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.     

Inter-male aggression with regard to polygynous mating system in Pampean grassland mouse, Akodon azarae (Cricetidae: Sigmodontinae)

Based on the hypothesis that, in Akodon azarae, polygyny operates through female defence, we studied inter-male aggression in order to test the following predictions: during the breeding period (1) resident males are more aggressive than intruder males in the presence of females (FP), and (2) aggressive behaviour is independent of male condition (resident or intruder) in the absence of females (FA). To test our predictions, we used the resident male behavioural response towards an intruder male in relation to FP or FA. We conducted 30 encounters in FP and 27 in FA in 0.79-m² round enclosures placed in the Espinal Reservation. Our results support the prediction that, in FP, the intensity of aggressive behaviour exhibited by males varied in relation to resident or intruder condition. Resident males showed high levels of aggression towards intruders, and intruders exhibited the greatest values of submissive behaviours with residents. In FA, the intensity of aggressive behaviour did not vary in relation to resident or intruder condition. Both resident and intruder males exhibited low aggressive behaviour and inter-male encounters resulted mainly in non interactive behaviours. Our results support the hypothesis that, in A. azarae, the polygynous mating system operates through female defence.     

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.     

Behaviour in a population of Columbian ground squirrels, Spermophilus columbianus columbianus

Field observations and films of laboratory encounters were used to describe and quantify behaviour in a population of Columbian ground squirrels. Activity correlated with ambient temperature. Feeding, the most frequent behaviour, decreased during mid-day whereas other behavioural patterns increased. Squirrels nearly always seemed alert, even during other behavioural patterns. Three of seven vocalizations indicated alarm. Possible adult scent-marking nerver exceeded 1 per cent of above-ground activity and decreased seasonally. Juvenile play was frequent; yearling play became more aggressive and was replaced by aggression by mid-season. Adult aggression rarely involved contact; aggression was most frequent during breeding and lactation among males and females, respectively. The behaviour of this population was similar to that observed in other populations of the same and other sciurids. Most differences cannot be explained without ecological investigation of the other populations.     

Effect of male-to-male aggressiveness and feed-restriction during rearing on sexual behaviour and aggressiveness towards females by male domestic fowl

This experiment was one part of a larger study investigating problems of aggression towards females by male broiler breeder fowl. To investigate causal mechanisms, we were interested in determining (1) if feed-restriction during rearing affects behaviour towards females at sexual maturity and (2) if aggressiveness towards females is correlated with general levels of aggressiveness. We compared broiler breeder males with commercial laying strain males, which were either fed ad libitum or were feed-restricted during the rearing phase, and with game strain males, bred for fighting. Differences in behaviour were determined by observing males during interactions with small groups of females.Laying strain males did not behave aggressively towards females, whether feed-restricted or fed ad libitum during rearing. Despite genetic selection for fighting ability, game strain males also were not aggressive towards females. Conversely, broiler breeder males displayed significantly higher levels of aggression towards females than did feed-restricted laying strain males (P<0.02). Broiler breeder males were rough with females during mating, whereas laying strain and game strain males were not. Females struggled more frequently during mating attempts by broiler breeder males (P<0.002) and interfered frequently when these males attempted to mate with other females.From our results, we conclude that (1) feed-restriction during rearing has little effect on the sexual and aggressive behaviour of laying strain males at maturity and (2) selection for aggressiveness has not resulted in males which are more aggressive to females. Aggression towards females appears to be a unique problem occurring in broiler breeder male strains and not a function of feed-restriction.     

Short Amplexus Duration in a Territorial Anuran: A Possible Adaptation in Response to Male-Male Competition

