Use of the song repertoire in intersexual and intrasexual contexts by male red-winged blackbirds

Summary Live and stuffed male and female red-winged blackbirds (Agelaius phoeniceus) were presented to territorial males in order to study the repertoire organization of males during intersexual and intrasexual encounters. Territorial male red-winged blackbirds switched song types more frequently and sang more song types in response to presentation of a female stimulus than during control periods, but decreased their switching frequency when a male stimulus was presented. Switching frequency in response to the female stimulus was three times that in response to the male stimulus. These results support the hypothesis that song repertoires of male red-winged blackbirds evolved primarily in response to intersexual rather than intrasexual selection. They also suggest that male red-winged blackbirds have been selected to de-emphasize their song repertoires during encounters with conspecific males as a result of some as yet unidentified cost. Offprint requests to: W.A. Searcy     

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.     

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.     

Ideal dominance distributions: a test using red-winged blackbirds (Agelaius phoeniceus)

Summary The ideal dominance distribution model predicts that competition between individuals of a species for territories will result in socially dominant individuals acquiring territories in higher quality habitat than their subordinates. Although the dispersion and relative reproductive success of male red-winged blackbirds (Agelaius phoeniceus) across habitats in eastern Ontario appears to conform to the ideal dominance distribution model, data from a study of three captive groups comprised of males from both high (marsh) and low (upland field) quality habitats failed to support the prediction that males from marsh habitat are dominant to those from upland habitat. Contrary to the prediction males from uplands were generally dominant to males from marshes. We found a significant positive correlation between dominance and both increased epaulet size and increased body size. Controlling for these positive effects, upland males remained generally dominant to marsh males. Measurements of independent samples of males from both habitats indicated that the overall distribution of males does not conform to ideal dominance. We suggest that the strong between-year territory fidelity shown by male red-winged blackbirds and chance events when they initially acquire territories may contribute to this lack of conformity.     

Polygyny in the red-winged blackbird: do females prefer monogamy or polygamy?

Five hypotheses have been proposed to explain polygyny in the red-winged blackbird (Agelaius phoeniceus). We categorized the hypotheses into three groups based on female preference for unmated versus monogamously mated males: (1) the “polygyny threshold” model, “sexy son” hypothesis and the “asynchronous settlement” model, which assume that females prefer unmated males to mated males on breeding situations of homogeneous quality; (2) the “neutral mate choice” hypothesis, which assumes that females have no preference; and (3) the “cooperative female choice” model, which assumes that females prefer monogamously mated males to unmated males. We tested the direction of female preference in two field experiments. In both experiments, newly settling females were given a choice of two adjacent territories, one defended by an unmated male and the other by a monogamously mated male. Male mating status was randomized with respect to the variation in territory quality and male quality. Early in the breeding season, significantly more females settled with the unmated males than with the mated males. Although more females settled with the unmated males than with the mated males late in the breeding season, the difference was no longer significant. Female settlement late in the season appeared to be related to the tenure of the resident females: the new females avoided territories where the resident females were in early stages of their nesting, but settled on territories where the resident females were in late stages. The pattern of female settlement shows that females prefer unmated males to mated males. The preference is consistent with the polygyny threshold model, sexy son hypothesis and the asynchronous settlement model, and inconsistent with the neutral mate choice hypothesis and the cooperative female choice model. For this reason, the latter two hypotheses are unlikely to explain the occurrence of polygyny in our population of red-winged blackbirds. Received: 1 December 1994 / Accepted after revision: 28 October 1995     

Results of dyeing male yellow-headed blackbirds solid black: implications for the arbitrary identity badge hypothesis

Summary Territorial male yellow-headed blackbirds that were dyed solid black were similar to controls in their ability to attract mates. They also had no difficulty in defending their territories from floating males. The only remarkable effect of this dramatic color manipulation was that five of the 12 blackened males that regained a territory after being dyed also later took over the better quality territory of another male. Not one of the 11 control males that regained a territory succeeded in such an eviction. This surprising result is interpretable under the theory that conspicuous and distinctive coloration sometimes evolves as an arbitrary recognition badge by making good fighters more memorable to their opponents. With insight gained from these results, we modify and considerably expand the theoretical treatment of this Arbitrary Identity Badge Hypothesis. We also explain why our results for yellow-headed blackbrids should differ so dramatically from those for red-winged blackbirds where blackened males often lost their territories to floating males.     

