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.     

Surplus territory contenders in male red-winged blackbirds: where are the desperados?

Summary We tested whether payoff asymmetries could explain why floater red-winged blackbirds seldom escalate contests to acquire territories. We removed territorial male \ldowners\rd and allowed floater replacements to claim territories. We then released owners to see how three currencies of payoff asymmetries (replacement occupancy) owners' likelihood and speed of reclaiming their territories (owner success). Neither the duration of an owner's internment nor the amount of time that a floater replacement had held the territory affected owner success (Figs. 1 and 2). Owner success was also not affected by the number of neighbors that they had (Fig. 3). Finally, owner success was the same irrespective of whether or not he was the likely sire of offspring on the territory, or the size of his harem (Fig. 4). Although these results are consistent with the proposal that payoff asymmetries will be irrelevant in contests for valuable resources, they are inconsistent with the proposal's corollary that excluded individuals should become \lddesperados\rd and escalate or even fight to the death in contests for those resources. Expected payoffs for passive acquisition of territories in this species may be higher than from a more aggressive desperado strategy.Correspondence to: P.J. Weatherhead     

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.     

Role of female territoriality in social and mating systems of Canthigaster valentini (Pisces: Tetraodontidae): evidence from field experiments

Sharpnose puffers, Canthigaster valentini (Pisces: Tetraodontidae) at Lizard Island, Australia, live in made-dominated haremic social and mating systems. The hypothesis was that mature females are restricted in their movements and can be monopolized by some males. Field experiments at Lizard Island, Great Barrier Reef, between January and March 1983 showed that mature females were still territorial in the absence of males and movements of females were not controlled by males. Males abandoned their territories when their females were removed. The territorial behavior of those males with access to females (territorial males) restricted the access of other males (bachelor males) to them. Bachelor males took over harems and became territorial males when established territorial males were removed. The results of the experiments thus supported the hypothesis.     

Residence duration influences the outcome of territorial conflicts in brown trout (<Emphasis Type="Italic">Salmo trutta</Emphasis>)

From the value asymmetry hypothesis, we predicted that increasing residence duration should increase perceived territory value, which should (1) increase the motivation of a replacement owner to defend the territory, (2) decrease the probability that the original owner regains the territory and (3) increase contest duration and aggression between the original and replacement owner. These predictions were tested on young brown trout. First, individual fish were allowed to establish territories for 3 days. These original owners were then moved to an adjacent territory, whereupon replacement owners were allowed to take up their territories for either 2 or 4 days. After 4 days, the original owners were returned to their old territories, now containing a new size-matched resident. The resulting conflict was observed and the initiator, duration, aggression levels and the winner of the contest were determined. Replacement trout that had been resident for 4 days initiated and won more contests against original owners than did 2-day replacements. This suggests that the motivation to defend the territory increases with residence time, which will determine the outcome of territorial contests between opponents of similar resource-holding potential (i.e. size). Moreover, contests with 4-day replacements were longer and aggression levels were higher than in contests with 2-day replacement trout. These results are consistent with the value asymmetry hypothesis, extending its validity for explaining the behaviour of territorial animals.     

Song repertoires and density assessment in red-winged blackbirds: further tests of the Beau Geste hypothesis

Summary The Beau Geste hypothesis proposes that song repertoires are advantageous in territory defense because nonterritorial males, who are prospecting for territories, use the number of song types they hear to assess the density of territorial males, and then avoid densely-settled areas. A territorial male can then inflate the apparent density of singers on his territory by singing several distinct song type. This hypothesis assumes (1) a positive correlation between the density of song types and the density of territorial males, and (2) a negative correlation between the rate of trespassing and the densities of both song types and territorial males. We studied the behavior of male red-winged blackbirds (Agelaius phoeniceus) and found support for the first, but not the second, assumption. Results showing a positive correlation between density of song types and density of territorial males show that the proposed method of density assessment is feasible in this species. However, we could find no support for the assumption that nonterritorial male red-winged blackbirds avoid densely settled areas. In contrast to the assumed negative correlation, rate of trespass per territory was not consistently correlated with male density, and trespass rate per unit area was positively correlated with male density. Further, these relationships were retained when we controlled statistically for the effects of territory quality. We conclude that prospecting male red-winged blackbirds to not attempt to avoid densely settled areas, and that, although they do avoid territories defended by many song types, they do not use song type density to assess the density of territorial males. It is thus unlikely that the Beau Geste hypothesis adequately accounts for the evolution of song repertoires in the red-winged blackbird.     