Mating duration is a reproductive behaviour that can impact fertilization efficiency and offspring number. Previous studies of factors influencing the evolution of mating duration have focused on the potential role of internal sperm competition as an underlying source of selection; most of these studies have been on invertebrates. For vertebrates with external fertilization, such as fishes and frogs, the sources of selection acting on mating duration remain largely unknown due, in part, to the difficulty of observing complete mating behaviours in natural conditions. In this field study, we monitored breeding activity in a population of the territorial olive frog, Rana adenopleura, to identify factors that affect the duration of amplexus. Compared with most other frogs, amplexus was short, lasting less than 11 min on average, which included about 8 min of pre-oviposition activity followed by 3 min of oviposition. We evaluated the relationship between amplexus duration and seven variables: male body size, male condition, operational sex ratio (OSR), population size, clutch size, territory size, and the coverage of submerged vegetation in a male’s territory. We also investigated the influence of these same variables, along with amplexus duration, on fertilization rate. Amplexus duration was positively related with clutch size and the degree of male-bias in the nightly OSR. Fertilization rate was directly related to male body size and inversely related to amplexus duration. Agonistic interactions between males in amplexus and intruding, unpaired males were frequent. These interactions often resulted in mating failure, prolonged amplexus duration, and reduced fertilization rates. Together, the pattern of our findings indicates short amplexus duration in this species may be an adaptive reproductive strategy whereby males attempt to reduce the risks of mating and fertilization failures and territory loss resulting from male-male competition.     

Familiarity and flexible mating strategies of a solitary rodent, Dipodomys ingens

Because mating is a product of individual reproductive strategies that may vary with changing conditions, we predicted variable mating behaviour in an arid-adapted, territorial rodent, the giant kangaroo rat, Dipodomys ingens. We also predicted that familiarity would facilitate nonaggressive courtship and mating in this solitary rodent. Through direct observations in the field, we found that mating varied from exclusive to multiple partners. Where densities were low, and on nights when multiple females were in oestrus, each animal mated with one member of the opposite sex. In conditions where the operational sex ratio was skewed towards multiple males, males footdrummed and competed for females. Males were able to mate with one or two females in adjacent territories, and they successfully competed for these same females throughout the breeding season. Females that mated exclusively with one male had more pups emerge from the burrow compared with females that experienced male competition. Females allowed nearest neighbour males to enter their burrows, and they engaged in more nonaggressive contact with close neighbours than with other males. Paired encounters in the field showed less aggression towards neighbours than strangers. In laboratory tests, females were less aggressive towards and allowed more contact with familiar than unfamiliar males. These results show that D. ingens can alter mating strategies as conditions change. Familiarity is an important factor in nonaggressive interactions between males and females and may be important to mate preferences in females during reproduction. The less aggressive behaviour to neighbours than to strangers (‘dear enemy’ phenomenon) is consistent with other solitary animals that defend multipurpose territories. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Aggression in the early phases of the reproductive cycle of the male three-spined stickleback (Gasterosteus aculeatus)

Male sticklebacks with nests were more aggressive towards other males than were males which had yet to build nests.Males with nests were more aggressive towards other males than towards non-gravid females. This was not true of fish that had yet to build nests.The increased aggression shown to other males by males with nests was reduced after gonadectomy to the level characteristic of fish that had yet to build nests. This reduction occurred in the first week after the operation. Gonadectomy had no immediate effect on the aggression shown to non-gravid females.The aggression of males that had still to build nests was not reduced by gonadectomy.Gonadectomy resulted in a complete absence of nest building and nest-directed behaviour.     

Sex-Specific Aggression and Antipredator Behaviour in Young Brown Trout

Sex differences in adult behaviour are often interpreted as consequences of sexual selection and/or different reproductive roles in males and females. Sex-specific juvenile behaviour, however, has received less attention. Adult brown trout males are more aggressive than females during spawning and juvenile aggression may be genetically correlated with adult aggression in fish. We therefore tested the prediction that immature brown trout males are more aggressive and bolder than immature females. Because previous work has suggested that precocious maturation increases dominance in salmonids, we included precocious males in the study to test the prediction that early sexual maturation increase male aggression and boldness. Aggression and dominance relations were estimated in dyadic contests, whereas boldness was measured as a response to simulated predation risk using a model heron. Independent of maturity state, males initiated more than twice as many agonistic interactions as females in intersexual contests. However, males were not significantly more likely to win these contests than females. The response to a first predator attack did not differ between sex categories, but males reacted less to a second predator attack than females. Sexual maturity did not affect the antipredator response in males. Since there is no evidence from field studies that stream-living immature male and female salmonids differ in growth rate, it appears unlikely that the sex differences demonstrated are behavioural consequences of sex-specific investment in growth. It seems more likely that sex-specific behaviour arises as a correlated response to sexually selected gene actions promoting differential behaviour in adult males and females during reproduction. Alternatively, sex differences may develop gradually during juvenile life, because a gradual developmental program should be less costly than a sudden behavioural change at the onset of sexual maturity.     