Does the “Teer” vocalization deter prospecting female red-winged blackbirds?

Summary A speaker-occupation experiment was conducted to determine whether female red-winged blackbirds (Agelaius phoeniceus) use the aggressive Teer vocalization to deter conspecific females that are prospecting for opportunities to settle. In each of 14 trials conducted during the breeding seasons of 1987, 1988, and 1989, tape recordings and a pair of loud speakers were used to defend suitable habitat. Observations of female red-winged blackbirds settling on these speaker territories and on adjacent control areas showed that speaker territories were settled significantly earlier than silent control territories. Each of the 14 first settlers on speaker territories included at least one speaker location within the bird's display area, and 9 of the 14 included both speaker locations. Additional observations showed that the rate at which prospecting females visited control and speaker territories, and the number of females eventually settling on them, were not significantly different. Finally, females that settled on speaker territories sang Teers at significantly higher rates than those on control territories. These results demonstrate that the Teer alone was not sufficient to deter prospecting female red-winged blackbirds. On the contrary, Teers given in the absence of an actual resident may have attracted settlers. It is suggested that the aggressive Teer vocalization might function to establish dominance relationships among the females sharing a polygynous male's territory.     

Site fidelity and territorial movements of males in a rapidly declining population of yellow-headed blackbirds

We tested several hypotheses to explain low between-year territory fidelity in a breeding population of yellow-headed blackbirds (Xanthocephalus xanthocephalus). During a 5-year study the population of territorial males declined by two-thirds and some of the marshes that supported territories significantly deteriorated. Individual males held territories and bred for an average of 1.9 years. Of males that bred for at least 2 years, 30% skipped owning a territory in the study area during at least 1 year of their breeding lifetimes. Our information suggests that they may have bred outside of the area in those years. Of males with territories in two or more breeding seasons, 60% changed breeding marshes at least once. Males changed territories during 42.9% of between-year opportunities to do so. We found no support for the hypotheses that male yellow-headed blackbirds: (1) are more likely to move when territory density is low; (2) are likely to abandon territories that are deteriorating; or (3) change territories to improve their reproductive success. We suggest three non-mutually exclusive explanations for the yellow-headed blackbird's weak site fidelity: (1) it is a response to habitat deterioration and to other factors that may be causing the population's decline; (2) the males, being migratory, make fresh settlement decisions each year after they arrive on the breeding grounds in the general vicinity of their previous year's breeding; (3) yellow-headed blackbirds may have evolved in, and be adapted to, highly unstable habitats, moving frequently in response to changes in local breeding site conditions. Correspondence to: L.D. Beletsky     

Breeding synchrony and extra-pair mating in red-winged blackbirds

Using data from a 6-year paternity study of red-winged blackbirds, I tested the hypotheses that increased nesting synchrony should either promote extra-pair mating by increasing the advantage of extra-pair mating to females, or decrease extra-pair mating by constraining males from seeking extra-pair copulations. Contrary to these hypotheses, the occurrence of extra-pair paternity did not vary with nesting synchrony over the breeding season, or vary with the number of synchronous nests within territories or within marshes, or with nesting order on territories. However, for nearly all nests with extra-pair young, there were fewer females synchronous with that nest on the cuckolder's territory than on the territory of the cuckolded male. This “advantage” of a synchrony difference was less pronounced for older males that cuckolded younger males, particularly when the two males were not neighbors. Collectively, these results suggest that breeding synchrony affects extra-pair mating by affecting mate guarding, but that breeding synchrony alone can not be used to predict which females are more likely to engage in extra-pair mating, nor with which extra-pair males they will mate. Understanding why extra-pair mating by older males is less affected by breeding synchrony may explain much about both the proximate and ultimate causes of extra-pair mating in red-winged blackbirds. Received: 7 June 1996 / Accepted after revision: 25 November 1996     

