Singing and the establishment and maintenance of dominance hierarchies in captive brown-headed cowbirds

Summary The importance of singing in the establishment and maintenance of dominance rank was investigated in captive flocks of brown-headed cowbirds (Molothrus ater). Each flock consisted of 3 males and 1 female. In one experiment the two largest males in each of 4 flocks were temporarily devocalized by puncturing the interclavicular air sac just before group formation. The third male in each group was sham-operated. There was a significant tendency for the non-devocalized birds to become the dominant member of their group. Only sham-operated males sang immediately after the operation. In a second experiment, dominant males from flocks with established hierarchies were temporarily devocalized. None of these males lost their dominant positions, despite their inability to sing. These results suggest that song is important in the establishment of dominance rank; however, once established, social hierarchies may be maintained by non-vocal means, such as social inertia or individual recognition.     

Red-winged blackbird parental investment following brood parasitism by brown-headed cowbirds: is parentage important?

Parental investment by red-winged blackbirds (Agelaius phoeniceus) in response to natural and experi‐mental parasitism by brown-headed cowbirds (Molothrus ater), and in response to freeze-dried, female cowbird mounts presented near redwing nests during the egg-laying period was measured. Two measures of redwing parental investment were used: nest defense effort toward a model predator, and rate of feeding nestlings. There were no significant differences in levels of parental investment among unparasitized nests, naturally parasitized nests, or experimentally parasitized nests. Similarly, parental investment did not differ between redwings that were exposed to the cowbird mount and those that were not exposed to the mount, or among redwings exposed to the cowbird mount at different distances from the nest. This suggests that red-winged blackbirds do not recognize when they have been parasitized, and hence do not associate parasitism with a decrease in their parentage, or that parentage is not an important predictor of parental investment in this species. Received: 24 January 1997 / Accepted after revision: 7 June 1997     

Functional reference in an alarm signal given during nest defence: seet calls of yellow warblers denote brood-parasitic brown-headed cowbirds

Field observations and model-presentation experiments have shown that yellow warblers (Dendroica petechia) produce seet calls preferentially in response to brood-parasitic brown-headed cowbirds (Molothrus ater). In this study, we investigated whether seet calls are functionally referential alarm calls denoting cowbirds by determining whether female warblers responded appropriately to seet calls in the absence of a cowbird, whether alarm calling by warblers varied with response urgency, and how warblers in a population allopatric with cowbirds responded to cowbird and avian predator models and seet playbacks. As a control, we presented chip calls, which are elicited by nest predators as well as by non-threatening intruders, but are not strongly associated with cowbirds. Yellow warblers responded differently to playbacks of seet than chip calls. To seet playbacks, almost 60% of females gave seet calls and rushed to sit in their nests, responses typically elicited by cowbirds, whereas these responses were given infrequently in response to chip calls. Yellow warblers seet called equally in situations that simulated low, medium and high risk of parasitism, which suggests that these calls did not vary with response urgency. In a population allopatric with cowbirds, seet calls were rarely produced in response to cowbird or avian nest predator models and never to seet playbacks. These results suggest that seet calls are functionally referential signals denoting cowbirds and that cowbird parasitism was a strong selective pressure in the evolution of functional referentiality in the seet call of yellow warblers.Communicated by W.A. Searcy     

Male contests in the scarlet rosefinch (Carpodacus erythrinus) in relation to asymmetries in resource holding power and pairing status

Summary Inter-male contests in relation to asymmetries in their pairing status and resource holding power were studied in a population of the Scarlet rosefinch. Since males in this species do not defend territories before nest building, and since most males have full adult plumage, possession of territory and age are unlikely to be used as cues in the settlement of contests. Rather, two other features are used, namely size and pairing status. In this study, contests between two unpaired males were usually short displacements, whereas 56% of contests between two paired males were escalated. In the first case, the smaller male withdrew in all contests, whereas in the second case there was an equal probability of withdrawal with regard to size. There was no relationship between size difference and the probability of escalation in the contests. Most of the fights between paired and unpaired males escalated, namely in 60% of the cases where the unpaired male was smaller and in 75% of the cases where he was larger. The unpaired male withdrew in 18 out of 23 contests, suggesting that pairing status is more important than size in determining the outcome of a contest. There was no variance in size differences between contestants in short and escalated contests. This study demonstrates that there is a high competition between males for females in this population.     

