Effect of morph types, body size and prior residence on food-site holding by males of the male-dimorphic stag beetle Prosopocoilus inclinatus (Coleoptera: Lucanidae)

Mate-securing tactics of small males in male-polymorphic species exhibiting male–male combat is an important issue in behavioral ecology. While most studies have focused on the outcomes of such combat encounters, the holding of a mating resource like a feeding site has a greater impact for obtaining reproductive success. We examined the effects of the prior residence at a feeding site on resource acquisition in the male-dimorphic stag beetle, Prosopocoilus inclinatus. More than 70 % of encounters did not result in combat. While larger males tended to occupy a food site after a combat, smaller males with prior residence tended to occupy food sites when no fighting occurred. Morph types or body size have no effect on the occurrence of combat, meaning that small males do not hesitate to fight with large males. These findings show that, under experimental conditions, the prior residence has a positive effect to hold food site in P. inclinatus.     

Hormonal Correlates of Dominance and Starvation-induced Aggression in Chicks of the Blue-footed Booby

In the blue-footed booby (Sula nebouxii), the first-hatched chick aggressively dominates its sibling and sometimes kills it when food is in short supply. To investigate the endocrine correlates of dominance-subordinance and hunger-induced agonism, we deprived 15–20-d-old single-chick and two-chick broods of food during 48 h by taping chicks' necks to prevent ingestion of parentally provided food (a protocol used previously and known to elicit escalated sibling fighting). We monitored weight and levels of circulating testosterone, oestradiol and corticosterone in deprived and normally fed broods comprising singletons, seniors (= dominants) and juniors (= subordinates), and observed behaviour to verify that aggression increased in deprived two-chick broods. During the 2 d of fasting, experimental chicks lost on average 6% of their baseline weight. After normal feeding was reinstated, seniors and singletons recovered normal (control) weight, but juniors remained significantly lighter than controls. No testosterone was detected in any nestling, but baseline corticosterone level was 109% higher in juniors than in seniors or singletons, implying that elevated corticosterone in juniors is a consequence of social subordination and may facilitate submissive behaviour. Although there was evidence that aggression of seniors increased under food deprivation, the increase was not accompanied by the rise in levels of testosterone expected under the ‘Challenge hypothesis’ (Wingfield et al. 1990). This result implies that this hypothesis probably does not apply to booby nestlings in the context of starvation-induced aggression. During the 2 d deprivation period, corticosterone levels of experimental chicks increased significantly, then declined to baseline levels a day after tapes were removed. The increase was probably due to the combined effects of starvation and frustration. We suggest that corticosterone could alter responsiveness of nestling blue-footed boobies to external stimuli, resulting in more aggression by dominants and greater readiness of subordinates to submit.     

Effect of food concentration on the production and viability of resting eggs of the rotifer Brachionus: implications for the timing of sexual reproduction

1. Life‐table experiments with Brachionus calyciflorus test several hypotheses related to the idea that sexual reproduction in monogonont rotifers should occur when food resources are favourable. 2. The food concentration necessary for a fertilised mictic female to produce one phenotypically normal resting egg was higher than that for an amictic female to produce one daughter. At the lowest concentration of Cryptomonas erosa (1.25 × 10³ cells mL^?1), the lifetime fecundity of these two types of females was 0.9 and 1.4, respectively. 3. The lifetime fecundity of both fertilised mictic females and amictic females increased with food concentration to 3.4 resting eggs and 15.2 daughters female^?1, respectively. The approach to maximal fecundity with increasing food concentration was more rapid for fertilised mictic females, such that their lifetime fecundity relative to that of amictic females gradually decreased from 0.64 (at 1.25 × 10³ C. erosa mL^?1) to 0.22 (at 2.5 × 10⁴ C. erosa mL^?1). 4. The probability of a fertilised mictic female producing one or more abnormal resting eggs during her lifetime was high (approximately 75%). The mean proportion of abnormal eggs produced per female varied among the different food‐concentration treatments (26–38%) but was not higher at the low food concentrations. 5. The proportion of normal resting eggs that hatched was high (51–71%); those produced at low food concentrations were no less likely to hatch than those produced at high food concentrations. No abnormal resting eggs hatched. 6. The probability of a fertilised mictic female producing an abnormal resting egg increased rapidly with her age at all food concentrations. The probability of a normal resting egg hatching declined with maternal age at the low food concentration in one of two experiments. 7. The results support the idea that induction of mictic females should occur when food resources are good. Coincidence of sexual reproduction with low food availability risks low production of resting eggs for several reasons. Population size may be small, with a low probability of encounters between young mictic females and males, and fertilised mictic females may be unable to mature and produce resting eggs.     

