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.     

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.     

Male collared lizards,Crotaphytus collaris (Sauria: Crotaphytidae), signal females by broadcasting visual displays

The signalling function of displays broadcast when animals are distant from conspecifics can be difficult to determine. I tested the extent to which visually transmitted broadcast displays given by free‐ranging territorial male collared lizards signalled same‐sex rivals or females. One test involved recording the frequency of broadcast displays, aggressive contests with rivals, and courtship encounters with females during ten reproductive seasons when local sex ratios varied markedly. The frequency of broadcast displays decreased as the ratio of male competitors to females increased. The frequency with which males initiated contests with rivals was not related to the ratio of competitors to females, whereas the frequency of courtship interactions decreased with sex ratio because there were fewer females to court. The behaviour of males that defended territories during two successive seasons showed a similar pattern. Broadcast display frequency was positively correlated with courtship frequency, but not with the frequency of contests with rivals. Lastly, individual males gave more broadcast displays during focal observations when they also engaged in courtship encounters with females than other observations when they engaged in aggressive conflicts with rival males. Although these results do not reject the possibility that broadcast displays may also signal male rivals, they support a major role of these displays in advertisement to females. © 2013 The Linnean Society of London     

Deferred agonistic behavior in a long-lived scincid lizard Eumeces laticeps

Summary A laboratory experiment with the broad-headed skink (Eumeces laticeps) involving staged agonistic encounters demonstrates that larger males have an advantage over smaller ones in agonistic bouts. Field data on head wounds produced by intraspecific fighting during the breeding season show a much higher frequency of new wounds among males over 100 mm in snout-vent-length than in smaller males. The significant difference in new-wound frequency strongly suggests avoidance of fights by the small males, which is corroborated by laboratory and field observations. Access by males to reproductively active females depends on the ability to defeat other males in aggressive contests virtually always involving head biting if the males are of nearly equal size. Because the probability of winning agonistic encounters increases with size, young males avoid fights with older males. Aggressive contests with larger males and reproductive attempts other than courtship in the absence of larger males are deferred.Aggressive behavior in E. laticeps may be employed in direct defense of females, but might also be expressed in defense of specific sites and/or territories. In the laboratory, males in their home cages were significantly more likely to win encounters with males of similar size than were males fighting in the home cages of opponents. This suggests that encounter site could be important in determining encounter outcome and that field study of possible site defense or territoriality is needed.     

Territorial behaviour of the red-backed salamander: Expulsion of intruders

Red-backed salamanders, Plethodon cinereus, were paired in laboratory chambers where they defended areas against each other by threat display and biting. Difference in sizes of paired individuals was not correlated with which one would enter the other's area or be more aggressive after an intrusion. We also found no sexual differences in agonistic behaviour. Both defenders and intruders assumed a threat posture for about 50% of the invasion periods, but defenders exhibited a higher rate of biting. Defenders expelled intruders in 74% of all encounters, while intruders expelled defenders in 18%, with 8% of the encounters a draw. Intruders frequently left the contested area even in the absence of biting or threat display by the defender, apparently in response to the defender's pheromones or to the high potential cost of an escalated aggressive contest (damage to the chemosensory structures). These data provide behavioural evidence that a salamander can evict intruders from a defended area and thus support the hypothesis of territoriality among terrestrial salamanders.     

Signaling by Protrusive Scent Glands in Cave Crickets,Troglophilus neglectus Krauss (Orthoptera: Rhaphidophoridae), is Primarily Involved in Male-male Agonism

Males of several species of cave crickets (Rhaphidophoridae) possess protrusive abdominal scent glands, which are generally presumed to influence female behavior before mating. The validity of this hypothesis, as well as of the alternative possibility of the signal being designed for the rivals, was tested in Troglophilus neglectus by describing the detailed context and the consequences of such signaling by gland protrusion in the mating period. Small groups of both sexes were observed under naturalistic conditions, using interval and focal behavioral sampling. The frequency of gland protrusion increased progressively in both experimental seasons, and was expressed in solitary males as well as in their interactions with both mates and rivals. Such signaling, however, was not an obligatory part of courtship and pre-mating behavior in general, and caused neither female attraction nor influenced the courtship success. The males protruded the glands significantly more frequently during the antennal contacts with another male than during contacts with a female. Of the different behaviors expressed in the inter-male encounters, only aggressiveness was significantly correlated with protrusion of the glands in the respective individuals. This strongly implies the function of the male scent as an agonistic signal, influencing the outcome of the intermale contests.     

