Owner and floater red-winged blackbirds: determinants of status

Summary We used sequential removal experiments to test whether the resource-holding potential (RHP) of territory owner red-winged blackbirds, Agelaius phoeniceus, was superior to that of their first replacements (shallow floaters) and subsequent replacements (deep floaters). Among the removals were secondyear males, which were morphologically inferior to adults and which also tended to be competitively inferior in aviary contests. The highest proportion of secondyear males occurred in the deep floater class. Thus, the RHP of some deep floaters was inferior to that of owners and shallow floaters. However, among adults, owners, shallow floaters, and deep floaters had equivalent morphological and competitive RHP. Furthermore, replacement males that had defended territories for many days were neither morphologically nor competitively superior to males that had defended territories for only a few days. Our results suggest that RHP distinguishes adults from second-year males, but does not separate owners from floaters. The only hypothesis that is potentially supported by our observations is that owners have a greater expected payoff from their territory than intruders, and in this way owners are able to maintain site dominance. The nature of that payoff remains to be determined. Offprint requests to: P.J. Weatherhead     

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.     

Habitat preference increases territorial defence in brown trout (Salmo trutta)

The striking ability of territory owners to repel intruders has generated a number of theoretical explanations as well as experimental studies in many animal species. However, effects of individual habitat preferences on territorial defence have rarely been studied. From the territory value hypothesis, we predicted that owners of preferred habitats should invest more resources in defence than owners of non-preferred habitats. We tested this prediction with young territorial brown trout in a two-stage experiment. First, trout were allowed to choose individually between gravel and a uniform bright substrate. As expected, they showed a significant (79%) preference for gravel. However, there was considerable variation between individuals in substrate preference, with a few fish preferring the bright substrate. Half of the tested fish were then transferred to a gravel substrate and the rest to a bright substrate, manipulating habitat type in relation to preference. Territory owners were then staged against size-matched intruders whereupon contest aggression was observed and the winner of each contest determined. Overall, owners won most of the contests. Satisfied owners won 86% and owners of less preferred territories, 74% of the contests. Furthermore, more satisfied owners attacked sooner and were more aggressive relative to the intruders. We conclude that brown trout show individual variation in habitat preference, which appears to be linked with their investment in territorial defence. These results suggest that understanding and modelling of animal contests could benefit from considering how territorial defence is influenced by individual habitat preference and specialisation. Received: 25 February 2000 / Revised: 22 May 2000 / Accepted: 25 June 2000     

Contest behaviour in the speckled wood butterfly (Pararge aegeria): seasonal phenotypic plasticity and the functional significance of flight performance

Although contemporary animal contest theory emphasises the importance of physical asymmetries in resolving disputes, such asymmetries do not obviously settle fights in all groups. Territorial male butterflies, for example, compete via elaborate non-contact aerial interactions in which success is determined by relative persistence. Prior research suggests that the resolution of these contests is not clearly related to physical variables such as body size or energy reserves. However, given that the contests involve elaborate aerial manoeuvres, one long-standing suggestion is that asymmetries in flight performance, and thus flight morphology, may be important. We addressed this hypothesis via a manipulative investigation into the biophysical correlates of contest success in the speckled wood butterfly, Pararge aegeria. This species possesses the ability for significant adaptive phenotypic plasticity in relevant flight morphological parameters. We took advantage of this plasticity to rear 90 individuals of markedly varying flight morphologies, which we then pitted against each other in a semi-controlled experimental fashion. Multiple logistic and lognormal analyses provided little evidence for the relevance of morphological parameters, including relative flight musculature, wing loading and wing aspect ratio (wing length relative to area), to the outcome and/or duration of experimental contests. Instead, we found a positive effect of age upon contest success. Given that ability for high acceleration is strongly linked to variation in these morphological parameters, our findings suggest that flight performance is not a strong determinant of resource-holding potential in this notably territorial butterfly.     

