Effects of female body size and age and male mating status on male combat in Anastatus disparis (Hymenoptera: Eupelmidae)

1. Aggressive behaviour is widely exhibited by animals to acquire important resources and usually shows a dangerous or nondangerous pattern. Dangerous fighting patterns are usually characterized by fights ending with contestants being severely injured or killed. Resource value is an important nonstrategic factor influencing fighting behaviour. Studies of many species addressing nondangerous fighting behaviour have shown that when resource values change, organisms usually adjust their fighting behaviour accordingly. Only a few species show dangerous fighting patterns. Thus, few relevant studies have addressed how variation in resource value affects aggression with a dangerous fighting pattern. 2. Here, an egg parasitoid wasp, Anastatus disparis, which exhibits a dangerous fighting pattern to acquire mating opportunities, was used as an experimental model to study the adjustment of fighting behaviour resulting from a change in resource value. 3. Our results show that the female properties of body size and age affect their objective resource value and that males increased their fighting intensity for relatively large and young females. However, male mating status in A. disparis may not influence the subjective value of mate resources, and fighting intensity did not significantly differ between mated and virgin males. In addition, the number of times a male had previously mated had no significant effect. These results suggest that mating opportunities are important for both virgin and mated males, resulting in neither of them showing any adjustment in fighting for mating opportunities. 4. Generally, A. disparis males with extreme fighting patterns adjust their fighting behaviour according to the variation in resource value, which avoids the meaningless costs of injury and death.     

Social selection and the evolution of a female weapon in queens of the ant Messor pergandei (Hymenoptera: Formicidae)

The evolution of weaponry occurs less frequently in females than in males and is most often important for protecting ecological resources or offspring rather than winning mates. The purpose of female weapons is often confounded by the presence of similar weapons in males, so cases where only females need weapons provide important tests of our understanding of how and why weapons evolve. In some populations of the ant, Messor pergandei (Mayr), newly mated queens initiate new nests in social groups that subsequently break down when queens engage in battles for control. The incipient social environment differs geographically, so that lethal fighting occurs in some populations but not others. Consistent with the hypothesis that queens in populations where lethal fighting occurs should show selection for weaponry (broad heads and strong mandibles), we found that heads of queens from sites where lethal fighting occurs were broader than those at sites with non‐fighting queens and a site with solitary queens. Evolution of weaponry is specific to queens, because regression results from workers often did not follow this pattern. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 1011–1020.     

Fighting in fig wasps: do males avoid killing brothers or do they never meet them?

1. In many fig wasp species, armoured wingless males regularly engage in lethal fights for access to females inside figs, which act as discrete mating patches. 2. Kin selection generally opposes killing brothers, because their reproductive success provides indirect genetic benefits (inclusive fitness). However, siblicide may be avoided if (i) brothers do not occur in the same figs, or (ii) males avoid fighting brothers in the same fig. Alternatively, (iii) siblicide may occur because intense mate competition between brothers at the local scale overcomes kin selection effects, or (iv) males do not recognise kin. 3. A fig may also contain wasps from other closely related species and it is not known if males also fight with these individuals. 4. Nine microsatellite loci were used in the first genetic analysis of fighting in fig wasps. We assigned species and sibling identities to males and tested alternative fighting scenarios for three Sycoscapter wasp species in figs of Ficus rubiginosa. 5. Approximately 60% of figs contained males from more than one Sycoscapter species and approximately 80% of fights were between conspecifics, but a surprising 20% were between heterospecific males. 6. Within species, few figs contained brothers, suggesting that females typically lay one son per fig. Overall, most males do not compete with brothers and all fights observed were between unrelated males.     

Strong Competition Does Not Always Predict Play Asymmetry: The Case of South American Sea Lions (Otaria flavescens)

Although play fighting has been studied for over a century in both human and non‐human animals, quantitative data on marine mammals are still scarce. Here, we investigated play fighting in South American sea lions (Otaria flavescens), one of the most sexually dimorphic species with an extreme polygynous mating system, high levels of both intra‐ and inter‐sexual competition. All these features make South American sea lions a good model species to test some predictions on play fighting. Our results indicate play is restricted to juveniles, being inhibited among adults, and as to be expected in a species that shows a high degree of sexual dimorphism, it is mainly expressed in males. Even though playful interactions were punctuated by competitive behaviours, animals played in a highly symmetric way and were able to adjust their competitive playful interactions in a flexible manner and so reduce the risk of escalation to a minimum level. They were highly selective in their choice of playmates by limiting the number of players per session and playing more with age‐matched companions and friends. All these factors taken together are probably at the basis of the low risk of escalation recorded during the study. This result is predictive of a high ability and motivation of these animals to engage in play behaviour which can have a possible role not only in the acquisition of dominance status, but also in establishing and maintaining social relationships, an unexpected role in a so highly competitive species.     