Territorial defence in the great tit (Parus major): Do residents always win?

Summary I removed resident pairs of great tits from their territories for short periods and released them after replacement pairs had occupied the spaces. When two pairs are manipulated in this way into occupying the same territory an escalated contest ensures. Contests between residents and replacements are longer, more likely to involve physical fights and longer fights or displays than contests between established neighbours or residents and intruders. The degree of escalation between residents and replacements is an increasing function of replacement time. It rises to a peak and then diminishes again. The probability that the replacement will defeat the former resident is an increasing function of replacement time, reaching 90% after several days. These results are consistent with the hypothesis that territorial residents win in contests against intruders because of an asymmetry in payoff rather than an asymmetry in resource holding potential or an arbitary convention. A possibly asymmetry in payoff in the great tit is the cost of defending the territory against neighbours. The cost is higher for replacements than for residents.     

Territorial fidelity of willow warbler (Phylloscopus trochilus) males and success in competition over territories

Summary Territorial fidelity and competition between males was studied in a population of individually marked willow warblers (Phylloscopus trochilus). After spending the winter in Africa males arrive in the study area in late April or the beginning of May. They can be separated into two categories, old colour-banded males that return for a new breeding attempt, and birds that are new to the study area. These new birds could either be birds that had bred the previous year returning to a different site, or more likely 1-year-old birds returning for their first breeding attempt. Old males show high site fidelity and are successful in retaining their territories from the preceding year. The few old males that switched territories between years seemed to do so in response to reduced breeding success in the preceding year. However, in 13 out of 14 cases when a new resident was already established on the previous resident's territory, the old male was successful in evicting the new resident. In a removal experiment both old and new birds were removed from their territories and then released when a replacement male had been established. All old males successfully managed to reclaim their original territories after escalated contests. None of the new males were successful and no fights were observed.     

How reliable are behavioral cues for assessment of male mating status in polyterritorial wood warblers,Phylloscopus sibilatrix?

It has been suggested that polyterritoriality in birds, i.e. mated males defending two spatially separated territories, deceives females into accepting polygyny against their best interest. Mated males singing on secondary territories should make it difficult for new arriving females to assess the mating status of unmated and mated males advertising to females. However, there have been objections to the deception hypothesis, suggesting that differences in territorial behavior between males of different mating status may make correct assessment possible. Polyterritorial males frequently leave their secondary territory to visit the female in the primary territory, thereby revealing their mating status to females in search of mates. The aim of this study is to investigate how reliable behavioral cues are in assessing the mating status of males in polyterritorial wood warblers. Our study shows that singing activity of mated males on secondary territories and unmated males was similar in the morning, while there was a significant difference in the afternoon. Unmated males spent, on average, more time on their territories than mated males spent on their secondary territories. However, there was considerable variation in both unmated and mated males in the frequency in which they left their territories. For instance, unmated males left their territories quite frequently early in the season compared to later. Hence, females must be able to interpret singing behavior and a male leaving his territory differently depending on time of the season and time of the day in order to correctly assess the mating status of advertising males. Estimating the increase in probability of finding an unmated male by using behavioral cues showed that females could increase the probability of finding an unmated male compared to random choice with respect to mating status. The uncertainty was, however, still considerable. The most important factor affecting the probability of finding an unmated male was the proportion of mated males on secondary territories compared to unmated males. Our study suggests that presence on a territory is not a reliable cue for the assessment of male mating status in polyterritorial wood warblers.     