Habituation and song repertoires in the great tit

Summary This paper is concerned with the idea that the song repertoires of passerine birds are an evolutionary adaptation to reduce habituation in listeners. In an experiment involving 16 territorial males I played either a single song or a repertoire of songs for 15 two-min trials through a single loudspeaker near the edge of the territory. In a second experiment with 10 birds I played the songs through one of two loudspeakers in different parts of the territory, alternating between loudspeakers on successive trials. The birds tended to habituate more rapidly to single song playback than to repertoires. In the second experiment the overall level of habituation was lower and the difference between the two treatments was less marked.Two features of song repertoire organisation are consistent with the habituation hypothesis (1) the avoidance of low recurrence intervals in switches between song types, and (2) the fact that within repertoire variability is as great or greater than between repertoire varability.The main problem with the habituation hypothesis is that habituation by listeners does not seem to be adaptive, so it is not clear why they should habituate. I suggest a hypothesis. Intruders may assess the density of birds in an area by listening to songs, so that habituation may be a mechanism by which this density assessment is achieved. Repertoires could be a mechanism by which resident birds cheat, through increasing the apparent density of singing birds.     

Carotenoid-based status signalling in red-shouldered widowbirds (<Emphasis Type="Italic">Euplectes axillaris</Emphasis>): epaulet size and redness affect captive and territorial competition

In breeding plumage, the African male red-shouldered widowbirds (Euplectes axillaris) are black except for red carotenoid-based epaulets ('shoulder patches'), similar to the well-studied American red-winged blackbirds (Agelaius phoeniceus). To experimentally test the signal function of the red epaulets in male red-shouldered widowbirds, we manipulated epaulet colour and size (within natural variation), and observed the effects in two competitive contexts. First, in captivity, unfamiliar males with epaulets of different size or redness were staged in dyadic contests over food. Only epaulet manipulations significantly affected contest outcome; red epaulet males out-competed orange and control males, which in turn dominated blackened (no epaulet) males. Epaulet size manipulations had particularly strong effects; enlarged epaulet males vigorously defeated smaller epaulet males within colour treatments, whereas the independent effect of 'redness' (i.e., within size treatments) was much weaker. Second, in a field experiment, epaulets were manipulated prior to territory establishment. Males with enlarged epaulets acquired territories, whereas most males with blackened or reduced signals failed to obtain territories. Furthermore, among the territorial males, birds with enlarged epaulets defended larger territories and spent less time in boundary signalling, yet they received fewer intrusions from conspecifics. In contrast, epaulet manipulations had no effect on either female visitation rates or subsequent female settlement. These results concur with recent studies and suggest that the carotenoid-based epaulets of red-shouldered widowbirds are status signals, sexually selected through male contest competition for territories.     

Competition for territories in red-winged blackbirds: Is resource-holding potential realized?

Summary In resource-defence mating systems we intutively expect that the most competitive males should win the best resources so that, in territorial species, male quality and territory quality will be highly correlated. In the polygyny threshold model (PTM) of mate choice this expectation has become an assumption. We performed a removal experiment using red-winged blackbirds to test the validity of this assumption. On the basis of two morphological and two behavioral indices of competitive ability we found only weak correlation between male competitive ability and territory quality. Factors potentially contributing to this result include habitat quality perception, site dominance, and site fidelity. However, both our experimental design and measurement of male competitive ability may have caused us to underestimate the real correlation between male quality and territory quality. Nevertheless, our results suggest that male quality is not perfectly correlated with territory quality as is assumed in the PTM. Therefore, male quality and territory quality should be treated independently when modelling mate choice in this and other territorial species.     