Cobreeding in the Burying Beetle,Nicrophorus vespilloides: Tolerance Rather Than Cooperation

Under intra‐ and interspecific competition, cooperative behaviour can provide direct fitness benefits if individuals work together to expel intruders. In the burying beetle Nicrophorus vespilloides, a relatively small species, multiple unrelated pairs can breed together, and individuals are weak competitors in interactions when competing with larger individuals of the same species and with larger species of the same genus. In field and laboratory studies, we found that two pairs together attracted significantly fewer congeneric and non‐congeneric competitors compared with single pairs. No other benefits were found. Communal breeding had large negative effects on fitness, as there were fewer offspring per pair and a higher chance of injuries. The higher chance of injuries reflected pairs fighting among themselves and not against competitors. These costs are much greater than the small benefit of fewer intruders. Why should a N. vespilloides breeding pair eventually allow another pair to join? A potential partial explanation is that these are not cooperative pairs in the traditional sense, but rather pairs have a higher tolerance for each other. From the resident's perspective, joining pairs are not expelled because the chance of injury makes the cost of fighting high. Joining others may have unusually low costs in this species because reproductive opportunities are rare, dependent on a resource that is unpredictable in time and space, and residents should be inclined to tolerating new pairs because of the cost of fighting. Thus, for N. vespilloides, communal breeding appears to be ‘making the best of a bad job’, providing some reproduction rather than none.     

Agonistic Displays and the Benefits of Fighting in the Field Cricket, Gryllus bimaculatus

Fighting is often composed of discrete agonistic displays. Few studies have partitioned fighting behavior into its component agonistic displays and evaluated the relationships between the frequency of the displays and the potential benefits of fighting, particularly mating success. In this study, we quantified the frequency of male field cricket, Gryllus bimaculatus, agonistic displays. The displays were quantified under three social environments which varied in the potential benefits of fighting: males with other males only, males with other males and female scents, and males with other males and females. We found that (1) the presence of females elicited an increase in agonistic displays characteristic of intermediate levels of escalation, (2) female scents did not produce a similar increase in the frequency of agonistic displays, and (3) in the presence of females, the frequency of agonistic displays was positively correlated with mating success. Aggressive stridulation, an energetically low-cost display, was more strongly associated with mating success than were more costly displays. The results are discussed in the context of the evolutionary theory of aggression and in the context of cricket mating systems.     

The role of social and/or ecological contexts influences assessment strategy use in Tilapia

Animals engage in costly agonistic contests during which winners procure resources. During these interactions, the combatants obtain and use information to make decisions on whether to persist or to withdraw from the fight, which is termed assessment. Recent theory and work have suggested that the types of assessment employed may be more variable than previously thought, with the use of different strategies possibly being influenced by social and ecological conditions during priming. This study addresses the contextual components (social and ecological) that affect the utilization of one assessment strategy over another. Male tilapia were primed with different combinations of social (large and small animals) and ecological (resource rich or poor) contexts 24 hr prior to fighting in staged, dyadic contests. When opponents were primed with the same context, a clear assessment strategy emerged and differed as a function of priming treatment. Conversely, when fish were primed with different treatment contexts, there was no discernible assessment. In addition, priming conditions had differing effects for large and small fish. Thus, assessment strategies in cichlids are dependent upon a combination of social, ecological contexts and size of the animal. Since assessment strategies change as a function of both of these contexts, as well as others, future framework investigating assessment strategies should include both intrinsic and extrinsic factors that may shape fighting dynamics.     

The evolution of social behaviour in sentinel crabs (Macrophthalmus): implications from molecular phylogeny