Aggression and worker control of caste fate in a multiple-queen wasp, Parachartergus colobopterus

Although famously cooperative, social insect colonies harbour considerable potential for genetic conflict among colonymates. This conflict may be expressed behaviourally as aggression by workers. We investigated aggression in 34 colonies of the wasp Parachartergus colobopterus, by evaluating the characteristics of both instigators and victims of aggressive interactions. We estimated genetic relatedness and queen number using DNA microsatellites and found that workers and emerging females should be most in conflict over the caste of the latter when there are many queens on the nest. We found that aggressive interactions are more likely to involve older workers attacking either males or younger workers, and that victim and aggressor females have more ovarian development than randomly sampled colonymates. Moreover, mated females with low levels of ovarian development relative to active queens were also more likely to be aggressors and victims than were randomly sampled females. Aggression among females supports the hypothesis that older workers use aggression towards younger females as a means of policing the development of emerging females into queens. Workers also may use aggression to suppress the reproduction of some mated females. Our findings thus support the hypothesis that genetic conflicts of interest motivate worker aggression in swarm-founding wasp colonies.     

AGONISTIC AND SPACING BEHAVIOUR OF THE NOISY MINER MANORINA MELANOCEPHALA, A COMMUNALLY BREEDING HONEYEATER

The Noisy Miner is a communally breeding Australian honeyeater in which several males feed the offspring of a single female. They reside year-round in colonies, which may number several hundred birds. Miners within colonies unite to mob predators and are successful in defending their colony area against all other species of birds. The species is also highly aggressive intraspecifically. Individuals in a colony in southeastern Queensland were present most of the time in small activity spaces. Most resident males showed extensive overlap of activity space from one season to the next. Males did not defend their activity spaces, so that coalitions of birds occurred whose membership changed with place and time. Certain assemblages of males, termed coteries, were of a more permanent nature. Coterie members showed aggression towards outsiders at border regions. Females' activity spaces were much smaller and less variable than those of males. They showed almost no overlap and were probably maintained by mutual avoidance. Females tended to nest within the activity space they occupied shortly before the breeding period. Thus the spacing of males and females within a colony was quite different. Interactions, often agonistic, were frequent between individuals, between an individual and a group, and between groups. Encounters involving two males in which participants normally lived farther apart were more often agonistic than when participants lived more closely, and more aggression was seen within coteries than between them. When larger groups of birds had agonistic encounters, they more frequently involved birds from different coteries. Very little male-female aggression was seen. Interactions between males and females of different coteries were usually sexual, sometimes involving attempted copulation. Colony sizes are probably too large to permit individual recognition of fellow members, but this could be more likely in coteries. Many males in coteries are doubtless closely related, but outsiders frequently infiltrate them. Coteries are not reproductive units.     

Adaptive significance of permanent female mimicry in a bird of prey

Permanent female mimicry, in which adult males express a female phenotype, is known only from two bird species. A likely benefit of female mimicry is reduced intrasexual competition, allowing female-like males to access breeding resources while avoiding costly fights with typical territorial males. We tested this hypothesis in a population of marsh harriers Circus aeruginosus in which approximately 40 per cent of sexually mature males exhibit a permanent, i.e. lifelong, female plumage phenotype. Using simulated territorial intrusions, we measured aggressive responses of breeding males towards conspecific decoys of females, female-like males and typical males. We show that aggressive responses varied with both the type of decoys and the type of defending male. Typical males were aggressive towards typical male decoys more than they were towards female-like male decoys; female-like male decoys were attacked at a rate similar to that of female decoys. By contrast, female-like males tolerated male decoys (both typical and female-like) and directed their aggression towards female decoys. Thus, agonistic responses were intrasexual in typical males but intersexual in female-like males, indicating that the latter not only look like females but also behave like them when defending breeding resources. When intrasexual aggression is high, permanent female mimicry is arguably adaptive and could be seen as a permanent 'non-aggression pact' with other males.     