The appeasement effect of sterility signaling in dominance contests among Bombus terrestris workers

The establishment of dominance hierarchies through aggressive interactions is very common in insect societies. In many cases, it is also mediated through pheromone emissions that enable individuals to evaluate the reproductive quality and level of aggressiveness of the dominant individual, thereby reducing the number and intensity of costly fights. Here, we studied these processes in the primitively eusocial bee Bombus terrestris, using a paired bee system. Specifically, we investigated the behavioral, reproductive, and pheromonal correlates of dominance establishment. Workers were shown to establish dominance hierarchies using overt aggression within 3–4 days. Thereafter, the aggression drastically decreased, and dominance was maintained mostly by ritualized agonistic behavior. The behaviorally dominant bee lost the ester compounds that workers produce in their Dufour's gland (the so-called “sterility signal”) concomitantly with the development of her ovaries. The other bee announced as subordinate by continuously producing high amounts of those esters. The hypothesis that sterility signaling serves as an appeasement signal to pacify the dominant bees is supported by the negative correlation found between the proportion of these esters and the level of aggression that the subordinate received from the dominant worker. Physical interactions, and presumably also the ensuing overt aggression between the bees, were essential for the above pheromonal change to take place and enabled the dominant workers to develop their ovaries and to lay eggs. The subordinate bee’s signaling of non-reproductive status may minimize energy expenditure in costly fights and help stabilize the reproductive division of labor among workers.     

Food sharing among retaliators: sequential arrivals and information asymmetries

Many animals share food, that is, to tolerate competitors at a defensible clump. Most accounts of resource sharing invoke special evolutionary processes or ecological circumstances that reduce their generality. Surprisingly, the Hawk–Dove game has been unable to address in a simple and general way why so many group foraging animals share food. We modify the Hawk–Dove game by allowing a finder the opportunity of retaliating if joiners escalate and by considering the consequences of information asymmetries concerning resource value among players. Introducing the first change, the retaliator strategy was sufficient to predict widespread sharing in habitats where food clumps are of intermediate richness. When information asymmetry between finder and joiner is created by allowing the quality of clumps to vary, we show that the conditions for sharing are even more easily met and apply to a wider range of resource qualities. Our model therefore offers one of the most parsimonious and potentially general evolutionary accounts of the origin of non-aggressive resource sharing.     

Interference asymmetries among age-sex classes of rufous hummingbirds during migratory stopovers

Summary Three age-sex classes of rufous hummingbirds (Selasphorus rufus) overlap temporally and defend feeding territories during migratory stopovers in the Sierra Nevada of California. We demonstrate that these classes differ in their ability to secure and maintain high-quality feeding territories for refueling, and that these differences result in differences in resource use. Data on acquisition of territories, territory characteristics, and responses of territory owners to intruders suggest that several mechanisms are involved in determining dominance, involving sex- and age-related differences in wing disc loading, coloration, and experience. We discuss the implications of these results for understanding intraspecific variation in migration strategies. Correspondence to: F.L. Carpenter     

Territoriality among male red-winged blackbirds

Summary We examined male site fidelity and territorial movements in a population of individuallyidentifiable red-winged blackbirds (Agelaius phoeniceus) over an 8-year period. Based on general theory and knowledge of the ecology of this population, we predicted that males would be site conservative, and that they would voluntarily change territories only when they could expect to improve significantly their reproductive success. Because males are absent from their territories for only short periods, and probably have accurate comparative information only about nearby territories on which to base their decisions, we predicted that most moves would be over short distances. About 70% of males that bred for more than 1 year retained their original territories between breeding years. Most of the 30% of males that changed territories moved less than 200 m, often to adjacent territories. As predicted, males moving less than 200 m tended to have better reproductive success after moving than before, whereas long distance movers did not improve their success after moving. Territorial male redwings appear to monitor breeding activity on nearby territories and move when significant potential benefits are perceived and opportunities exist. Long-distance moves may be involuntary ones. The amount of information possessed about present and alternative sites, and the time and opportunity required to collect the information, are probably major constraints influencing site conservatism in this and other territorial species.     

Territoriality among male red-winged blackbirds

Summary We tested two hypotheses to explain territorial dominance in male birds. Male red-winged blackbirds were removed from their territories for 7 d and then released after replacement owners had held their territories 2 to 7 d. Original owners regained territories from short-term replacements, but could not defeat 6 to 7 d replacements. This outcome suggests that replacement males relinquished their territories to persistent original owners after 2 to 3 d of ownership because the territory lacked sufficient value to them, but not after 7 d, when its value was greater. This result supports the Value Asymmetry Hypothesis of territorial dominance and provides strong evidence in birds that differences in the extent of knowledge of or investment in an area and, hence, willingness to escalate contests, contribute to territorial dominance.     