Crabs of the genus Macrophthalmus are known to exhibit highly developed and diverse social behaviour, such as allocleaning, fighting and waving display behaviour, the first being observed widely throughout the genus. Fighting behaviour between males has been classified previously into grasping fighting and claw‐extending fighting, and male waving display into four patterns, the vertical non‐forward‐pointing type, vertical forward‐pointing type, lateral non‐forward‐pointing type and lateral forward‐pointing type, on the basis of interspecific behaviour comparisons. To understand the evolutionary pathways of these social behavioural activities, 978‐bp nucleotide sequences from mitochondrial 16S rRNA genes of 21 species, including two outgroup taxa, were analysed and a molecular phylogeny was reconstructed. The resultant tree demonstrated striking inconsistencies with the relationships inferred from morphological features. Species with similar habitat conditions showed similar morphological features, although they were not phylogenetically close relatives. Phylogenetic analysis of allocleaning behaviour suggested that it evolved once in the early history of the lineage. The analysis of fighting behaviour demonstrated that species with claw‐extending fighting, being a more complex behaviour than grasping fighting, are found in the most ancestral part of the phylogeny. The analysis also revealed that claw‐extending fighting has evolved secondarily on two occasions, suggesting that fighting behaviour is not characterized by sufficient phylogenetic components. The superimposition of a waving pattern on to the phylogeny indicated that the lateral non‐forward‐pointing type has evolved from the vertical non‐forward‐pointing type, the lateral forward‐pointing type having evolved from the vertical forward‐pointing type. This scenario also appeared reasonable with respect to the behavioural trends of cheliped movements in waving. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 88 , 45–59.     

Intra-sexual Dimorphism in Male Mandibles and Male Aggressive Behavior in the Broad-Horned Flour Beetle Gnatocerus cornutus (Coleoptera: Tenebrionidae)

In the stored-product beetle, the broad-horned flour beetle, Gnatocerus cornutus (Fabricius), all males possess enlarged mandibles, widened gena, and a pair of small horns on the vertex, but females lack these completely. Observations of male-male interactions of G. cornutus showed that larger individuals won male-male fights, and that the mandibles were used as weapons. Morphological analysis based on the non-linearity test of Eberhard and Gutierrez's model (1991) showed that intra-sexual dimorphism in males was only found in the mandibles used in male-male combat, but not in the gena and horns. This beetle can be an ideal model for evolutionary studies of exaggerated weapons for male combat, because rearing successive generations and observing male fighting are easy.     

The coevolution of juvenile play–fighting and adult competition

Although play–fighting is widespread among juvenile mammals, its adaptive significance remains unclear. It has been proposed that play is beneficial for developing skills to improve success in adult contests (motor‐training hypothesis), but the links between juvenile play–fighting and adult aggression are complex and not well understood. In this theoretical study, we investigate the coevolution between juvenile play–fighting and adult aggression using evolutionary computer simulations. We consider a simple life history with two sequential stages: a juvenile phase in which individuals play–fight with other juveniles to develop their fighting skills; and an adult phase in which individuals engage in potentially aggressive contests over access to resources and ultimately mating opportunities, leading to reproductive success. The simulations track genetic evolution in key traits affecting adult contests, such as the level of aggression, as well as juvenile investment in play–fighting, capturing the coevolutionary feedbacks between juvenile and adult decisions. We find that coevolution leads to one of two outcomes: a high‐play, high‐aggression situation with highly aggressive adult contests preceded by a prolonged period of juvenile play–fighting to improve fighting ability, or a low‐play, low‐aggression situation in which adult contests are resolved without fighting and there is minimal investment in play–fighting before individuals mature. Which of these outcomes is favoured depends on the mortality costs and on the type of societal structure: societies with strong reproductive skew, favouring monopolization of resources, show high levels of adult aggression and high investment in juvenile play–fighting, whereas societies with low reproductive skew have both low adult aggression and low levels of play–fighting. A review of empirical evidence, particularly in the primate genus Macaca, highlights some limitations of our model and suggests that other, complementary functional explanations are needed to account for the full range of competitive and cooperative forms of play–fighting. Our study illustrates the power of evolutionary simulations to shed light on the long‐standing puzzle of animal play.     

Humpback whales interfering when mammal‐eating killer whales attack other species: Mobbing behavior and interspecific altruism?

Humpback whales (Megaptera novaeangliae) are known to interfere with attacking killer whales (Orcinus orca). To investigate why, we reviewed accounts of 115 interactions between them. Humpbacks initiated the majority of interactions (57% vs. 43%; n = 72), although the killer whales were almost exclusively mammal‐eating forms (MEKWs, 95%) vs. fish‐eaters (5%; n = 108). When MEKWs approached humpbacks (n = 27), they attacked 85% of the time and targeted only calves. When humpbacks approached killer whales (n = 41), 93% were MEKWs, and ≥87% of them were attacking or feeding on prey at the time. When humpbacks interacted with attacking MEKWs, 11% of the prey were humpbacks and 89% comprised 10 other species, including three cetaceans, six pinnipeds, and one teleost fish. Approaching humpbacks often harassed attacking MEKWs (≥55% of 56 interactions), regardless of the prey species, which we argue was mobbing behavior. Humpback mobbing sometimes allowed MEKW prey, including nonhumpbacks, to escape. We suggest that humpbacks initially responded to vocalizations of attacking MEKWs without knowing the prey species targeted. Although reciprocity or kin selection might explain communal defense of conspecific calves, there was no apparent benefit to humpbacks continuing to interfere when other species were being attacked. Interspecific altruism, even if unintentional, could not be ruled out.     