Interactions between behavior and plasma steroids within the scramble mating system of the promiscuous green turtle, Chelonia mydas.

We measured plasma androgen (combined testosterone and 5alpha-dihydrotestosterone) (A) and corticosterone (B) in the promiscuous green turtle (Chelonia mydas) during courtship in the southern Great Barrier Reef. This study examined if reproductive behaviors and intermale aggression induced behavioral androgen and adrenocortical responses in reproductively active male and female green turtles. Associations between reproductive behavior and plasma steroids were investigated in green turtles across the population and within individuals. Levels across a range of both asocial and social behaviors were compared including (a) free swimming behavior; (b) initial courtship interactions; (c) mounted behavior (male and female turtles involved in copulatory activities); (d) intermale aggression (rival males that physically competed with another male turtle or mounted males recipient to these aggressive interactions); and (e) extensive courtship damage (male turtles that had accumulated excessive courtship damage from rival males). Behavioral androgen responses were detected in male turtles, in that plasma A was observed to increase with both attendant and mounted behavior. Male turtles who had been subjected to intermale aggression or who had accumulated severe courtship damage exhibited significantly lower plasma A than their respective controls. No pronounced adrenocortical response was observed after either intermale aggression or accumulation of extensive courtship damage. Female turtles exhibited a significant increase in plasma B during swimming versus mounted behavior, but no change in plasma A. We discuss our results in terms of how scramble polygamy might influence behavioral androgen interactions differently from more typical combative and territorial forms of male polygamy.     

Does size matter? Assessment and fighting in small and large size‐matched pairs of adult male convict cichlids

Contests between matched pairs of sexually mature convict cichlids Archocentrus nigrofasciatus were staged to determine if size alone affected the ways in which the males fought. Both small and large contests began with a period of visual displays. As fights progressed, displays diminished, and were replaced by more escalated behaviours. Small and large contests were also of similar duration. Despite these consistencies with sequential assessment, large and small contests were not entirely alike. For example, large males engaged in more lateral displays and were slower to escalate than small males. Small males displayed less and escalated to biting sooner than large males. Two possible explanations are suggested for these contrasts in contest structure: absolute opponent size and differences in the males' ages coupled with differences in experience.     

Don't count your eggs before they're parasitized: contest resolution and the trade-offsduring patch defense in a parasitoid wasp

Although aggressive conflicts over hosts occur among femalesin many species of insect parasitoids, few studies have examinedthe mechanisms by which these contests are resolved. In Trissolcusbasalis, a parasitoid of pentatomid bug egg masses, femalesco-exploiting an egg mass (patch) encounter one another repeatedlyand fight for possession of the patch. We investigated the resolutionof pairwise contests by experimentally varying the release timeof females and the size of the patch. Logistic regression showedthat the female arriving first on the patch was more likely towin both the first agonistic encounter and to retain overallpossession of the patch. This advantage to the first femalesuggests a resource-correlated asymmetry in favor of the firstfemale, due to her having invested more offspring in the patch.Although escalations were more common when the asymmetry inarrival times was small, the majority of encounters within all contestswere nevertheless resolved without escalated fighting, withthe resident attacking and the intruder backing down. Thus contestresolution basically followed a "bourgeois" rule. Residentstolerated intruders more frequently when the patch size waslarger and the second female was released later, illustratingthe trade-off faced by residents between defending the patchand continuing to exploit it.     

