Breeding Season Aggression of Female and Male Eastern Bluebirds (Sialia sialis) to Models of Potential Conspecific and Interspecific Egg Dumpers

Naturally occurring aggression between female eastern bluebirds (Sialia sialis) is dramatic, resulting in severe injuries and even death. Furthermore, aggression among bluebirds is usually sex specific: males attack males, females attack females. We hypothesized that the primary function of female-female aggression is to guard against the threat of intraspecific egg dumping and that, in this context, same sex aggression is related to the possibility of advantages for males of parasitism (egg dumping) of their nests. Our hypotheses to explain variation in naturally occurring aggression predict temporal variation in aggressive tendency within nest cycles and between the sexes depending on asymmetries in threats to the residents. We report the results of experimental trials in the field designed to determine temporal variation in the aggressive tendencies of resident females to models of intruder females of two species, eastern bluebirds and brown-headed cowbirds (Molothrus ater). Both species dump eggs in the nests of bluebirds. Female aggression to eastern bluebird models is greatest during early stages of nest cycles; the patterns are most consistent with protection against egg dumping and protection of nest sites from usurpation. Male residents seldom attack female eastern bluebird models, but often attack models of female brown-headed cowbirds, a result inconsistent with the hypothesis that patterns of differential parental care control aggression of female and male residents.     

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.     

Behavioral and endocrine correlates of multiple brooding in the semicolonial house sparrow Passer domesticus. II. Females

Behavioral and endocrine changes associated with reproductive events were studied in free-living female house sparrows (Passer domesticus). Circulating levels of luteinizing hormone (LH) and estradiol (E2) were maximal during egg-laying, declined during incubation, and began to rise during the nestling stage. This pattern was repeated three to five times during the prolonged breeding season of this species. Multivariate statistical analysis demonstrated that elevated levels of LH and E2 were associated with maximal sexual activity, high rates of intrusion at nests by conspecifics, and elevated rates of agonistic activity. Patterns of sexual activity varied during the season and perhaps reflected differing strategies for protecting paternity adopted by the male. Curiously, elevated levels of testosterone (T) and dihydrotestosterone (DHT) were not associated with high levels of intrusion or agonistic response, but rather were associated with incubation stages. As in males, concentrations of corticosterone (B) were high during each egg-laying and nestling stage and were associated with high or rising levels of reproductive hormones, suggesting that reproductive activity was not overly stressful to these birds. Body mass and fat depots declined during the season and may regulate termination of reproduction. It appears that access to abundant food resources found in association with human dwellings allows house sparrows to maintain reproductive activity longer than most other temperate-zone dwelling bird species.     

Sex of opponent influences response to a potential status signal in house sparrows

We evaluated the possible use of a sexually dimorphic plumage trait as a status signal in inter- and intrasexual social interactions in free-living house sparrows, Passer domesticus. Males with larger black throat patches (bibs) tended to be dominant regardless of the opponent's sex. This relationship was maintained after controlling for body size and age in multivariate analyses. However, the nature of social interactions varied according to the sex of the opponent and a male's bib size. Males with larger bibs tended to be less aggressive to male opponents, and received significantly less aggression from those birds, a key predicted benefit of a status signal. However, when the opponent was a female, males with large bibs displayed significantly more aggression than males with small bibs, and this relationship persisted even after controlling for the dominance status of the participants. Females were significantly more aggressive towards male opponents than towards female opponents, and females interacting with two males of known bib size were more aggressive to the one with the larger bib. These results suggest that the bib of male house sparrows may be a signal of status, but that in interactions with females, the bib may also have other functions. Frequent testing by females of the underlying quality being signalled by the bib may be linked to later mate choice, and might be adaptive if the information value of the bib varies among years. Such testing may also contribute to mechanisms for maintaining the bib as a reliable signal of status. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Effects of testosterone on song, aggression, and nestling feeding behavior in male great tits, Parus major