Sound Characteristics and Outcome of Contests in Male Croaking Gouramis (Teleostei)

A correlative study using similar-sized males of the croaking gourami Trichopsis vittata was carried out to investigate whether sound characteristics influenced winning and if relative fighting ability was assessed by acoustic signals. Pair-wise contests between males were decided using lateral displays (LD) and vocalization in 26 cases, whereas 66 fights escalated to the frontal display (FD) phase. Physical fighting (mouth wrestling) and injuries were rarely observed in this species. Winners were generally larger than their opponents, and this effect was more pronounced in non-escalated than in escalated contests. Sounds of fight winners had a higher sound pressure level and also a lower dominant frequency. Neither number of acoustic signals nor duration of lateral and frontal displays were predictors of contest outcome. Acoustic measures were highly correlated to body weight. These results indicate that traits correlated with RHP (such as sound pressure level and dominant frequency) were predictors of the outcome, while traits not correlated with size (such as number and duration of displays) did not influence winning. In accordance with the main prediction of assessment models, the contest duration (cost) increased with the decrease in asymmetry of body length as well as sound pressure level. No such relationships were found for weight and dominant frequencies in LD- and FD-contests. The present study indicates that morphological and sound characteristics influence winning in fish. Moreover, the results suggest that croaking gouramis settle conflicts without damaging combats by assessing asymmetries in different components of RHP such as body weight and length, which may reliably be signalled by acoustic and visual assessment signals.     

Decision Making and Assessment in the Fighting Behaviour of Nannacara anomala (Cichlidae,Pisces)

Predictions from ESS-theory concerning assessment of fighting ability and decision making are tested using a cichlid fish Nannacara anomala. Pairs with a pronounced difference in weight were allowed to interact. Fight duration increased when the difference in weight between the contestants decreased and the probability of an escalation from tail-beating to mouth-wrestling also increased when the weight difference decreased. Despite the fact that weight difference very well predicts the outcome of fights in this species it was found that fishes weighing less than 50 % of the opponent engaged in fighting, which suggests that visual assessment alone provides only limited information about relative fighting ability. The possible functions of various behaviour patterns are discussed in relation to assessment and escalation.     

Influence of body reserves and eye opacity on foraging behaviours of tiger snakes

Physiological states such as reproductive stage, parasite load, body condition, and environmental conditions, influence behaviours in complex ways. According to the Clark's asset-protection principle (Clark, 1994. Behav Ecol 5:159-170), individuals in good body condition take fewer predation risks than conspecifics in lesser condition. In many ectotherms, foraging, digestion, and moulting require an elevation of the metabolism achieved through intensive basking, thereby increasing the risk of being detected by predators. Using four experimental groups of snakes, we showed that two independent physiological states, (1) the presence of prey in the stomach and (2) eye opacity associated with moulting, increase predator vulnerability. In a parallel experiment, two groups of snakes were maintained on contrasting diets (high food intake vs. low food intake) for 6 months. Apart from sloughing periods, most individuals (89%) accepted their meal irrespective of food treatment. Consequently, well-fed snakes exhibited higher body condition (one third physiological state) relative to less-fed individuals. During sloughing events, however, well-fed individuals often refused (78%) to eat while the less-fed individuals readily accepted (86%) their meal despite eye opacity. Individuals with less body reserves accepted the cumulative risks owing to eye opacity and stomach fullness. By contrast, well-fed snakes remained more "prudent" (i.e., exhibited behaviours that depended on their internal state). Our results show that snakes can adjust their foraging "decision" by combining different physiological informations.     