Mating system shift in a pronghorn population

Summary At the National Bison Range (western Montana, USA), pronghorn (Antilocapra american) maintained a territorial mating system from as early as 1965 through 1978. Preliminary observations in 1981 suggested that the mating system had changed. Data from the ruts, 1982–1984 revealed a progressive decay in territoriality. In this paper, data from the territorial years 1969–1978 are contrasted with data from the decay years 1982–1984, with results as follow: 1. In 1982–84, fewer territories were defended than in 1969–78. This was attributable to a smaller proportion of males defending territories in 1982–84, not to a smaller number of males. 2. In 1982–84, most territory owners either abandoned their territories early in ruts, or lost control of females on them following frequent, persistent intrusions by non-territorial males. Abandonment and loss of control did not occur in 1969–78. 3. In 1982–84, territorial males that maintained control of females on their territories did so by shrinking their zones of defense to small areas around female groups. 4. In 1982–84, following the disruption or severe disturbance of all territories, many females left territories, and mated elsewhere, with non-territorial males. In 1969–78, most females remained on territories throughout rut, and mated with territory owners. —The mating system change followed catastrophic winter mortality, 1978–79, that removed 75% of the males, including all males older than 5 years, and all male fawns, from the population. In 1982–84, the number of males present was not different from the number of males in 1969–1978, but the frequency distribution of male ages was strongly shifted toward younger ages. The small number of older males, 1982–84, likely resulted in smaller proportions of males initially defending territories, and in less effective territory defense. When females then clustered on the few territories where defense was at first successful, they attracted large numbers of non-territorial males. The resulting high rates of raids on these territories, coupled with reduced defense radii by territorial males, allowed females only 12% reclining time (summer percentage was 39%). This increased energy cost, plus an apparently greater risk of injury on weakly defended territories, appeared to prompt many females to seek calmer matings elsewhere. Also, if female pronghorn practiced mate selection based upon horn or body size, they may have reduced their efforts to remain on territories in 1982–84. Males from 1969–78 were larger than males from 1982–84, and showed greater variance in horn size. At least two conditions appear to influence the tendency of males to be territorial. First, males must be at least three years old before they attempt to defend a territory. Second, the declining proportion of males defending territories, 1982–84, that coincided with an increasing number of males three years and older, suggests that males also decide whether or not to attempt territory defense based upon the frequency of territorial defense in the population.     

Mate choice in the willow warbler — a field experiment

Summary Females may choose a mate on his own quality or on the quality of his resources, i.e., his territory. We removed willow warbler males and allowed new males to settle, before the arrival of females, in order to test whether the proximate cue for female choice was any male trait or territory characteristics. The experiment indicates that females base their choice on some male trait. No correlation was found between male settlement order and size (tarsus length, wing length), but males arriving early were in better body condition than males arriving late. The most likely male trait for female choice was singing rate, which was a good indicator of male settlement date. The duration of time between mating and egg laying was shorter in early territories, both in the observational and the experimental data. Since male song rate was correlated with territory quality (e.g., food abundance) the ultimate benefit to females choosing males with high song rates could be a high quality territory.Offprint requests to: B.L. Arvidsson     

Dominance in feeding territories relates to foraging success and offspring growth in brown skuas <Emphasis Type="Italic">Catharacta antarctica lonnbergi</Emphasis>

Territoriality should lead to strict dominance, as territory holders typically control access to resources and exclude others from their use. In feeding territories, dominance should be reflected in foraging success and ultimately in reproduction differences; however, these successive links have rarely been made explicit. Therefore, we investigated a population of brown skuas Catharacta antarctica lonnbergi, in which only part of the breeding population occupied feeding territories within penguin colonies. We identified the dominance hierarchy and determined the foraging success of the participants in fights for access to penguin carcasses within the territories. Furthermore, we monitored offspring growth from parents with and without feeding territories. Our results indicated a clear dominance hierarchy with territorial birds in their own territory dominating over territorial breeders from other territories, non-territorial breeders and non-breeding birds. However, territory owners could not completely exclude others from access to food. Foraging success was positively related to dominance scores: The dominant territory owners received 63% of a carcass, whereas non-territorial pairs could get less than 10%. The link between foraging success and offspring development was less clear: Although male chicks of non-territorial parents suffered from lower growth rates and, thus, delayed fledging, there were no such differences in female chicks. Territoriality in skuas did not imply a complete occupation of food, but guaranteed optimal growth conditions for offspring. Non-territorial individuals were forced to search for alternative resources, and the restricted access to the preferred food resulted in inferior conditions for offspring development, making this foraging strategy less rewarding.     