As outlined in the trade-off hypothesis of testosterone (T) secretion, fluctuations in T during the breeding season might reflect how males allocate their time and energy to competitive behaviors for mates and territories, associated with high T levels, and parental activities, associated with low T levels. In the present study, great tit, Parus major, males were implanted with T-filled or empty silastic capsules at the start of the breeding season and the behavior of these two male categories was compared during the entire breeding season. As a measure of competitive behavior we looked at song behavior and territorial responsiveness to a male decoy, during the three main stages of the breeding period (the egg-laying, incubation, and nestling stages). As a measure of parental care we looked at feeding behavior during the nestling stage. Our results only partly supported the trade-off hypothesis. T implants increased plasma androgen levels and enhanced spontaneous song activity and the production of aggressive vocalizations in response to a decoy. However, our results suggest that the degree of physical aggression might be less than fully coupled with T. First, approach to the decoy was not affected by the treatment. Second, although T levels are known to vary from high during egg laying to low while feeding young, control and T-treated males spent similar amounts of time close to the decoy in the three breeding stages. Our results thus suggest that vocal and physical aggression might be regulated differently in the great tit. Furthermore, in contrast with most other studies on temperate bird species but in agreement with a previous study on the great tit, T treatment did not affect male feeding rates. As the dose of T we used was lower than that typically used in other studies, we cannot at present completely exclude the possibility that the latter result reflects this lower dose of T rather than the species used.     

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.     

Non-migratory stonechats show seasonal changes in the hormonal regulation of non-seasonal territorial aggression

In many birds and mammals, male territorial aggression is modulated by elevated circulating concentrations of the steroid hormone testosterone (T) during the breeding season. However, many species are territorial also during the non-breeding season, when plasma T levels are basal. The endocrine control of non-breeding territorial aggression differs considerably between species, and previous studies on wintering birds suggest differences between migratory and resident species. We investigated the endocrine modulation of territorial aggression during the breeding and non-breeding season in a resident population of European stonechats (Saxicola torquata rubicola). We recorded the aggressive response to a simulated territorial intrusion in spring and winter. Then, we compared the territorial aggression between seasons and in an experiment in which we blocked the androgenic and estrogenic action of T. We found no difference in the aggressive response between the breeding and the non-breeding season. However, similarly to what is found in migratory stonechats, the hormonal treatment decreased aggressive behaviors in resident males in the breeding season, whereas no effects were recorded in the non-breeding season. When we compared the aggressive responses of untreated birds with those obtained from migratory populations in a previous study, we found that territorial aggression of resident males was lower than that of migratory males during the breeding season. Our results show that in a resident population of stonechats T and/or its metabolites control territorial aggression in the breeding but not in the non-breeding season. In addition, our study supports the hypothesis that migratory status does modulate the intensity of aggressive behavior.     

Repeatable nest defense behavior in a wild population of Eastern bluebirds (Sialia sialis) as evidence of personality

Consistent individual differences in behavior, or personality, have been demonstrated in a variety of species other than humans, including mammals, birds, and invertebrates. Behavioral consistency has been shown to affect dispersal, foraging, exploration, and antipredator responses, which may have an impact on parental and offspring survival. Despite increasing research in behavioral consistency, the repeatability of nest defense behavior has rarely been assessed in wild bird populations. Furthermore, previous studies investigating nest defense behavior have utilized laboratory studies or mounted predators to elicit defensive behavior. It is important to assess personality in wild populations to fully understand the fitness consequences of behavioral consistency across natural contexts and to utilize live predators or competitors for accurate assessment of defensive behavior. We used an ecologically relevant, live, invasive, nest site competitor, the house sparrow (Passer domesticus), to elicit nest defense behavior in a wild population of Eastern bluebirds (Sialia sialis) to determine if nest defense behavior is repeatable within and between years for males and females (males: 2009, N?=?17; 2010, N?=?18; both years, N?=?9. Females: 2009, N?=?22; 2010, N?=?15; both years, N?=?11). We also determined if individual behavior changes as a function of season, parental age, brood size, or the number of house sparrows around the nest site. We found that females demonstrated repeatable behavior both within and between years. Male nest defense behavior was only repeatable in 1 year and was influenced by season in the other year. Parental age, brood size, and the number of house sparrows around the nest site did not affect nest defense behavior. We conclude that Eastern bluebirds demonstrate consistent nest defense behavior, or personality, although males are more plastic than females.     