Predation risk and the territory value of cover: an experimental study

In an experiment on territorial brown trout (Salmo trutta), we addressed the novel hypothesis that protective cover increases the value of a territory in relation to the perceived level of predation risk. We predicted that territory holders should invest more resources defending territories with cover than territories without cover, and that defence should increase as predation risk increases. First, trout were allowed to establish ownership in territories with or without overhead cover. Second, predation risk was manipulated by simulating aerial predator attacks in half of the territories of each type, whereupon the preference for cover was estimated. Third, owners of the four types of territories were staged in dyadic contests against size-matched intruders. Territory owners showed a preference for cover, which increased further after simulated predator attacks. In subsequent contests, conflicts over territories with cover were settled faster than conflicts over territories without cover, which may suggest that the value of cover increases the motivational asymmetry between owner and intruder. Consistent with our hypothesis, owners of territories with cover were much more aggressive if they had been subjected to predator attacks the day before the conflict. These results suggest that territory owners are able to estimate the value of protective cover in response to variation in the level of predation risk in the habitat.Communicated by J. Krause     

Fighting without weaponry: a review of male-male contest competition in butterflies

The males of many butterfly species compete via pair-wise intrasexual contests, yet lack any obvious morphological traits conventionally associated with animal aggression. The evolution of contest behavior in this group is therefore interesting because the means by which individuals are able to inflict costs upon each other during combat are unclear. Here we review the relevant empirical information within a game-theoretical context in order to analyze contest-related costs and mechanisms of dispute settlement. Territorial butterfly contests carry obvious role asymmetries, and individuals in the 'resident' role are more likely to win across all studied species. We use this phenomenon as a framework for gaining further insight into the evolution of contest settlement in this group. Four principal hypotheses are evaluated: (1) that residency is used as an arbitrary convention, (2) that residents stand to gain a greater payoff if successful, (3) that residency bestows higher resource-holding potential (RHP) on males in that role, and (4) that residents are males of intrinsically higher RHP. We found most support for the latter explanation (the 'superior-competitor' hypothesis), partly because the alternatives require a restrictive set of ecological conditions, but also due to positive support for the idea of intrinsic RHP. The strongest evidence is provided by species in which repeated interactions (between the same individuals) always have the same outcome, regardless of changes in residency status. However, we also found a consistent effect on contest duration due to apparent 'confusion over residency,' which suggests a conventional element to contest behavior. Although butterfly contests must be costly in some way, the literature provides limited insight into how costs accrue, and hence the determinants of contest RHP remain obscure. We suggest that careful experimentation, using 'natural' manipulations where possible, should prove most beneficial in identifying costs and exploring schedules of contest settlement in this group. Moreover, future investigations should be sensitive to the fact that butterfly territoriality and associated male-male interactions are seated within a broader ecological and life-historical context.     

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.     

How do fighting ability and nest value influence usurpation contests in <Emphasis Type="Italic">Polistes</Emphasis> wasps?

Most research on animal contests has focused on the factors that influence the intensity and outcome of aggressive contests within nonsocial species, while relatively little is known about contests in social taxa. Here, we examine contests among queens of the social paper wasp, Polistes dominulus. Queens use multiple reproductive strategies, including nesting alone, usurping established colonies, and cooperatively joining other queens. We stage contests between a nesting queen and a challenger to test how resource value (RV) and resource holding potential (RHP) influence (a) who occupies the nest at the end of the contest and (b) the extent of conflict between the queen and challenger. We found that RHP, as measured by individuals’ facial patterns and body size, influenced the outcome of the contest. Challengers with high RHP were more likely to successfully usurp the nest than challengers with low RHP. Interestingly, queens with relatively high RHP were more likely to form a cooperative association with the challenger than queens with lower RHP, suggesting that queens may evict individuals that are an aggressive threat. RV influenced the intensity of conflict. There was more aggressive conflict over large nests than over small nests. Overall, social taxa have complex contest dynamics with important parallels to contests in nonsocial taxa. Studying contests in social taxa provides an important perspective on the factors that influence individual decisions about conflict versus cooperation.     

Northern harriers on feeding territories respond more aggressively to neighbors than to floaters

Summary Territory owners often respond less aggressively towards intruding neighbors than towards intruding floaters, an observation termed the dear enemy phenomenon. Comparisons of territory owners' responses to intruding neighbors versus their responses to intruding floaters usually have been made for owners of multi-purpose and/or breeding territories. Here, I describe responses of female northern harriers Circus cyaneus (owners) on winter feeding territories towards three types of intruders (female neighbors, female floaters, and male floaters) and show that the dear enemy phenomenon does not occur. Owners' responses towards neighbors were more intense (mostly flights rather than calls) than responses towards female floaters, which in turn were more intense than responses towards male floaters. The greater intensity of owners' responses towards neighbors compared to owners' responses towards male and female floaters may be related to differences in the threat posed by each of the three intruder types in terms of fighting ability (RHP) and potential losses from intrusion. Hence, whether owners respond more aggressively towards neighbors or floaters, and whether the dear enemy phenomenon is observed, may depend upon the relative magnitude of threat presented by neighbors and floaters to owners in terms of fighting ability and potential losses from intrusion.     