Territoriality among male red-winged blackbirds

Summary To test theories recently proposed to explain territorial dominance in animals, we performed several versions of experiments in which male red-winged blackbirds were removed from their territories, held in captivity for varying periods, and then released to challenge their replacements. Males removed for 7 to 49 h recovered their territories from replacement males, either when released or over the following few days or weeks. The duration males were held off territory, the duration replacement males occupied territories, and the original owners' awareness before fighting that they had been replaced, apparently did not influence contest outcomes, but whether the new owner was a neighbor or a previously non-territorial male had some effect. The pattern of territory recovery observed most closely supports the hypothesis that territorial dominance in redwings arises from asymmetries in local knowledge and experience between owners and challengers, although another hypothesis, the Resource Holding Potential hypothesis, was not entirely ruled out. We discuss design of removal experiments to test territoral dominance, and propose that ecologies of particular species may powerfully influence outcomes of these experiments.     

Resource-holding power asymmetries,the prior residence effect,and reproductive payoffs in male northern elephant seal fights

The effect of resource-holding power (RHP) and prior residency asymmetries on fight outcome and subsequent seasonal copulatory success was analyzed for fights between marked male northern elephant seals (Mirounga angustirostris). RHP asymmetries were measured as differences in estimated mass and prior residency asymmetries were measured as differences in beach tenure prior to the fight. The principal results were: (a) Neither differences in mass nor differences in beach tenure had any effect on fight outcome as separate factors. (b) Mass and tenure differences had an interactive effect on fight outcome; fight winners were either heavier males present for shorter periods (intruders) or lighter males present for longer periods (prior residents). (c) Winners of fights copulated more often than losers after a fight throughout the breeding season; this difference was smallest for low-ranking males, larger for high-ranking males in short fights, and greatest for high-ranking males in long fights. (d) Prior resident males who won long fights obtained significantly more copulations after a fight than the males they defeated, but this was not true for intruder males who won long fights. These results suggest that male northern elephant seals will incur greater contest costs (i.e., fight for longer periods and/or against heavier males) for higher reproductive payoffs. They also imply that, at least for males in long fights, differences in prior residence represent payoff asymmetries, with higher reproductive payoffs for winning prior residents than for winning intruders.     

Association patterns among male and female spotted hyenas (Crocuta crocuta) reflect male mate choice

Although female animals tend to be choosier than males in selecting mates, sexual selection theory predicts that males should also be choosy when female fecundity varies. Reproductive success among female spotted hyenas varies greatly with social rank. Our goals were therefore to determine whether male hyenas preferentially associate with high-ranking females, and whether male preferences are affected by female reproductive state. Interactions between adult males and females were observed intensively, and association indices calculated for all male-female pairs, over a 7-year period in one population of free-living hyenas. Males initiated most affiliative interactions with females, and males associated most closely with females that were likeliest to be fertile. High- and middle-ranking males associated most closely with high-ranking females, but low-ranking males associated equally closely with females in all rank categories. We used molecular markers to determine the paternity of cubs born during the study period, and found that sires associated more closely with the mothers of those cubs than did non-sires, particularly during the last months before conception. These association data indicate that male spotted hyenas do indeed exhibit selective mate choice, and that they prefer females likeliest to maximize male reproductive success.     

Shiny cowbirds share foster mothers but not true mothers in multiply parasitized mockingbird nests

Obligate brood parasitic birds, such as cowbirds, evade parental care duties by laying their eggs in the nests of other species. Cowbirds are assumed to avoid laying repeatedly in the same nest so as to prevent intrabrood competition between their offspring. However, because searching for host nests requires time and energy, laying more than one egg per nest might be favoured where hosts are large and can readily rear multiple parasites per brood. Such ‘repeat parasitism’ by females would have important consequences for parasite evolution because young parasites would then incur indirect fitness costs from behaving selfishly. We investigated shiny cowbird (Molothrus bonariensis) parasitism of a large host, the chalk-browed mockingbird (Mimus saturninus), in a population where over 70 % of the parasitized mockingbird nests receive multiple cowbird eggs. We assessed egg maternity directly, using cameras at nests to film the laying of individually-marked females. We also supplemented video data with evidence from egg morphology, after confirming that each female lays eggs of a consistent appearance. From 133 eggs laid, we found that less than 5 % were followed by the same female visiting the nest to lay again or to puncture eggs. Multiple eggs in mockingbird nests were instead the result of different females, with up to eight individuals parasitizing a single brood. Thus, while cowbird chicks regularly share mockingbird nests with conspecifics, these are unlikely to be their maternal siblings. Our results are consistent with shiny cowbird females following a one-egg-per-nest rule, even where hosts can rear multiple parasitic young.     