Balancing the competing requirements of air-breathing and display behaviour during male–male interactions in Siamese fighting fish Betta splendens

Air-breathing fish of the Anabantoidei group meet their metabolic requirements for oxygen through both aerial and aquatic gas exchange. Siamese fighting fish Betta splendens are anabantoids that frequently engage in aggressive male–male interactions which cause significant increases in metabolic rate and oxygen requirements. These interactions involve opercular flaring behaviour that is thought to limit aquatic oxygen uptake, and combines with the increase in metabolic rate to cause an increase in air-breathing behaviour. Air-breathing events interrupt display behaviour and increase risk of predation, raising the question of how Siamese fighting fish manage their oxygen requirements during agonistic encounters. Using open-flow respirometry, we measured rate of oxygen consumption in displaying fish to determine if males increase oxygen uptake per breath to minimise visits to the surface, or increase their reliance on aquatic oxygen uptake. We found that the increased oxygen requirements of Siamese fighting fish during display behaviour were met by increased oxygen uptake from the air with no significant changes in aquatic oxygen uptake. The increased aerial oxygen uptake was achieved almost entirely by an increase in air-breathing frequency. We conclude that limitations imposed by the reduced gill surface area of air-breathing fish restrict the ability of Siamese fighting fish to increase aquatic uptake, and limitations of the air-breathing organ of anabantoids largely restrict their capacity to increase oxygen uptake per breath. The resulting need to increase surfacing frequency during metabolically demanding agonistic encounters has presumably contributed to the evolution of the stereotyped surfacing behaviour seen during male–male interactions, during which one of the fish will lead the other to the surface, and each will take a breath of air.     

Interspecific aggression in captive male lemmings

Two experiments were conducted with lemmings to determine which of two species, Lemmus trimucronatus and Dicrostonyx groenlandicus, was capable of exercising aggressive social dominance over the other in staged, dyadic encounters, and which was able to outcompete the other for a single shelter. Agonistic behaviour occurred in all dyadic encounters, but dominance was not established by either species. A sequential analysis of encounters revealed that each species had a distinct fighting strategy. In shelter competition, prior residence was a crucial factor in the outcome. However, one species did not completely exclude the other. Further evidence to support species differences in fighting strategies was derived from the shelter experiments. These results are discussed in relation to ecological considerations.     

Spatial distribution patterns of sheep following manipulation of feeding motivation and food availability

We hypothesised that (i) increased feeding motivation will cause sheep to move further apart as a result of individuals trying to find food and (ii) in conditions of high food availability, sheep will move less and show greater social attraction. The effects of both feeding motivation and food availability on spatial distribution was examined in eight groups of food-deprived (high feeding motivation) and satiated (low feeding motivation) sheep in good or poor food resource plots in a 2 × 2 design. Distance travelled was assessed using Global Positioning System collars, grazing time using scan sampling and social cohesion using proximity collars that record the number and duration of encounters within 4 m. Food-deprived sheep in the good-resource plots grazed the most, whereas satiated sheep in the poor-resource plots grazed the least (P = 0.004). Food deprivation had no significant effect on the number or duration of encounters and feeding motivation appeared to have little effect on spatial distribution. Contrary to expectation, sheep had more encounters (P = 0.04) of a longer total duration (P = 0.02) in poor-resource plots than in good-resource plots, indicating that sheep were showing more social cohesion if food was scarce. Our findings suggest that when food is scarce, animals may come together in an attempt to share information on food availability. However, when a highly preferred food is abundant and well dispersed, they may move apart in order to maximise the intake. It is concluded that the particular details of our experiment, namely the even distribution or absence of a highly preferred food, affected spatial distribution patterns as sheep tried to find this food and maximise the intake.