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.     

Non‐Conceptive Sexual Behavior in Spiders: A Form of Play Associated with Body Condition,Personality Type,and Male Intrasexual Selection

In the socially polymorphic spider Anelosimus studiosus, males mature early in the reproductive season and recruit to the webs of juvenile females and guard them until they mature. During the period before females mature, males and females engage in repeated bouts of non‐conceptive (play) sexual behavior, where the pair courts and engages in mock copulation; both males and females gain performance‐enhancing experience via these encounters. In this study, we examined the factors that underlie individual variation in the tendency to engage in non‐conceptive mating and determine whether it impacts male–male competition for females. We found that docile females, being less resistant to mating in general, are more likely to accept male courtship and non‐conceptive copulation as juveniles. Personality type influenced the exhibition of non‐conceptive sexual behavior in males as well. High body condition males of the aggressive phenotype were more likely to engage in non‐conceptive sexual behavior than males with lower body condition. Body condition did not influence docile males’ propensity to engage in non‐conceptive sexual behavior, but female size did. Docile males engaged in more non‐conceptive sexual displays with larger females. Engaging in non‐conceptive sexual displays negatively impacted male performance in staged male–male contests for access to females. This cost was greatest for males of the aggressive phenotype, which are otherwise favored in male–male contests. Our findings indicate expression of non‐conceptive sexual displays is linked to personality and results in reproductive performance trade‐offs for male A. studiosus.     

Production of acoustic signals during aggression in Coosa bass, <Emphasis Type="Italic">Micropterus coosae</Emphasis>

Although sound production has been described for sunfishes, it is previously unknown for basses, both groups of fishes in the family Centrarchidae. We document production of acoustic signals during aggressive encounters in Coosa bass, Micropterus coosae. During dyadic encounters, presumptive winners of contests produced sounds associated with a variety of behaviors, including chases, lateral displays, circle swims, nudging, jerking and during post aggression, while fish were stationary. These sounds are low-frequency, non-harmonic and consist of one to 41 separate pulses. In most trials larger fish won contests, regardless of territory ownership (which fish was put into the test tank first) and size difference.     

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.     

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.     

Aggression in harlequin frogs: male-male competition and a possible conflict of interest between the sexes

Long-term field observations and experimental manipulations were made on a population of harlequin frogs, Atelopus varius, in Costa Rica to document seasonality of intra- and intersexual aggression, determine the function of aggression, and identify the factors that influence success in aggressive encounters. Both males and females defended portions of their home range and exhibited site fidelity. Males were more aggressive towards other males during the pre-breeding and breeding seasons, whereas females were more aggressive towards other females during the post-breeding season. For up to several months prior to oviposition, males attempted to amplex (clasp) females without courtship behaviour, while females chased males from their territories and attempted to dislodge males that amplexed them. In staged encounters the winner was usually the apparent resident, regardless of body size. Frogs from a high-density population were more aggressive than individuals from a low-density population. Subadult males were as aggressive and as likely to win encounters as adult males. The function of aggression seemed to be related to the bizarre reproductives behaviour observed in these frogs. There seemed to be male-male competition for mates, as the operational sex ratio strongly favoured males, oviposition was asynchronous, and amplexus lasted for at least several weeks. Female-female aggression may have been related to defence of foraging or shelter sites, but aggression towards males was likely to be an attempt to thwart unwanted amplexus.     