Impact of contaminant exposure on resource contests in an invasive fish

There is increasing concern for the disruptive effects seen in aquatic species exposed to environmental contaminants. However, few studies have investigated the impact of such contaminants on the behavior of individuals living in exposed waters. Contaminant exposure can affect animal populations by disrupting behaviors including feeding, locomotion, and mating. In this study, we examined how living in an ecosystem polluted by combinations of polycyclic aromatic hydrocarbons, polychlorinated biphenyls, and heavy metals (arsenic, cadmium, iron, lead, zinc) impacts contest behavior in the round goby (Neogobius melanostomus). Fish collected from heavily contaminated and cleaner sites in Lake Ontario were subjected to a resource contest to determine the effect of these contaminants on aggression and the establishment of dominance hierarchies, which in turn influence access to food, shelter, and mating opportunities. Dominance establishment (a clear resource winner) was less obvious among fish from the contaminated site compared to the more stable hierarchies that formed between pairs of fish from the clean site. Pairs of fish from the contaminated site performed more assessment displays compared to fish from clean sites. These results suggest that the costs of living in an environment under exposure can shape behavioral repertoires. The altered conflict resolution strategies of contaminated fish may reflect impaired cognitive function, sensory perception, and/or higher metabolic load associated with aggression. This study provides support for the utilization of quantifiable behavioral differences as ecologically relevant measures of contaminant exposure.     

Apparent resource competition among embryos in the brood pouch of a male pipefish

Embryo success was studied in the paternally brooding pipefish Syngnathus typhle. During brooding, which lasts about a month, males provide embryos in their brood pouch with nutrients and oxygen via a placenta-like structure. Egg size depends on female size. In aquaria, males were mated with differently sized females to give the following treatments: M, mixed-egg-size broods of approximately half large and half small eggs; L, single-egg-size broods of large eggs; S, single-egg-size broods of small eggs; and F, field mated males. All males were kept in aquaria for a full brooding period. For each egg-size category, the number of newborn was compared with the number of eggs the male initially fertilized in his brood pouch. Within mixed-egg-size broods, a higher proportion of large eggs survived and large eggs resulted in heavier newborn than small eggs. Indeed, small eggs from a mixed-egg-size brood had significantly lower relative success (proportion of embryos surviving to birth) than those from a brood entirely composed of small eggs. The implication is that embryos compete for resources within the brood pouch, and that competitive success depends on egg size. Given that females produce eggs corresponding in size to their body size, and that females are known to compete indirectly for access to mates (i.e., the sex-roles are reversed), this intrabrood competition could be seen as an extension of female-female competition, but alternative explanations are discussed. Received: 28 April 1995/Accepted after revision: 28 October 1995     

Female brown-headed cowbirds' (<Emphasis Type="Italic">Molothrus ater</Emphasis>) social assortment changes in response to male song: a potential source of public information

In many species, females' behavior appears to be influenced by that of other females, particularly regarding mate choice. Females theoretically can reduce the costs associated with independent male assessment by observing conspecifics. Studies of brown-headed cowbirds suggest that females pay attention to other females' behavior. Group-housed females modify their song preferences, whereas females housed in pairs do not. What information is available to females in a group environment? To address this question, we studied two groups of juvenile (i.e. hatch-year birds) and adult female cowbirds in a naturalistic group setting. We used a longitudinal ABA design, consecutively introducing and removing males that differed in age, amount of song production and stage of song development, to isolate the male characteristics that related to changes in female behavior. Juvenile and adult females assorted by age class when singing adult males were in the aviary, but not when singing juveniles or silent males of any age class were in the aviary. Results from playback tests confirmed that adult male song alone influenced female age class assortment. Videotape analysis from playback tests revealed that females also wing stroked in response to male song. Other females sometimes approached females who wing stroked and observed them. We hypothesize that group-level changes in social organization and individual females' responses can serve as visual signals for other individuals.     

Testosterone predicts future dominance rank and mating activity among male chacma baboons

Despite the many benefits that testosterone has on male reproduction, sustaining high levels of testosterone for long periods can be costly. The challenge hypothesis predicts that males will show temporarily sustained elevations of testosterone at critical periods, counterbalanced by decreased levels during noncritical periods. We investigated male testosterone measures extracted from fecal samples in a group of chacma baboons (Papio hamadryas ursinus) living in the Okavango Delta, Botswana. Because rank serves as a proxy for competition for mates, we examined how male testosterone was related to dominance rank, age, aggression, and mating activity. Males showed an elevation in testosterone at maturity; young adult males had the highest testosterone levels followed by a steady decline with age. Among dispersing males, testosterone was temporarily elevated in the month following dispersal. After controlling for age, testosterone and rank were unrelated, but testosterone and changes in rank were positively correlated, such that males rising in rank had higher testosterone than males falling in rank. Thus, for males in this group, testosterone was predictive of a male's rank trajectory, or future rank. Similarly, male testosterone levels predicted future, rather than current, mating activity. Finally, male testosterone and aggression rates were unrelated during stable periods in the dominance hierarchy but positively related during unstable periods when high ranks were being contested. In general, our results support the challenge hypothesis with males exhibiting elevated testosterone in association with the acquisition of high rank (ensuring access to mates), rather than with mating itself.