Male Demography,Female Mating Behavior,and Infanticide in Wild Patas Monkeys (Erythrocebus patas)

Infanticide by males has been hypothesized to be a naturally selected behavioral strategy that increases the infanticidal male's reproductive success. The sexual selection hypothesis has been challenged via alternative, nonadaptive hypotheses that dispute its empirical and theoretical bases. Two of the most widely recognized alternatives are the social pathology hypothesis, in which infanticide results from overcrowding or recent human disturbance, and the generalized aggression hypothesis, in which infanticide is an epiphenomenon of increased male aggression. We report the first case of infanticide in wild, seasonally breeding patas monkeys (Erythrocebus patas) living at a low population density in a stable habitat, conditions which do not support the social pathology hypothesis. Its exceptional occurrence is consistent with the sexual selection hypothesis: over a 7-year period the infanticidal male was the only one of 13 resident males that was not present during the actual conception season but was present during the following birth season. Also consistent with this hypothesis, mothers were differentially targeted for male aggression, which increased sevenfold during the days surrounding the infanticide and then decreased to baseline levels after the infanticide. Aggression targeted at mothers does not support the generalized aggression hypothesis. As predicted by the sexual selection hypothesis, females began soliciting mating immediately after the infanticide, despite its occurrence in the nonconceptive season.     

Territoriality and testosterone in an equatorial population of rufous-collared sparrows, Zonotrichia capensis

Territorial aggression, displayed by male vertebrates in a reproductive context, is generally thought to be mediated by testosterone. The challenge hypothesis predicts that in socially monogamous species, territorial challenges should induce an increase in plasma testosterone concentrations, which will enhance aggressive behaviour and territory defence. This hypothesis is based on northern latitude birds and needs to be tested in tropical birds before it can be universally accepted. We tested the challenge hypothesis in an equatorial population of rufous-collared sparrows in Papallacta, Ecuador. This population shows an extended breeding period during which males aggressively guard territories. During the early breeding season, males were challenged with conspecific or heterospecific simulated territorial intrusions (STIs) lasting 10 min. Conspecific-challenged males responded more aggressively than heterospecific-challenged males. However, there was no increase in plasma testosterone in response to the conspecific STI. During the breeding season, males were challenged with conspecific STIs lasting 0, 10 or 30 min. Males behaved aggressively regardless of STI duration, and did not differ in plasma testosterone or luteinizing hormone concentrations. During the breeding season, males were implanted with testosterone-filled or empty silastic tubes and subsequently challenged with a conspecific STI. Testosterone implants significantly raised plasma testosterone concentrations, but testosterone-implanted males were not more aggressive than blank-implanted controls. Combined, these findings suggest that testosterone concentrations above breeding baseline are not related to territorial aggression in this population and therefore do not support the challenge hypothesis.     

Reproductive Consequences of Aggression in a Territorial Songbird

Territorial aggression can influence males’ ability to obtain high‐quality resources and access to mates; however, in many species, the reproductive consequences of variation in aggression are unknown. In this study, we investigated how individual variation in aggressive behavior relates to reproductive success in socially monogamous, genetically polygynous song sparrows (Melospiza melodia). Prior research in this species shows that male song sparrows differ in their willingness to engage in agonistic interactions with territorial intruders and that individual variation in aggression appears to have functional significance. Aggressive males have been shown to obtain territories where females produce larger clutch sizes, suggesting that individuals who display high levels of territorial aggression are defending high‐quality territories or females. Further, aggressive males are considered a greater threat to territory‐holding males than less aggressive males. In this study, we ask whether individual differences in aggression are linked to differences in extra‐pair reproductive success, annual reproductive success, and offspring quality. We did not uncover a relationship between aggression and annual reproductive success or patterns of extra‐pair paternity. However, we found that the nestlings of aggressive males grew at a faster rate than the nestlings of less aggressive males. Future studies should attempt to identify mechanisms to explain the relationship between offspring growth rate and male aggression and investigate whether faster offspring growth rates translate to greater survival and recruitment of offspring.     

The role of androgen receptors in regulating territorial aggression in male song sparrows