Detailed analysis of fighting in polecats (Mustelidae) using ciné film

The phenomenon of fighting was investigated under laboratory conditions using polecats ( Mustela putorius, M. furo and interspecific hybrids). Diadic interactions between male polecats were recorded on 16 mm cine film in an unfamiliar area of 16 m², one individual having been introduced ten minutes before its opponent.
The behaviour patterns involved in fighting are described and their frequencies of occurrence, duration and average bout lengths specified. Biting, which occupied 41% of the animal's time in fighting, and attacking, which made up 27% of the bouts of behaviour, were the most important actions involved in fighting. Polecats most commonly bite their opponent's neck and bites in this region were the longest in duration. Success in gaining a bite was influenced by the opponent's behaviour at the time of the attack.
Polecats fight when one individual bites its opponent and the opponent retaliates by biting; bite was shown to be the commonest response to being bitten. The length of a fight appears to be determined by the levels of motivation to bite of the two opponents, while biting and being bitten operate as a positive feed back mechanism. The fight terminates when the more aggressive opponent ceases to make spontaneous attacks. The reasons why one individual wins a fight are that it consistently spends more time than its opponent in biting, it persists for longer in making spontaneous attacks and its method of attacking is more efficient.     

Effects of starvation on the fighting ability of invasive and autochthonous ants

Ants are widespread in all terrestrial habitats, and competitive interactions between species are common. Although redistribution of food within a colony may buffer the negative effects of temporary resource shortages, colony functionality can be compromised when famine is prolonged. One of the possible effects of famine is impairment of the fighting ability of species, with cascade effects on community. Here, we investigated whether food shortage alters the fighting ability of workers of three dominant species in the Mediterranean area: the invasive alien species, Lasius neglectus and Linepithema humile, and one highly polydomous autochthonous species belonging to the Tapinoma nigerrimum complex. We performed laboratory tests of interspecific one-on-one aggression and pairwise group contests between species, with all possible combinations of artificially satiated and starved groups. Results showed that starvation had a scarce effect on the individual aggressiveness in all three species. Similarly, the outcomes of the group fights were only lightly affected, but with an important exception. The positions of species in the fighting hierarchies were in most cases clear and linear, with L. neglectus at the top. However, we found that L. humile and L. neglectus showed equal mortality when one of the two species was starved and the other satiated. Although we investigated only one aspect of competition, that is, fighting ability, our results provide a piece of the complex jigsaw of competitive interactions of ants, and suggest that food deprivation can be a determinant that alters the relationships between ants and promotes or hampers the coexistence of dominant species.     

Scent may signal fighting ability in male Iberian rock lizards

Intrasexual competition favours the evolution of conspicuous fighting ability badges. However, in spite of the fact that chemoreception is important in sexual selection of many animals, such as lizards, the role of chemical signals in males' contests is relatively unknown. Here, we show that proportions of cholesterol in femoral gland secretions of male Iberian rock lizards were related to their body size (which confers a competitive advantage in fights). Males discriminated chemically and responded aggressively to cholesterol stimuli presented on swabs. Moreover, we experimentally increased cholesterol in the scent of males, and staged encounters in neutral cages between two unfamiliar and size-matched males. Focal males lost more agonisitic interactions against males manipulated with cholesterol than in control tests. We suggest that differences in scent composition may reliably signal fighting ability in many lizard species, which would help to avoid the costs of fighting.     

Pair territoriality in the longnose filefish,oxymonacanthus longirostris

The longnose filefish,Oxymonacanthus longirostris, usually lives in heterosexual pairs, the male and female swimming together and sharing the same territory. Pair territoriality in the species was examined in detail in relation to sexual differences in territorial defense activities. Rigorous pair territoriality was maintained only during the breeding season, although pairs used their home ranges exclusively to a certain extent, during the non-breeding season. The frequency of aggression against other conspecific pairs in the breeding season was higher than in the non-breeding season. Agonistic interactions appear to be over both mates and food resources, the strict pair territoriality in the breeding season possibly being due to mutual mate guarding. In intraspecific aggressive interactions, males usually led their partner females when attacking intruders. The feeding frequency of males was much lower than that of females in the breeding season. Mate removal experiments indicated that females could not defend their original territories solitarily and their feeding frequency decreased. Conversely, males could defend territories solitarily without a decrease in feeding frequency. These results suggest that males contribute most to the defense of the pair territory, with females benefiting from territorial pair-swimming with their partner males.     

