Body Size, Leg Autotomy, and Prior Experience as Factors in the Fighting Success of Male Crab Spiders, Misumenoides formosipes

Game theory predicts which characteristics of male crab spiders should affect their success in mate-guarding contests. Previous data on Misumenoides formosipes suggested that body size and leg number influenced fight outcome, but the study had methodological problems. Using improved methods, we corroborated the size advantage but refuted the disadvantage of leg reduction. We also conducted the first test for an invertebrate on the effect that a single contest has on the outcome of subsequent contests. When size and residency status were controlled, prior winners defeated prior losers after only one contest experience. We discuss the importance of controlling for previous experiences in studies of contest resolution and the ramifications for both past and future studies.     

Inter-male competition and mating success in the field cricket,Gryllus bimaculatus (de Geer)

Males of the field cricket Gryllus bimaculatus compete aggressively for the acquisition of burrows and females. The effect of population density on the nature of encounters between males, and factors affecting individual competitive ability were examined. Aggression was reduced in high population densities, both in terms of the proportion of encounters leading to aggression and in the intensity of aggressive disputes. The addition of burrows and females increased aggressiveness and removed population density effects. Individual competitive ability was determined primarily by body size and secondly, by an individual's past experience of winning (‘confidence’). Competitive ability, prior residence in burrows and the presence of females all appear to influence the outcome of aggressive disputes. Variation in calling frequency of males was influenced both by population density and competitive ability; males who called more often had a greater encounter rate with females. Females were more likely to mate with burrow residents. However, amongst residents, larger males gained more matings. The potential for female choice in this species is discussed.     

Rats Benefit from Winner and Loser Effects

Prior fighting experience of opponents can influence the outcome of conflicts. After a victory, animals are more likely to win subsequent contests, whereas after a defeat animals are more likely to lose, regardless of the identity of opponents. The underlying mechanisms and the adaptive significance of these winner and loser effects are as yet unknown. Here, we tested experimentally whether agonistic behavior of male wild‐type Norway rats is influenced by social experience, and we investigated whether this might reduce fighting costs (duration of contest, risk of injury) in subsequent encounters. Rats were randomly assigned to receive either a losing or a winning experience and subsequently tested with unfamiliar, naïve opponents. We found that most rats with a winning experience won the subsequent encounter, and all rats with a losing experience lost the next contest. Previous winners attacked more rapidly in the subsequent encounter and reduced their aggressive behavior sooner; the contests were decided more quickly, which saved time and behavioral effort to the winner. Previous losers received less aggression in the next encounter, despite emitting fewer submissive ultrasonic calls than in the preceding contest, thereby reducing the risk of being injured by the opponent. Thus, anonymous social experience influenced rats’ subsequent behavior toward size‐matched, naïve, unknown social partners. Furthermore, apparently, they benefit from showing winner and loser effects in intraspecific contests by saving time, energy, and risk.     

Food availability in beach and marsh habitats and the size of the fiddler crab claw,a sexually selected weapon and signal

Males of the sand fiddler crab Uca pugilator possess a greatly enlarged claw that is used as a weapon in ritualized contests for control of breeding burrows and is waved to attract females to breeding burrows. Approximately 5400 crabs were collected along the Atlantic coast of North America at 14 localities, all of which had both beach and salt marsh habitats. Five measurements were made on each claw. Principal components analysis was used to generate a single measure of claw size from the seven correlated measures and scores of the claw. Carapace width was measured to index body size. Claw size was greater in beach than marsh habitats, controlling for body size. However, body size did not differ by habitat type. Claw size was also greater in laboratory‐reared males receiving more food, suggesting that differential access to food could influence habitat‐associated differences in claw size. Chlorophyll a concentration and total organic content, reflecting, respectively, the abundance of benthic algae and other food, were greater in beach than marsh habitats. Moreover, feeding opportunities were greater in the wetter beach habitat because crabs there, but not in marsh habitat, can feed at breeding burrows. Adult fiddler crabs continue to molt and grow in both body and claw size as they age. Energetic investment in the claw relative to the body is plastic. It appears that the availability of food can affect the amount of energy invested in the claw.     

Territorial interactions between larvae of the dragonfly Pyrrhosoma nymphula: outcome of encounters

The behaviour of final-instar larvae of the aquatic predator Pyrrhosoma nymphula (Odonata: Coenagrionidae) during laboratory-staged territorial interactions is described. Occupants, as distinct from intruders, won 72% of encounters. The behaviour of occupants during contests won by intruders was more like that of intruders during all contests than that of occupants during contests won by occupants. Contest outcome was little affected by either size differences between the contestants or the use of the Labial strike. Contest duration was not correlated with outcome, with size differences between contestants or with the use of the Labial strike. Winners and losers differed significantly in the number of acts used during encounters won by occupants (losers using more acts), but not during encounters won by intruders. Winners and losers also differed in their use of the behavioural acts Slow wave, Lateral display and Lamellae swipe during contests won by occupants, but not during contests won by intruders. The results are discussed in terms of the asymmetric war of attrition.     