Does signalling mitigate the cost of agonistic interactions? A test in a cricket that has lost its song

Prevailing models of animal communication assume that signalling during aggressive conflict mitigates the costs of fighting. We tested this assumption by staging dyadic encounters between male field crickets, Teleogryllus oceanicus, under three conditions: (i) both males could sing aggressive songs, (ii) neither male could sing, and (iii) one male could sing but the other could not. We conducted experiments on males from a Hawaiian population from Kauai that has recently evolved signal loss, and males from a Hawaiian population from the Big Island that has not. Among both populations, interactions between two silent males were characterized by higher levels of aggression than interactions involving one or two singing males. Because the level of aggression is strongly related to the cost of fighting, these data demonstrate that signalling mitigates the cost of fighting. In mixed trials, we found no statistically significant differences between the behaviour of calling and non-calling males in either population. We conclude that there is no evidence that the Kauai population exhibits special adaptations to alleviate the costs of signal loss. Finally, we found that males were much more likely to signal after their opponent''s retreat than after their own retreat. Aggressive song therefore meets the definition of a ‘victory display’.     

Territorial behaviour in corn mice, <Emphasis Type="Italic">Calomys musculinus</Emphasis> (Muridae: Sigmodontinae), with regard to mating system

We studied territorial behaviour of Calomys musculinus, a promiscuous-polygynous species, in order to test the predictions that during the breeding period, home-range-owner females are more aggressive than home-range intruders and that males do not display territorial behaviour. At the core of the breeding season, we conducted 136 intrasexual and 66 intersexual encounters. The study was carried out in four 0.25-ha enclosures, each containing one independent population. We studied territorial behaviour utilising focal animal sampling to record behaviour. Encounters were conducted in a circular opaque arena (COA) near the home-range’s activity mathematical centre (AMC). Amicable behaviour was not observed in any of the 78 trials among females. The rates of aggressive and submissive behaviour per female–female encounters varied in relation to whether they were home-range owners or intruders, the former being aggressive, the latter submissive. So we could link these drives to the territorial behaviour of C. musculinus. This result fits well with the already known territoriality of C. musculinus during the breeding season. In contrast, the 58 intrasexual male encounters did not show direct aggressive interactions between them. The rates of the different behaviours between males did not vary in relation to whether they were home-range owners or intruders. In our study, which took place during the breeding period, C. musculinus males were nonterritorial, and they never exhibited aggressiveness towards females.     

The effects of Trichinella infection on intersexual interactions between mice

Parasitic infection may influence mating behaviour in at least two ways: (1) by creating a selection pressure for mate choice based on apparent resistance to infection and (2) by reducing the likelihood of an infected individual mating through induced changes in its physiology or behaviour. These two possibilities were tested using a nematode infection in laboratory mice. Female CFLP mice were infected with larvae of the aphasmid nematode Trichinella spiralis and allowed to interact with uninfected males which were either experienced or inexperienced in term of having previously copulated. Both the responses of males towards infected and uninfected females and the responses of females towards males were recorded. Overall, males were significantly less likely to mount, copulate and show other behaviours related to copulation with infected females. When experienced and inexperienced males were considered separately, experienced males showed significantly more investigatory activity towards infected females, but infection did not appear to affect responses by inexperienced males. Infected females were more likely to avoid or respond aggressively to attempts to mount by males and the latency to the first attempt to mount correlated positively with the frequency of avoidance/aggression towards approaching males. The results suggest that infection in females influences intersexual interactions by inducing a disinclination to accept mounts and copulate. The functional implications for parasite transmission are considered.     

Female aggression predicts mode of paternity acquisition in a social lizard

Individual differences in behaviour are ubiquitous in nature. Despite the likely role of selection in maintaining these differences, there are few demonstrations of their fitness consequences in wild populations and, consequently, the mechanisms that link behavioural variation to variation in fitness are poorly understood. Specifically, the consequences of consistent individual differences in behaviour for the evolution of social and mating strategies have rarely been considered. We examined the functional links between variation in female aggression and her social and mating strategies in a wild population of the social lizard Egernia whitii. We show that female Egernia exhibit temporally consistent aggressive phenotypes, which are unrelated to body size, territory size or social density. A female''s aggressive phenotype, however, has strong links to her mode of paternity acquisition (within- versus extra-pair paternity), with more aggressive females having more offspring sired by extra-pair males than less aggressive females. We discuss the potential mechanisms by which female aggression could underpin mating strategies, such as the pursuit/acceptance of extra-pair copulations. We propose that a deeper understanding of the evolution and maintenance of social and mating systems may result from an explicit focus on individual-level female behavioural phenotypes and their relationship with key reproductive strategies.     