This paper examines the role that androgen receptors (ARs) play in modulating aggressive behavior in male song sparrows, Melospiza melodia morphna. Song sparrows are seasonally breeding, territorial birds that maintain year-round territories with male-female pair bonds formed during the spring breeding season. Plasma testosterone levels peak as territories are established and mates acquired. In late summer, testosterone levels fall and remain basal during the non-breeding season. We examined the role of ARs in regulating territorial aggression in captive song sparrows under short- and long-day conditions as well as just prior to, and at the start of the breading season in freely living birds using the nonsteroidal antiandrogen flutamide to block AR function. Birds were implanted with either empty or drug filled silastic implants for 18 to 42 days and then challenged with a novel male decoy to assess the individual birds level of male-male aggression. Freely living birds remained on their home territory and underwent a simulated territorial intrusion, whereas laboratory-held birds were assessed using a laboratory simulated territorial intrusion and remained in their home cage. Experimental treatment of male song sparrows decreased aggressive behavior during the pre-breeding life history substage (March-April) in freely living birds as well as in laboratory-held birds under long-day (16L:8D) conditions. During the early breeding substage (April-May) there was no measurable effect of flutamide treatment on aggressive behavior, nor was there a difference in behavior in the (8L:16D) laboratory birds. This demonstrates that ARs are an important component of the neuroendocrine control of aggressive behavior. Given that flutamide only affected aggression during the pre-breeding substage and in LD birds, the results suggest that AR dependent control of aggressive behavior changes as song sparrow life history states change.     

Predation by sparrowhawks Accipiter nisus on male and female pied flycatchers Ficedula hypoleuca in relation to their breeding behaviour and foraging

Bright plumage, song display, and aggressive resource defence in males may cause higher predation on males than on females during the breeding season. However, in birds, higher predation on females is sometimes observed. Parental investment may be high in females (egg-laying, incubation and feeding of offspring), which might lead to a high risk of predation. We studied predation by sparrowhawks Accipiter nisus in relation to behaviour in pied flycatchers Ficedula hypoleuca where breeding males are more conspicuous than females in plumage and behaviour. Male pied flycatchers generally occupied more exposed perches than females. Females were more mobile and foraged more than males, especially prior to and during incubation. During the incubation and nestling stages, when predation on the sexes could be directly compared, sparrowhawks took about the same number of male and female pied flycatchers. During incubation, however, females spent about 77% of the day in the nest and were 4.7 times more vulnerable than males per unit of time available (i.e. outside the nest). A comparison with the chaffinch Fringilla coelebs , where hawks took more females than males, indicates that timing of breeding, foraging behaviour and parental roles of males and females affect predation risk.     

Consequences of breeding system for body condition and survival throughout the annual cycle of tidal marsh sparrows

An individual's body condition and probability of survival can change throughout the annual cycle, based on the combined effects of many factors, including reproductive investment during breeding, colder temperatures during winter, and elevated risks during migration. We evaluated body condition and survival during breeding and non‐breeding periods in two closely related species with notably different reproductive systems. Male and female saltmarsh sparrows Ammodramus caudacutus represent extremes in parental care: males perform none, leaving females to do everything from build nests to care for fledglings. In contrast, male and female seaside sparrows A. maritimus have bi‐parental care and similar levels of reproductive investment, intermediate between male and female saltmarsh sparrows. Our results are consistent with the idea that females experience non‐lethal effects of reproduction, and that differences between the breeding season and winter affect condition. In both species, females had lower scaled mass index (SMI) values than males during both breeding and non‐breeding seasons, and female saltmarsh sparrows had lower SMI values than female seaside sparrows. Females carried more fat than males during the breeding season, and female, but not male, fat and muscle scores decreased over time, which is consistent with the adaptive mass hypothesis. In winter, all groups carried more fat and had higher muscle scores than when breeding, despite having lower SMI scores. Although we observed variation in body condition, within‐season survival was uniformly high in both seasons, suggesting that sex, species, season, body size, and body condition have little impacts on within season survival. Comparisons with previously‐published estimates of annual adult survival suggest that most mortality occurs during migration, even in these short‐distance migrants. The importance of considering multiple aspects of body condition, multiple seasons, and difficult‐to‐monitor events, such as migration, should not be ignored when thinking about the events and processes that cumulatively determine population dynamics.     

How do floater male eastern bluebirds benefit by filling vacancies on territories where females already have young?

When male eastern bluebirds (Sialia sialis) are killed or deserttheir mates, replacement males usually take over the territory.We captured 25 eastern bluebird males and held them in captivityto study the behavior of replacement males and their responseto young already present in the nest. Of 17 replacement males,most courted the females, and 23% of experimental females renested,a proportion not different from that of control females. Noreplacement males were infanticidal, and none fully adoptedthe young. Replacement males spent significantly less time onthe territory and defended it significantly less than controlmales. We discuss breeding options available to a floater maleeastern bluebird and conclude that (1) floater male bluebirdsthat fill territory vacancies benefit by gaining breeding opportunitiesin the current breeding season, and (2) indifference to theyoung is the least costly option for replacement male easternbluebirds.     