Experimental evidence that bright coloration is not important for territory defense in purple martins

Summary I tested the hypothesis that bright breeding plumage in territorial males acts as a badge of fighting ability or aggressive motivation to intimidate intruders. Territorial male purple martins (Progne subis) whose iridescent blue plumage was lightened to mimic the appearance of subadult males did not suffer an increase in intruder pressure or loss of territory compared with control males. Bright plumage color itself did not deter intruders and was not important for successful territory defense. Furthermore, a bright coloration of owners was not associated with an increased level of aggression toward intruders. Results from parallel studies on this species suggest that bright coloration is important in territory acquisition. The effectiveness of badges of fighting ability and aggressive motivation in territory defense is limited by whether intruders benefit from assessing these traits in owners. Differences in signaling systems between species are due in part to differences in floater tactics and the mode of territory acquisition.     

The reproductive behavior of Anthidium manicatum (Hymenoptera: Megachilidae) and the significance of size for territorial males

Summary Males of the wool-carder bee, Anthidium manicatum, patrol clumps of garden plants. Females of this species visit these plants for pollen, nectar, and pubescence; they also mate there. Females are polyandrous, with intervals between copulations as short as 35 s. Patrolling males defend their territories (0.1–1.3 m²) against other males and against other species of flower-visiting insects. Honey bees may be rendered unable to fly by the attacks of A. manicatum.Territory owners perform exploratory flights to other males' territories, changing territories often (median ownership 4–7 days; maximum 30 days) and flying up to 450 m to establish new territories. Territorial usurpations are nearly always by larger males.Female visitation rate is significantly correlated with number of flowers on a territory. The head size of territory-owner males shows significant correlation with territorial quality (measured by number of flowers, not area) and thus with number of female visits and copulatory opportunities. Some males fail to maintain territories and instead attempt to forage and copulate in other males' territories while the owners are otherwise occupied. Nonowner males are significantly smaller than owners, forage less often and from fewer flowers, and achieve significantly fewer copulations than owners. Females, however, do not reject smaller, nonowner males at a higher rate than they do larger, owner males; their choice for male size appears to be indirect, based instead on choice of food resource.The interval between a copulation and the male's next attempt with a different female is not shorter than that involving the same female. Males do not escort just-mated females about their teritories, as observed in Anthidium maculosum. Territorial behavior in this species most likely evolved through intrasexual competition for reproductive success which led to sexual dimorphism. The defense of a resourcebased territory is the mechanism used by a male to maximize his reproductive potential.     

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     

The adaptive significance of male subadult plumage in purple martins: plumage dyeing experiments

Summary I tested two hypotheses for the adaptive significance of subadult plumage in male purple martins (Progne subis) : the female mimicry and subordinance signaling hypotheses. Subadult males were at a competitive disadvantage in obtaining territories, as they arrived later in the spring than adult males. Contrary to the predictions of both hypotheses, adult male territory owners were not less aggressive toward subadult male than adult male intruders. The subadult plumage was not effective in mimicking females, as adult male owners were significantly more aggressive toward subadult male than female intruders. Summer adaptation hypotheses predict that young males in subadult plumage are more successful in acquiring territories and mates than they would be with an adult plumage. I tested this prediction by dyeing the plumage of floater subadult males to mimic the appearance of adult males. In 13/17 paired experiments, dyed subadults obtained territories before control subadults. There was no difference in the time it took dyed and control males to attract a mate after they obtained a territory. These results suggest that the subadult plumage is not an advantage to young males in competing with adult males for breeding resources. In late winter, subadult males were growing mostly femalelike feathers on their underside, suggesting that the subadult plumage is not the result of a molt constraint. The subadult plumage could enhance survival of yearlings in winter roosts if it improves access to good roost sites or reduces the risk of predation.     