Color bands function as secondary sexual traits in male red-winged blackbirds

Summary We present the first experiment to assess band color effects in a natural bird population. 38 territorial male red-winged blackbirds (Agelaius phoeniceus) were randomly assigned to 1 of 2 banding treatments. They received either all red bands (to match their epaulets) or all black bands (controls). Over half the red-banded males lost their territories while all black-banded males retained their territories. Red and black-banded males did not differ morphologically. However, among red-banded males, those that lost their territories had larger epaulets and were in poorer condition than those retaining their territories. Red-banded males suffered much higher intrusion rates, particularly by neighbors, than black-banded males. We propose that red color bands exaggerated the males' natural aggressive signal beyond the point where the signal was reliable. Our results suggest that signal reliability is maintained by regular testing, particularly of those males most likely to be signalling dishonestly. Offprint requests to: K.J. Metz     

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.     

An experimental study of song type function in the bobolink (Dolichonyx oryzivorus)

Summary We tested the hypothesis that the alpha and beta songs of male bobolinks (Dolichonyx oryzivorus) have separate intrasexual and intersexual functions by presenting caged males and caged females on the territories of male bobolinks. Although all males in our study population share both alpha and beta songs, we found no evidence for a specialized intersexual function of alpha song or a specialized intrasexual function of beta song. Territorial males increased their singing rate of both alpha and beta songs during both the male presentations and the female presentations. The proportions of the two song types did not change between control and experimental periods for either male or female presentations. We found no evidence in support of specialized intrasexual and intersexual functions of note types. Only 10% of the vocalizations initiated by territorial males were complete alpha or beta songs, while 87% were song fragments, and the remaining 3 % were compound songs. We did find that territorial males responded to simulated male intruders by increasing the proportion of fragmented songs and decreasing the proportion of compound songs. Moreover, males included fewer notes per song initiated and sang a higher proportion of songs that included only introductory notes during male presentations. We conclude that the vocal response of territorial male bobolinks to conspecific male intruders is to sing short, simple sequences of either song type. We found no consistent changes in vocal behavior of territorial males in response to female presentations other than increases in song rates.     

Pair and extra-pair mating success relative to male quality in red-winged blackbirds

We tested whether the reproductive success of male red-winged blackbirds (Agelaius phoeniceus) varied with male secondary sexual traits or with haematozoa prevalence, and whether these patterns were consistent with females preferring genetically superior males. We also determined whether the traits that correlated with male success on their own territories were also correlated with male success at siring young on other males territories. Our analysis included data from a 6-year study involving 617 nestlings for which paternity was determined by DNA profiling. Larger males sired more young on their territories, principally because they obtained larger harems. The success of larger males at acquiring more mates did not appear to be a consequence of larger males holding larger or better-quality territories. Older and longer-lived males sired more young by extra-pair fertilizations. Larger males sired the most offspring overall (on territory + off territory). Variation in epaulet size and color, responses to male and female models, nest defence and parasitism was not correlated with male success either on or off their territories or overall. Male success in a given year was significantly correlated with success the previous year, as expected if females were selecting genetically superior males. The male that was by far the most successful individual in this study was highly consistent from year to year. Because male body size is positively correlated with survival in this population (although not within the sample of males included in this study), female preference for larger males may have reflected a general preference for males with superior Survival ability. We propose that the direct advantage realized by older males in extra-pair matings might indicate that experience is important, such that experienced males are better at creating or exploiting the opportunities for extra-pair mating. This hypothesis is consistent with a pairwise analysis of cuckoldry that showed that cuckolders were most often older than the males they cuckolded.     

Playback and speaker-replacement experiments on song-based neighbor,stranger, and self discrimination in male Red-winged Blackbirds

Summary Two experiments were conducted to deterine whether territorial male Red-winged Blackbirds discriminate among neighbor, stranger, and self songs. In the first experiment, recordings of the three classes of songs were played to territorial males. Responses to playback of stranger song, as measured by song rate, intensity of Song Spread display, and closest approach, were significantly stronger than to playback of neighbor song. In addition, stranger song elicited significantly more intense Song Spread displays than did self song.In the second experiment, territorial males were removed from their territories and each was replaced with speakers broadcasting recorded songs of a stranger, and of the removed male. Broadcasts of the removed male's own songs were more effective in discouraging trespassing by neighboring male redwings than were broadcasts of songs of a stranger. However, the two broadcasts did not differ in their ability to deter nonneighboring male redwings from flying through or trespassing within the speaker territory.We suggest that neighbor-stranger discrimination in the Red-winged Blackbird develops as a consequence of associative learing.     