Honesty of a dynamic female aggressive status signal: baseline testosterone relates to bill color in female American goldfinches

Status signals are linked to fighting ability and enable competitors to gain access to resources without risking injury in aggressive combat. The relationship between testosterone (T), a hormone that mediates aggression, and signals of status is well studied in males, but little is known about the relationship between T and female signals of status. Female and male American goldfinches Spinus tristis express a dynamic carotenoid‐based orange bill color during the breeding season and previous work has demonstrated that females use orange bill color to communicate competitive ability during intrasexual competition. We test the hypothesis that female bill color reflects baseline T, which would allow receivers to directly assess a competitor's aggressive potential. We found a positive relationship between T and bill coloration in females, indicating that bill color has the ability to signal female competitive status. This finding is consistent with the hypothesis that female bill color is a reliable signal of fighting ability, and indicates that females, like males, may use coloration to signal their hormonally mediated aggressive potential.     

Effects of bill pigmentation and UV reflectance during territory establishment in blackbirds

During intrasexual interactions, the visual properties of conspicuous male sexually selected traits may be important in social signalling. We used intruder models to examine the effect of pigmentation and ultraviolet (UV) reflectance of a single male trait on the responses of male blackbirds, Turdus merula. Stuffed models of male blackbirds with brown, yellow or orange bills were presented to residents in their natural habitat; we altered UV reflectance from the bills by applying nail varnish. This UV reflectance did not appear to affect the response of resident males during simulated territorial intrusions under natural signal and viewing conditions. However, models with brown bills received less attention from resident males than models displaying yellow or orange pigmentation. Brown bills are typical of first-year males and may be an effective signal of subordinate status, reducing aggression from adult males. Furthermore, resident males came closer, and initiated aggressive displays more quickly, in the presence of orange- versus yellow-billed models, suggesting that orange-billed models may be perceived as more of a threat to territory ownership. In 44% of presentations, females also responded to models (a response rate half that of males). Females spent more time more than 10 m from models when they lacked UV, suggesting that UV may be more important during intersexual interactions. However, bill pigmentation did not affect female responses to the models. Males and females may be attentive to different properties of bill appearance when establishing the territory. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Disruption of the fifth melanocortin receptor alters the urinary excretion of aggression-modifying pheromones in male house mice.

The preputial glands of house mice express the gene for the fifth melanocortin receptor (MC5-R) and are a primary source of urinary pheromones involved in inter-male aggression. A 'resident-intruder' behavioral model was used to compare the responses of resident males to urine from mice with an engineered disruption of the fifth melanocortin receptor (MC5-RKO) with residents' responses to urine from wild-type mice (WT). Each type of urine was presented in combination with a castrated intruder male to provide the appropriate biological context. Resident males responded with a longer latency to bite when the urine was from gonadally intact WT males compared with urine from MC5-RKO mice. These results are consistent with the hypothesis that activation of the fifth melanocortin receptor in the preputial glands of male house mice causes excretion of urinary pheromones that delay aggressive responses by other males.     

Pair bonding and "the widow effect" in female prairie voles

We conducted field and laboratory experiments with the well-studied monogamous prairie vole, Microtus ochrogaster, to distinguish among three hypotheses for the failure of females that lose their mates to bond with a new male ("the widow effect"). The reproductive value hypothesis predicts that males prefer young to older females because they potentially have a longer reproductive lifespan. The mate rejection hypothesis predicts that females will prevent repairing by aggressively deterring males that might harm their current offspring. The misdirected paternal care hypothesis assumes that females will mate during postpartum estrus and thus will be pregnant and/or nursing young throughout the breeding season; males will avoid pairing with older females to avoid providing care to unrelated offspring and/or because of a delay to the next breeding opportunity. Males associated and bred more with older than young females, allowing us to reject the reproductive value hypothesis. Our results were consistent with the male rejection hypothesis in that females were aggressive toward unfamiliar males. Our results were most consistent with the misdirected paternal care hypothesis in that once females started breeding, they continued to become pregnant and nurse young throughout the study period. Thus, our findings suggest that the potential of misdirected paternal care and delayed mating opportunity in conjunction with the aggressive behavior of females toward unfamiliar males are likely explanations for the lack of repairing for widow females.     

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.