Density dependence and habitat quality modulate the intensity of display territory defence in an exploded lekking species

We evaluated the effect of conspecific abundance and habitat quality of leks on the territorial behaviour of males in an exploded lekking species, the Little Bustard (Tetrax tetrax). The hypothesis that males more intensely defend territories with higher conspecific abundance and better habitat quality was evaluated experimentally analysing the agonistic response of experimental males to male decoys placed on their displaying areas. Decoy experiments showed that the intensity of display territory defence by little bustard males is density dependent. The time experimental males took to return to their display sites after decoy placement decreased with abundance of both males and females. The strength of their final response was positively associated to local male and female abundance in the vicinity of their display sites. Habitat quality also influenced males’ display territory defence since the intensity of male response increased with the degree of natural vegetation cover. Habitat quality was particularly relevant in explaining variation of experimental males’ snort call rate, which decreased with the degree in plough cover and increased with the number of fields in the lekking area. Snort call rate decreased with the level of male aggregation and was lowest in males exhibiting the strongest aggressive response to decoys. These results add new evidence for the density dependence of species’ breeding territorial behaviour, supporting density-dependent models of lek formation and reinforcing the role of resources defence in exploded lek mating systems.     

Territory defence in tropical birds: are females as aggressive as males?

Much of our knowledge concerning the functions of territorial behaviour and how territories are defended by individuals comes from research on birds. The vast majority of this work has focused on temperate zone breeding territoriality in which territories are defended most obviously by males. Our understanding of the female role in territory defence is limited because they are less conspicuous and much harder to observe. We studied sex roles in territory maintenance and defence in a duetting, resident neotropical passerine, the white-bellied antbird (Myrmeciza longipes). This species maintains territories and pair bonds year round and both sexes sing and actively participate in territory defence. We performed a series of playback experiments throughout the dry (non-breeding) and wet (breeding) seasons. We exposed territorial pairs to three types of stimuli including: (1) single sex, male only songs, (2) single sex, female only songs, and (3) both sex songs/duets. Contrary to findings for most other tropical species, individuals defended their territories with equal levels of aggression regardless of stimuli. Furthermore, sex roles were very different, with males responding more aggressively than females to all stimuli throughout both seasons. Both males and females consistently responded more aggressively to territorial intrusions during the dry season than during the wet season, likely because food abundance is low in the dry season and territory value is high. Our analysis of duetting behaviour suggests that duets do not serve a significant role in mate guarding, or territory defence.     

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     

Effectiveness of territorial polygyny and alternative mating strategies in northern fur seals, <Emphasis Type="Italic">Callorhinus ursinus</Emphasis>

We conducted a 6-year longitudinal behavioral and genetic investigation of a highly polygynous pinniped, the northern fur seal (Callorhinus ursinus), to determine the contribution of terrestrial polygyny to male fertilization success and to assess the occurrence of alternative mating strategies. Genetic samples from 37 adult males, 50 adult females, and 85 pups were collected and genotyped using five polymorphic microsatellite loci. Pup paternity was assigned using Cervus 2.0 at 99% confidence level. Paternity of 83 pups (98%) was assigned to terrestrial males who held territories or stayed temporarily in the study area during the breeding season when fertilization occurred. For 56 pups of which attendance records of their mothers were available, paternity of 45 pups (80%) was assigned to the associate males in whose territory their mothers stayed during the perioestrus period. In addition to defending breeding territories, territorial males have often been observed attempting to forcibly abduct adult females from adjacent territories (female stealing): We observed a total of 95 such cases, in which the stealers had significantly fewer females than the territorial males from whose territories they stole females. Our results indicate that terrestrial resource-defense polygyny is the major mating system in this species and that nonassociated paternity occurs mostly as a result of alternative mating strategies of less successful males. Male northern fur seals thus appear to adopt conditional alternative strategies that depend on their current social status to maximize their life-time reproductive success.     

Acoustically mediated neighbor recognition in the North American bullfrog,Rana catesbeiana

Summary Playback experiments were conducted to determine whether territorial male bullfrogs (Rana catesbeiana) were capable of discriminating between advertisement calls of neighbors and strangers. Territorial males adjacent to subject males were removed and replaced with a speaker which broadcast calls of a stranger or the removed male. Ten of 11 males responded more strongly to calls of strangers than to calls of neighbors. When advertisement calls of neighbors were broadcast to subject males from positions within the removed neighbor's territory and from positions opposite the subject male, all 9 males tested responded more strongly to calls of a neighbor broadcast from a new position than to playbacks of the same call broadcast within the removed neighbor's territory. Because male bullfrogs recognize familiar calls and associate these calls with a particular direction or location, these data provide the first evidence for acoustically mediated neighbor recognition in frogs.     