Insight into invasion: Interactions between a critically endangered and invasive crayfish

Invasive species are known to competitively displace native species via direct behavioural interactions, or contests, which can result in limited access to valuable resources such as food or shelter for the native. Here, we aimed to determine the dynamics and outcome of agonistic contests between the critically endangered Fitzroy Falls Spiny Crayfish (Euastacus dharawalus) and the invasive yabby (Cherax destructor) in a controlled laboratory setting. Although C. destructor has been listed as a key threatening process for E. dharawalus, the magnitude and nature of its impact have not been formerly assessed. We recorded the frequency and intensity of aggressive interactions between the species, as well as the winner of the contest and subsequent dominance during staged encounters. In addition to this, we recorded the time individuals spent consuming food and actively moving on the substratum. Contrary to our predictions, the frequency of aggressive behaviours performed by the two species did not significantly differ, but E. dharawalus engaged in more intensely aggressive behaviours and was less submissive than C. destructor. As a result, E. dharawalus won significantly more contests than its invasive rival and subsequently achieved a higher dominance score overall. Despite this result, C. destructor spent significantly more time consuming food than did E. dharawalus. These results suggest that dominance in agonistic contests did not translate into dominance over food resources in our study. Ultimately, reduced access to food, combined with the costs of agonistic contests including injury and the relatively low reproductive rates of E. dharawalus indicate that C. destructor may have a negative effect on the survival and persistence of the critically endangered E. dharawalus.     

Information and Patch Exploitation Strategies of the Eastern Chipmunk,Tamias striatus (Rodentia: Sciuridae)

We examined to what extent 15 chipmunks foraging from dishes in the field varied load sizes depending on a predetermined sequence of resource renewal/removal. At 10-m distances from burrows, chipmunks collected smaller loads when seeds were replenished over sequential trials and larger load sizes when trials were interrupted by removing all seeds for one visit. Similar sequence effects were less obvious at 0 m. Load size also varied with travel time but not in a manner consistent with either long-term rate maximization or resource tracking. When travel time effects were removed, sequence effects became more pronounced at distances of both 0 and 10 m. The results suggest that chipmunks use past experience in patches to gauge expected returns in the future, and that they devalue resources according to their uncertainty (“future discounting”). That chipmunks varied load sizes with food renewal/removal more at 10 than 0 m from burrows suggests that the degree to which chipmunks devalue future resources may depend on how defendable those resources are.     

Territoriality and microhabitat selection in two intertidal New Zealand fish

Aspects of territoriality and microhabitat selection in Acanthoclinus fuscus and Forsterygion nigripenne robustum were studied in intra- and interspecific confrontation experiments under tidal and constant conditions. In both species aggressive responses were closely related to activity with maximal frequencies during the first 2 h of the experiments. Under tidal conditions activity was higher than at constant water level. Size predominantly affected contest outcome in both species. Initiation of the first bout and past experience were further important factors, whereas prior residence did not influence dominance relationships. Dominant individuals defended access to shelter and low tide pools as critical resources. A. fuscus preferred tubes matching the body size for cover, whereas F. n. robustum spent more time in caves and on open areas. Furthermore, A. fuscus preferred shallower water and was able to sustain long periods out of water. The different microhabitat preferences may reduce interspecific competition. Although A. fuscus did not win significantly more trials, higher aggressive tendencies were observed in this species.     

Size,sex and social experience: Experimental tests of multiple factors mediating contest behaviour in a rockpool fish

Aggressive contests amongst conspecifics are important to understand from an ecological and evolutionary perspective as contest dynamics can directly influence individual fitness. For some species, individual attributes such as relative body size closely predict the outcome, intensity and duration of contests, whereas for others, prior social experiences play a key role. However, disentangling the relative effects of individual attributes and social experiences is challenging from an experimental perspective, and because of this, few studies have investigated relative effects of multiple factors. Rockpool fishes have been well studied in terms of factors governing abundance, distribution and community structure, but much less so in terms of contest behaviour. This is surprising because contest dynamics are likely to directly affect the distribution of fishes along the rocky shore, and hence indirectly govern population and community composition. Here, we explored multiple factors potentially influencing contest behaviour in a numerically dominant, resident intertidal fish species, Bathygobius cocosensis (Gobiidae). Using a series of staged pairwise contest trials, we investigated the effect of size, sex and social experiences (namely prior residency and winner–loser experiences) on contest dynamics. We found no evidence that prior residency influenced contest outcome, suggesting social experiences play a minor role in contest dynamics. Previous winner/loser experience also did not influence contest outcome, although this is likely a product of low sample size. In addition, the likelihood of winning was unrelated to contestant sex, and the combination of sexes in paired contests did not influence contest intensity or duration. Instead, body size was related to contest outcome, intensity and duration in the majority of experimental trials. These results suggest that body size rather than sex or social experiences is the key determinant of contest dynamics in this species. We suggest that the dynamic biotic and abiotic environment inhabited by intertidal fish may reduce the influence of prior social experiences in modulating contest dynamics.     