Investment in territorial defence depends on rearing environment in brown trout (Salmo trutta)

In many animals, territoriality will arise or cease depending on environmental factors such as intruder rate and resource availability. We investigated the effect of rearing environment on territorial behaviour in ~1.5-month-old brown trout. In the laboratory, wild-caught (reared at a low density) and hatchery-reared (high density) trout were allowed to defend a territory against a size-matched intruder reared in the same or the other environment. Because territorial behaviour should be relaxed at high-rearing densities, we hypothesized that hatchery-reared trout should value their territories less and therefore invest less in defence compared with wild-caught trout. However, in all cases, territory owners were more likely to win the contest and hatchery-reared trout were just as likely as wild-reared to win mixed contests. Furthermore, pairs of hatchery-reared trout initiated contests sooner, fought longer and were more aggressive during the contest compared with pairs of wild trout. When hatchery-reared owners met wild intruders, the contest ended sooner compared with when the roles were reversed. We conclude that territorial behaviour in brown trout is largely innate, but that the hatchery environment has promoted more aggressive individuals. These results suggest that hatchery-reared trout invest more time and energy to obtain the same contest success as wild trout. In conclusion, the lack of experience of territorial defence in a high-density rearing environment seems to reduce the efficiency of territorial behaviour. In turn, this may have negative consequences for the performance of released hatchery fish in the wild.     

The downfall of mating: the effect of mate-carrying and flight muscle ratio on the escape ability of a pierid butterfly

In many species, the ability to evade predators is known to be periodically impaired by increased weight loads due to feeding and reproduction. Not only may extra weight reduce escape speed, but feeding and mating can also make the prey more noticeable to visually hunting predators. A number of butterfly species mate for hours, and if a mating couple is disturbed, one of the butterflies is responsible for flying, whereas its partner remains still. This study investigated the ability of male Pieris napi butterflies to fly while mating, with the prediction that mate carrying impairs flight ability compared to single flying males and that males with relatively high flight muscle ratios (FMR; male thorax mass/male + female body mass) will have better flight performance in copula. Our results clearly show that whereas single males always take off at steep angles and fly upwards, couples invariably have a negative take-off angle and rarely gain height. Moreover, landing height of the couples is positively associated with higher FMR. Hence, male flight ability when in copula is positively associated with a high relative thorax mass. Butterfly pairs may thus be at greater risk of predation as a consequence of their impaired flight ability, especially couples with critically low FMRs (<16%).     

Territory quality and territorial behavior in two sympatric species of hummingbirds

Summary Changes in territorial behavior of blackchinned hummingbirds (Archilochus alexandri) in response to experimental changes in territory quality were investigated using artificial feeders and simultaneous, pair-wise observations of owners. Some of the responses of A. alexandri were similar to those documented by a previous study of the Anna's hummingbird (Calypte anna): A. alexandri defending rich territories spent more time on their territories, encountered a greater percentage of intruders, and chased more intruders than did A. alexandri defending poor territories. In contrast to C. anna, A. alexandri supplemented chases with energetically inexpensive threat vocalizations more extensively when territory quality was increased. This difference may be related to A. alexandri's more tenuous control of rich territories. When both species were observed on very poor territories, A. alexandri chased a greater percentage of intruders, consumed a greater proportion of available food, and obtained a greater net energy gain from their territories. When observed simultaneously on territories with ad lib food, both species defended highly exclusive territories but A. alexandri suffered higher intrusion pressure and spent more time and energy in defense. These interspecific differences in territorial behavior may facilitate sympatric coexistence of the two species through aggressive partitioning of flower patches according to productivity: the greater payoff received by C. anna on rich territories and A. alexandri on poor territories should favor a corresponding monopolization of rich areas by C. anna and poor areas by A. alexandri.     

Interspecific aggression in Leucorrhinia dragonflies: a frequency-dependent discrimination threshold hypothesis

Summary Male Leucorrhinia dragonflies defend territories from conspecific and heterospecific intruders. Defense against heterospecifics is surprising, as mating and oviposition are the only activities that occur on the territories, and heterospecific males are not expected to pose a reproductive threat. L. frigida and L. intacta males respond aggressively with equal frequency and intensity against intrusions by conspecifics and heterospecifics. In contrast, L. proxima males respond more aggressively against conspecifics. The apparent lack of species discrimination shown by L. frigida and L. intacta males may result because territorial males that assess intruders (as do L. proxima) suffer a tactical disadvantage from hesitating when an intruder flies in. This assessment process may lead to reduced fighting success by the territorial male. Thus there is a tradeoff between assessment cost incurred when the intruder is a conspecific and benefit gained from avoiding conflict when the intruder is a heterospecific. Given this assessment cost, males of species that only rarely encounter conspecific intruders are more likely to evolve species discrimination than males of species that commonly encounter conspecific intruders.     