Recognition of individual neighbors by song in the song sparrow,a species with song repertoires

Summary Previous theory and research have suggested that bird species with song repertoires in general, and song sparrows (Melospiza melodia) in particular, cannot readily discriminate between the songs of neighbors and strangers. In a recent study (Stoddard et al. 1991) we showed that song sparrows can in fact discriminate neighbors from strangers on the basis of song. In this study we sought to demonstrate that song sparrows can make the finer discrimination between individual neighbors and that they can do so on the basis of a single song type. We compared the response of territorial males to song playback of neighbors and strangers at three locations: the neighbor's regular boundary, the opposite boundary, and the center of the territory. The birds showed strong neighbor-stranger discrimination at the regular boundary but not at the opposite boundary, nor in the center of the territory. The differences in song discrimination between different boundary locations indicate that song sparrows associate particular songs with particular territories, effectively discriminating between individual neighbors on the basis of song. Song repertoires themselves do not interfere with neighbor recognition to the extent originally postulated. As speakers are moved inside the territory from the border, however, the degree of discrimination diminishes. We believe that differences in speaker placement may have contributed to the variability in neighbor-stranger discrimination observed in previous studies of the song sparrow and perhaps other repertoire species as well. This interpretation is consistent with data from another song sparrow population showing that half the territory takeovers are by immediate neighbors.     

Multiple song repertoire characteristics in the willow warbler (Phylloscopus trochilus): correlations with female choice and offspring viability

Male willow warblers have song repertoires which vary in complexity along several dimensions. We examined whether female choice, as measured by date of pairing, was based on these song characteristics in 4 different years. Pairing date was negatively correlated with song repertoire size in 1 year, and with song versatility in another year, but there was no consistent effect of any song characteristic on pairing throughout the years or in the pooled sample. The variable that best explained how soon a male pairs is male arrival date (only males that had settled territories before the first female arrived were considered in the analysis). This correlation is consistently significant in all years. This is most parsimoniously interpreted as females choosing some habitat characteristic in the same way that males do. A small percentage of males (8.3%) attracted and paired with a second female. The likelihood of becoming polygynous was not explained by any measured song characteristic, but it was related to arrival date: early males were more likely to pair with two females. Males with large repertoires fledged more young in their primary nests, and there was a trend for the offspring of these males to have a greater probability of being recruited into the population. In conclusion, the results show that in most years there is no sexual selection by female preference on song characteristics, although the data on reproductive success is consistent with the idea of repertoire size being an indicator of male quality. Received: 4 June 1999 / Received in revised form: 1 December 1999 / Accepted: 31 December 1999     

Song overproduction and selective attrition lead to song sharing in the field sparrow (Spizella pusilla)

Summary Male field sparrows settling on particular territories for the first time often sing two or more song types. Vocal experience with neighboring males then leads in most cases to the deletion from the male's repertoires of all but one song type. In 2 years, settling males retained the song type that most closely resembled that of a neighbor. Focal observations in 1 year revealed that the most actively singing neighbor was the one the new birds resembled. Songs were not selected on the basis of their absolute acoustic characteristics, but rather by their resemblance to neighbors' songs. After the attrition process is complete, males retain their song for the rest of their lives. Evidence supports the hypothesis that the actual acquisition of a song need not take place at the time and place as males occupy territories. Rather, males may acquire an abundance of song models earlier in life and then select songs from this repertoire that most closely resemble those of their neighbors. The development of mature repertoires and of local dialects is the outcome of one process in this species.