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.     

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.     

Contest dynamics for food and reproductive resources are defined by health condition in a dung beetle

Male dung beetles compete to obtain food and females, and early resource recognition and appropriation increase the probability of mating. The outcome of such encounters is expected to be defined by self‐ and the opponent’s health condition. In this study, we analyzed the effect of body condition and immune defense on the contest dynamics between males that rolled a food ball with a partner female (owner males) and intruder males of dung beetle Canthon cyanellus. Body condition was measured as body size, body dry mass, lipid mass, and muscle mass; immune defense was estimated via phenoloxidase activity. Owner males with higher lipid mass contacted the food ball significantly earlier than owner males with lower lipid mass. Individuals with lower phenoloxidase activity started to roll food balls earlier than individuals with higher phenoloxidase activity. Owner males that had higher body dry mass, compared to female partners, began to roll the food ball significantly earlier than male–female pairs with lower differences in dry mass. Heavier males won significantly more contests than lighter males. Our results suggest that the health condition is a key factor related to the dynamics and outcome of male–male contests for resources and females in C. cyanellus. Consequently, differences in individual condition are main determinants of contest outcomes in dung beetles.     

The influence of contests on optimal clutch size: a game-theoretic model

We develop a game-theoretic model to predict the effect of size-dependent contest outcomes on optimal-clutch-size decisions. We consider the case where larger individuals develop from smaller clutches and, as adults, are advantaged in competition for limiting resources. The relationship between fitness and size thus depends on the sizes of other members of the population. We show that clutch-size optima are decreased by body-size-dependent contest outcomes, with larger effects when body size is most affected by clutch size, when prior resource ownership has less influence on contest outcome and when contests occur more frequently. We also show the existence of polymorphisms in clutch-size optima and that clutch-size driven changes in population density can, via an effect on the probability of host finding, further influence optimal clutch size. Our model is formulated to match the life history of a parasitoid wasp, in which clutch size affects offspring size and females engage in direct contests for host ownership, which larger females tend to win; we confirm that female-female competition is likely to influence clutch size in this species. However, the model is also relevant to clutch size in other taxa and supports recent suggestions concerning reproductive decisions in great tits.     

Behavioural and ecological investigations of the crabOcypode ceratophthalmus (Crustacea: Ocypodidae)

Observations conducted at different places along the Moçambique coast indicate that the behaviour of Ocypode ceratophthalmus is extremely plastic, permitting a high degree of adaptation to the often vastly different shores on which they are found. When the crabs are nocturnal and when, as is usual, the burrows are intertidally placed, the greater part of the crab's life is spent beneath the sand and the periods of activity are much influenced by the tidal cycles. Apart from feeding on items of prey conveyed into the burrows and stridulating when disturbed there is no evidence that they are anything but inactive when underground. On emergence from the sand the sole major activities observed were searching for food and the digging and clearing of burrows. They are very aggressive and although cannibalism is common, conflicts between equal sized crabs are generally ritualized. Social behaviour appears limited and, except for protest stridulation and ritualized posturing during intraspecific encounters, none was observed.     

Competition for resources mediated by intrinsic social dominance in sympatric finches

Gouldian Erythrura gouldiae and Long‐tailed Finches Poephila acuticauda are morphologically and ecologically similar sympatric species, for which social dominance relationships are likely to determine access to critical limiting resources. Building on previous research showing that Long‐tailed Finches dominate competition for nest‐sites, I staged dyadic contests for food between the two species in captivity in order to test social dominance dynamics under controlled conditions. Long‐tailed Finches were likely to dominate interactions with Gouldian Finches and expressed higher levels of aggression while competing for access to food. These results suggest a stable dominance relationship between the two species which could be affecting Gouldian Finches' access to food resources in the wild, potentially constraining their ability to recover from recent population declines.     