Courtship song, male agonistic encounters, and female mate choice in the house cricket,Acheta domesticus (Orthoptera: Gryllidae)

Whether female crickets choose among males based on characteristics of the courtship song is uncertain, but in many species, males not producing courtship song do not mate. In the house cricket,Acheta domesticus, we examined whether a female chose or rejected a male based on his size, latency to chirp, latency to produce courtship song, or rate of the high-frequency pulse of courtship song (“court rate”). We confirmed that females mated only with males that produced courtship song, but we found no evidence that the other factors we measured affected a female’s decision to mate. In addition, we investigated whether the outcome of male agonistic encounters affected the subsequent production of courtship song. In one experiment, we observed courtship and mating behavior when a single female was placed with a pair of males following a 10-min interaction period between the two males. Winners of male agonistic encounters had higher mating success. However, winners and losers of agonistic encounters were not different in their likelihood or latency to produce courtship song or in the number of times they were disrupted by the other male in the pair. In a second experiment, we allowed two males to interact for a 10-min period, but following this interaction period, we placed a female with each male separately and observed courtship and mating behavior. The mating success of winners and losers was not different under these circumstances, and we found no differences between winners and losers in any subsequent courtship or mating behavior examined. We conclude that winning agonistic encounters influences a male’s mating success in ways other than his production of courtship song and this effect is lost when winning and losing males are separated and each is given an opportunity to mate.     

Vibratory Communication in the Jumping Spider Phidippus clarus: Substrate‐borne Courtship Signals are Important for Male Mating Success

Recently, work has shown that multimodal communication is common throughout the animal kingdom but the function of multimodal signals is still poorly understood. Phidippus clarus are jumping spiders in which males produce multimodal (visual and vibrational) signals in both male–male (aggressive) and male–female (courtship) contexts. The P. clarus mating system is complex, with sex ratios and the level of male competition changing over the course of the breeding season. Vibrational signal components have been shown to function in male aggressive contests but their role in courtship has not been investigated. Here, we performed an experiment to test the role of vibrational signaling in courtship by observing mating success for males that were experimentally muted. We show that vibratory courtship signals, and in particular signaling rate, is an important component of mating success and potentially a target of female choice. While the ability to produce vibratory signals significantly increased mating success, some muted males were still able to successfully mate. In these trials, signaling rate also predicted mating success suggesting that redundant signal components may compensate for errors and perturbations in signal transmission or that vibratory signals function to enhance the efficacy of visual signals.     

The breeding behaviour of a northern temperate orconectid crayfish, Orconectes rusticus

Observations of breeding activities of male and female crayfish (Orconectes rusticus) in a natural stream population during the explosive spring breeding period, and supplemental laboratory observations, indicate a promiscuous mating system characterized by intense intermale aggression. Males wandered extensively over the substrate, fought frequently with each other, and attempted to disrupt copulating pairs. Larger males (as distinguished by large chela size) more frequently initiated and won aggressive encounters and interrupted copulating pairs more successfully than smaller males, but were subjected to greater predation. Intense intermale aggression ceased when females sequestered themselves under rocks in the stream or were removed from the laboratory population. Brief resistance by females to approaching males, along with sequestering behaviour, suggests that female choice may also occur.     

Familiarity influences body darkening in territorial disputes between juvenile salmon

Escalated contests between animals are potentially costly because of increased energy expenditure and risk of predation or injury. Hence we would expect selection to favour any mechanism that avoids unnecessary prolonged fighting. One such means of avoiding escalated fights could be the use of information gained through individual recognition. Previous work has shown that a darkening of the body colour is closely associated with submission in contests between juvenile Atlantic salmon, Salmo salar, and it has been hypothesized that this may act as a visual signal to the opponent. We tested the hypothesis that body darkening is used to reduce the cost of contests between familiar fish such that losers darken more quickly when faced with familiar than unfamiliar opponents. In contests between unfamiliar fish, submissive darkening occurred after more escalated contests in which the loser incurred more aggression, whereas the opposite occurred when familiar fish were in conflict. In addition familiar fish either submitted quickly or engaged in protracted conflicts in which neither fish signalled submission, whereas in unfamiliar fish contests were of intermediate duration regardless of whether either fish darkened. We suggest that body darkening is used by familiar fish to signal submission to familiar dominants in order to avert a costly escalated fight, but familiarity can lead to escalation without submission if perceived competitive asymmetries are finely balanced. Copyright 2000 The Association for the Study of Animal Behaviour.