Characteristics of social interaction between unfamiliar male rats (<Emphasis Type="Italic">Rattus norvegicus</Emphasis>): comparison of juvenile and adult stages

Social interaction between juvenile male rats that were unfamiliar with one another (JNI) was compared with that of adult unfamiliar animals (ANI). In their home cages, subjects were confronted with a stimulus conspecific and their agonistic behavior patterns were analyzed. It was found that behaviors that characterize "play fighting" (pouncing, submissive posture, on-top posture, etc.) were observed more in JNI than in ANI, whereas behaviors found in serious fighting (pawing, upright posture, etc.) were shown more in ANI. Analyses of the transitions between behaviors also revealed that the transitions constituting play fighting were found only in JNI. On the other hand, transitions that characterize serious fighting were shown in both JNI and ANI. These findings are discussed along with those of a previous study that observed the agonistic behavior between littermates, and the existence of some cue that signals the play-fighting situation among rats is suggested. Electronic Publication     

Who's Watching Me: Female Siamese Fighting Fish Alter Their Interactions in Response to an Audience

It is well established that interactions between conspecifics are often influenced by the presence of passive bystanders. Individuals have been found to alter their behavior in a variety of contexts, from foraging to aggression, based on the presence, sex, or identity of an audience. This audience effect may influence not only the nature of a signaling event but also the evolution of signal structure as signals may have to convey information across a distance. Additionally, audience individuals may use information obtained by watching in later encounters with these individuals, which may act as a selection pressure on communication. Communication networks in Siamese fighting fish, Betta splendens, are particularly well studied, with audience effects influencing both male–male interactions and male–female interactions. However, the effects of an audience on female–female interactions have not been examined in this species or any other. This study examined the interactions of pairs of females in three different audience conditions (male, female, and no audience). The results suggest that female–female interactions are affected by the presence of an audience as interactant‐directed gill flaring, the most commonly performed behavior, was performed more with an audience present. Additionally, the sex of the audience seemed to be influential, reflected by a difference in the frequency of interactant‐directed behaviors when a female vs. a male audience was present. This study is one of the first to demonstrate that females modify their behavior as a result of being watched and stresses the importance of examining audience effects in a variety of social contexts.     

Features of the aggressive behavior of victorious mice in inter-male interactions

Latency of the first attack, number of attacks and total attacking time in aggressive male C57BL/6J strain mice were studied during agonistic interactions with submissive mice which had a great defeat experience. Then, aggressive mice were placed together in order to find out which will be the winner in each pair. It has been shown that males with a shorter latency of the first attack and greater total attacking time gained the victory.     

Fighting between two females for a male in the hoolock gibbon

I observed fighting between two adult females for an adult male in a group of hoolock gibbons (Hylobates hoolock)during fieldwork conducted between November 1988 and December 1990 at Lawachara in the West Bhanugach Reserve Forest, Sylhet, Bangladesh. I discuss the history of the episodes and the consequences of fighting.     

Draglines and Assessment of Fighting Ability in Cannibalistic Jumping Spiders

The frequency of injury and death during female-female aggression varies in the jumping spider genus Portia, with interactions being more violent (likely to end in death or injury of one of the combatants) in P. labiata (from Sri Lanka) than in another two species (P. fimbriata from Australia and P. schultzi from Kenya). To investigate the role of draglines in the assessment of fighting ability, two types of tests were carried out: 1) dragline discrimination and 2) mirror image response (Portia's reaction to mirror images is comparable to interaction with conspecific rivals). For both types of testing, triplets of equal-size conspecific females were used: one female (the test spider) was exposed to draglines of two equal-size conspecific females they had not encountered before (donor spiders). The fighting abilities of donor spiders were determined directly by staging intraspecific contests between them. In dragline-discrimination tests (spider placed in petri dish containing draglines from two conspecific females with different fighting ability), females of P. labiata, but not the other two species, avoided draglines of the superior fighter (i.e., they spent the majority of their time on draglines of donor spiders with lesser fighting ability). For mirror-image testing, the test spider was placed in a petri dish containing a mirror and draglines. Each test spider was tested on two successive days, with donor draglines in the two tests coming from conspecific females with different fighting ability. In these tests, females of P. labiata (but not the other two species) spent less time embracing (each spider pressing its forelegs, palps and front of body against the other spider) and more time in a part of the petri dish where view of the mirror was obstructed when on the draglines of donor spiders with greater fighting ability than when on the other conspecific's draglines. Findings from this study suggest that P. labiata females use signpost cues associated with draglines to assess the relative fighting abilities of unknown opponents.