Influence of territoriality on adult density in two rockfishes of the genus Sebastes

Nonterritorial Sebastes carnatus and S. chrysomelas existed, along with territorial individuals, at 3 tagging sites off southern California, USA, which were monitored for nearly 1 yr. To test the hypothesis that territoriality affected adult density in these species, territorial fish were removed and the subsequent utilization of vacated territories by other fish was monitored. Intrusion of other fish into vacated territories increased significantly in 90% of the removals. Other fish colonized both the feeding and sheltering parts of the vacated territories, indicating that the previous owners had successfully defended both parts of their territories. Many of the colonizers had already possessed territories; they expanded their territories or moved into presumably better havitat. Several previously-nonterritorial fish also moved into vacated areas, and at least some of them appeared to establish territories. These fish, then, had previously been capable of establishing territories, but were prevented from doing so by resident territory holders. Thus territoriality, rather than such other factors as predation or low recruitment, limited the number of territorial fish at each site. However, territorial fish did not inhibit the settlement of larval recruits, and the relative mortality rates of older territorial vs nonterritorial fish were not determined. Thus the question of whether territoriality was a major factor controlling total density remains unresolved.     

Breeding territory quality and agonistic behavior: effects of energy availability and intruder pressure in hummingbirds

Summary The relative importance of food a vailability and intruder pressure on breeding territorial behavior was studied in two experiments with male calliope hummingbirds. In the first experiment, extra food was provided in inconspicuous feeders. Territory owners who fed from the feeders increased their display rates (power-diving and hovering) when on their territories, and spent less time out of sight (mostly off their territories). Food addition by this method had no significant effect on intruder pressure, as measured by number and duration of chases. In the second experiment, intruder pressure was increased by attracting feedertrained males to feeders in territories of individuals who did not feed from feeders. In this experiment, display activity of residents did not change but chasing activity increased. These results suggest that display rates by male calliope hummingbirds depend at least in part on the amount of food available, but are not strongly affected by the number of feeding attempts by male intruders. No attempt was made to determine whether displays can deter competitors, attract mates, or do both. However, since these energetically expensive displays depend at least in part on the energy available to owners, they provide readily available indices of male and/or territory quality which could potentially be used for different purposes by different individuals.     

A same-sex stepparent shortens a prebreeder's duration on the natal territory: tests of two hypotheses in Florida scrub-jays

Prebreeders of the Florida scrub-jay (Aphelocoma coerulescens) are less likely to be found on their natal territories with a same-sex stepparent than with parents or an opposite-sex stepparent. We tested two models that had been proposed to account for this sexual asymmetry. The dominance hypothesis states that stepparents perceive same-sex prebreeders as competitors, primarily for a mate, so behave aggressively toward them. The pair-formation hypothesis states that prebreeders remain home to pair eventually with the opposite-sex stepparent. Predictions from these two models were tested by analyses from a quarter-century of records on a study population at Archbold Biological Station and by new behavioral field observations at the Station. Results clearly rejected the latter and strongly supported the former hypothesis. No prediction from the pair-formation hypothesis was confirmed: no pairing by a prebreeder of either sex has ever occurred with a stepparent; remaining home was equally frequent in age-one males with stepmothers and genetic mothers, and in females with stepfathers and genetic fathers; and the same results were found in age-two prebreeders. By contrast, the data strongly supported the dominance hypothesis. Field data showed higher aggression rates by stepfathers to male prebreeders and stepmothers to female prebreeders than by parents to their same-sex genetic offspring. Fewer age-one males remained home with a stepfather than with the genetic father, and the same was found for age-one females with a stepmother and genetic mother; at age two, the effect occurred only in female prebreeders. The two hypotheses make different predictions about prebreeders with two stepparents versus with both parents, thus providing a critical test. The dominance hypothesis correctly predicted decreased duration at home by age-one males and females; data for age-two females were in the direction of predicted difference but not significant; and, as in other tests, no effect was found for age-two males. We propose that age-two male prebreeders remain home despite elevated aggression from stepfathers because these prebreeders retain the possibility of budding from or inheriting the natal territory. Received: 16 July 1997 / Accepted after revision: 26 June 1998