Power to Hold Sheltering Burrows by Juveniles of the Signal Crayfish,Pasifastacus leniusculus

Juveniles of the signal crayfish reside during daylight hours in shelters. At twilight they leave for food, at daybreak they either return or find another shelter. We examined conflicts over burrow ownership. At low densities with equal numbers of shelters and crayfish 1/5 of the burrows were occupied. Increasing both the number of crayfish and shelters improved the occupancy close to 50%. Doubling the number of crayfish in relation to the number of shelters increased the occupancy up to 75%. In an experiment with 30 randomly selected crayfish and 15 sheltering holes available the burrow holders were about 1–2 mm longer in carapace length than those found freely moving in the aquarium. Shelter owners were rarely newly moulted individuals. Size asymmetry (2 mm difference in carapace length) between owner and intruder affected the outcome of the contest over burrow ownership. When intruders were larger than owners, takeovers occurred in about 80% of the cases tested. If the owner was large and the intruder small the takeover frequency was about 20%. When the owner and the intruder were of matching size takeovers were still observed in about 45% of the cases. The outcome of the ownership contest has a true meaning. In a nursery-pond experiment, where low-protein food was randomly scattered all around, no size differences were found in carapace lengths of crayfish juveniles residing in different shelters. However, when high-protein food was introduced in a single spot, owners of burrows were significantly larger and more numerous in the nearby shelters than in the shelters furthest away from the food source. Therefore, when food is unevenly distributed the burrow ownership contests may potentially lead to size asymmetries between individuals. This may lead to large individuals nearby the food growing faster than small individuals further away from the food source, a process likely to further enhance size differences.     

Condition,not eyespan,predicts contest outcome in female stalk‐eyed flies,Teleopsis dalmanni

In contests among males, body condition is often the key determinant of a successful outcome, with fighting ability signaled by so‐called armaments, that is, exaggerated, condition‐dependent traits. However, it is not known whether condition and exaggerated traits function in the same way in females. Here, we manipulated adult condition by varying larval nutrition in the stalk‐eyed fly, Teleopsis dalmanni, a species in which eyespan is exaggerated in both sexes, and we measured the outcome of contests between females of similar or different body condition and relative eyespan. We found that females in higher condition, with both larger bodies and eyespan, won a higher proportion of encounters when competing against rivals of lower condition. However, when females were of equal condition, neither eyespan nor body length had an effect on the outcome of a contest. An analysis of previously published data revealed a similar pattern in males: individuals with large relative eyespan did not win significantly more encounters when competing with individuals of a similar body size. Contrary to expectations, and to previous findings in males, there was no clear effect of differences in body size or eyespan affecting contest duration in females. Taken together, our findings suggest that although eyespan can provide an honest indicator of condition, large eyespans provide no additional benefit to either sex in intrasexual aggressive encounters; body size is instead the most important factor.     

Fighting Strategies in Crayfish Orconectes rusticus (Decapoda, Cambaridae) Differ with Hunger State and the Presence of Food Cues

Crayfish, bearing dangerous weapons in the form of chelae, resolve intraspecific conflicts using stereotyped behaviors and structured, escalated encounters. According to predictions of game theory models, any decision to resort to unrestrained combat without prior careful behavioral assessment of the opponent's fighting abilities carries great risks. The present study examines the significance of internal hunger states and the presence of chemical food cues in this decision process using a 2 × 2 factorial design. Hungry crayfish escalated more rapidly, and thus took greater risks, during agonistic encounters, while the presence of a food source reduced the rate at which fights increased in intensity. However, there were no significant differences in fighting behavior as a result of the interaction between these two variables. We then address the complex trade-offs that individuals face in fighting with respect to increased risks of injury, appetitive states, and opportunities for resource access.     

Is male homosexuality maintained via kin selection?

We tested the kin selection hypothesis of male homosexuality using questionnaire data in a community sample of homosexual and heterosexual men. Homosexual men were no more likely than heterosexual men to channel resources toward family members. Indeed, heterosexual men tended to give more financial resources to siblings than homosexual men. Furthermore, homosexual men were somewhat more estranged from family members, especially from fathers and oldest siblings.     

Dominance hierarchies in group-living cleaning gobies: causes and foraging consequences

In species living in social groups, aggression among individuals to gain access to limiting resources can lead to the formation of stable social hierarchies. We tested whether dominance rank in social groups of sponge-dwelling cleaning gobies Elacatinus prochilos in Barbados was determined by physical attributes of individuals or by prior experience of dominance, and examined the foraging consequences of dominance rank. Intraspecific aggression within groups resulted in stable dominance hierarchies that were strongly correlated with fish length. Dominant individuals maintained exclusive territories while subordinate fish occupied broader home ranges. Larger, competitively dominant fish were able to monopolize areas inside the sponge lumen with the highest abundance of the polychaete Haplosyllis spp., a favoured prey item, and achieved the highest foraging rates. The removal of a territorial individual from large groups resulted in a domino-like effect in territory relocation of the remaining fish as individuals moved to the territory previously occupied by the individual just above them in the group hierarchy. Individuals added to existing groups generally failed to gain access to territories, despite being formerly dominant in their original groups. When given the opportunity to choose a location in the absence of larger competitors, gobies frequently preferred positions that were previously defended and that had abundant food. These results suggest that intraspecific competition for resources creates the observed dominance structures and provides support for the role of individual physical attributes in the formation and maintenance of dominance hierarchies.