Twilight fighting in the evening brown butterfly,<Emphasis Type="Italic"> Melanitis leda</Emphasis> (L.) (Nymphalidae): age and residency effects

Butterflies are conspicuous among animals that fight for mating opportunities because it is not clear how contest costs could accrue. Nevertheless, the bulk of research in this group suggests that contests are settled on the basis of asymmetries in fighting ability (the superior competitor hypothesis). There is also a consistent effect on contest escalation due to apparent 'confusion' over residency, which could result from resident butterflies playing different tactics than non-residents, or from a causal link between residency and fighting ability. I addressed these hypotheses by studying contest behavior in the crepuscular species Melanitis leda (L.) (Nymphalidae) over a 5-month period in tropical Australia. Males competed via conspicuous two-stage maneuvers of a form unique among butterflies. Prior residents won 77% of all contests. Non-resident males arrived and perched in occupied sites until challenged by incumbent residents, and the aggressiveness of these interlopers increased as a function of the time before they were detected. Contest winners also tended to be younger than losers, and contest escalation was negatively related to both the age of the losing male and the magnitude of the between-combatant age asymmetry. These results are consistent with the superior competitor hypothesis based upon age as a determinant (or correlate) of fighting ability, but also suggest residency has a unique influence on contest behavior. Further research is clearly required to unravel the effects of age, residency and other potential biophysical determinants of fighting ability in this species.     

Inter-sexual contests in the hermit crab <Emphasis Type="Italic">Pagurus bernhardus</Emphasis>: females fight harder but males win more encounters

Contests often occur between members of the same sex when they compete for access to mates, but inter-sexual contests may occur over access to other essential resources such as food or shelter. Despite the possibility that such contests are common, most studies focus on male fighting, and very few have analysed fights between males and females. Because males and females differ physically and physiologically, fighting ability or resource-holding potential (RHP) may also be subject to inter-sexual variation. In this study, we investigate size-controlled inter-sexual contests over the ownership of empty gastropod shells in the hermit crab Pagurus bernhardus. During these fights, there are two roles, attacker and defender, and we use a fully orthogonal experimental design to compare the performance of males and females in each role, when fighting either a male or female opponent. Although females fight more intensely, male attackers have an advantage when compared to females playing the attacker role, as they are more likely to evict the defender from its shell and thus win the resource. Further, in the defender role, male defenders are subject to shorter attacks than female defenders. The differences in agonistic performance could not be attributed to differences in perceived resource value between the sexes or to differences in body or weapon size. There are clear differences in the agonistic behaviour of males and females, and this possibility should be incorporated into models of contest behaviour. In particular, evolutionarily stable strategies may be expected to vary with sex ratios.     

Lek structure and territoriality in the chryxus arctic butterfly,Oeneis chryxus (Satyridae)

Summary The territorial and mating systems of the satyrid butterfly Oeneis chryxus were studied for 3 years on individually marked populations in algonquin Provincial Park, Ontario. Males defended territories in a dry open forest clearing by means of chases and spiral flights with conspecifies. Defended areas did not contain food or oviposition sites required by females. Territories were not uniformly distributed over the study area; the highest density of territories was clustered over bare ground covered with sand and small stones rather than areas of vegetation. This pattern of distribution of territories occurred during each year of the study. The patterns of occupancy of territories showed a high degree of consistency from one day to the next, some males defending a given territory for as many as 11 consecutive days. Removal experiments showed that a surplus of non-territorial males did not exist. Vacant territories in the high-density area were absorbed by neighboring males; those in the low-density area remained empty. Male size was not correlated with territory position in the study are. Wing patterning (i.e. size of spots) differed between males occupying territories on sandy/stony versus vegetated areas. The territorial and mating systems of O. chryxus resemble lek polygyny, and are more similar to some vertebrate lek systems than to hilltopping or landmark territoriality.