Inter-group encounters and female transfer in mountain gorillas: Influence of group composition on male behavior

This paper examines the behaviors used by male gorillas (G. gorilla beringei) to influence female choice during inter-group encounters. Encounters are related to the acquisition of females rather than to the defense of a group's range. Data on 58 independent encounters confirms that encounters are generally aggressive, but that contact aggression is less frequent than previously reported. A high number of potential migrants in one of the groups involved, rather than the newness of either group, predicts the likelihood of contact aggression between males. Encounters with a newly formed unit last longer than encounters involving established groups. Male herding serves to prevent female transfer. Females without dependent offspring are more likely than others to be herded. Proceptive females are the target of herding more often than non-proceptive cycling females. Groups where herding has been observed are more likely to be newly formed, and they include significantly more males. This last result confirms that there are advantages to male cooperation in gorillas. It also raises the question of why such cooperation does not occur more often. © 1993 Wiley-Liss, Inc.     

`Little Chipmunk' Syndrome? Male Body Size and Dominance in Captive Yellow‐Pine Chipmunks (Tamias amoenus)

In mammals, large males are often assumed to have higher mating success because they have greater success at contest competition. This relationship is often used to explain the prevalence of male-biased sexual size dimorphism in mammals. However, in many small vertebrates, large individuals are not always dominant. Using staged dyadic encounters, we examined the relationship between male body size and social dominance in captive male yellow-pine chipmunks ( Tamias amoenus ), a species with female-biased sexual size dimorphism. The yellow-pine chipmunk has a mating system in which males participate in mating chases and dominant males may have an advantage in acquiring matings with oestrous females. Captive male chipmunks were aggressive in only 28% of 144 paired encounters; however, several lines of evidence indicated that smaller chipmunks were dominant over large chipmunks: (1) small males were dominant in more dyads than large males; (2) within dyads, dominant males were smaller than subordinate males; and (3) small males performed more aggressive behaviour than large males. These results are not consistent with the prediction that large males are typically dominant. If large chipmunks are able to gain matings with females because of qualities other than dominance (such as the ability to successfully find and/or chase receptive females), then the costs of aggression to large chipmunks may outweigh any potential benefits. Small males, but not large males, may improve their mating success by being aggressive.     

Description and behavioural biology of two Ufens species (Hymenoptera: Trichogrammatidae), egg parasitoids of Homalodisca species (Hemiptera: Cicadellidae) in southern California

The morphology, distribution, and hosts of two egg parasitoids, Ufens principalis Owen sp. n. and U. ceratus Owen sp. n. (Hymenoptera: Trichogrammatidae), are described for the first time. These species are compared to U. niger (Ashmead), the only species of Ufens s. str. currently described from the Nearctic, and diagnostic differences are presented. The behavioural biology of U. principalis, and U. ceratus to a smaller extent, is also described for the first time. Ufens principalis exhibited a rapid and long-distance response in the form of directional flight toward freshly laid eggs of Homalodisca species, its primary hosts in southern California. Parasitism involved aggregations of female U. principalis on fresh Homalodisca egg masses, which remained attractive to U. principalis for a relatively short time. The level of oviposition by U. principalis females was low during most of the day and peaked before sunset in tandem with a peak in Homalodisca oviposition. Oviposition behaviour of U. principalis is described and the distribution of ovipositor probe durations showed that most probes were generally of very short duration. Mating of both Ufens species occurred on the egg mass, with males showing aggressive behaviour towards each other as they competed for emerging females. Ufens ceratus males displayed greater aggression towards other males than U. principalis males. By contrast, fights among U. principalis males involved more individuals and lasted longer than corresponding fights between U. ceratus males.