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.     

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.     

Male Iberian rock lizards may reduce the costs of fighting by scent matching of the resource holders

Animals should adopt strategies to minimize the costs of intraspecific aggressive interactions. For example, individuals should be able to identify resource holders in advance and avoid fighting with them because residents are generally more likely than intruders escalate aggression. It has been suggested that scent marks function mainly to allow competitor assessment by conveying the costs of entering a scent-marked area. Individuals may identify territory owners by comparing the scent of substrate marks with the scent of any conspecific they encounter nearby, assessing whether these two scents match or not, a mechanism known as scent matching. Here, we examined the response of male Iberolacerta cyreni lizards to areas scent-marked by other males and the potential role of scent matching in agonistic interactions. We designed a laboratory experiment where we allowed a male to explore the scent-marked substrate of another male, and then we immediately staged agonistic encounters in a nearby clean neutral area with either the male that had produced the scent marks (matching treatment) or with a different non-matching individual male. The higher chemosensory exploratory rates of substrate scent marks in comparison to clean substrates suggested that males detected and spent more time exploring scent marks to obtain information on the donor male. Moreover, this information was later used to decide the fighting strategy. Intruding males delayed time until the first agonistic interaction, reduced the intensity of fights and the number of aggressive interactions, and won less interactions with males which scent matched that of scent marks (because they would be considered as the territory owners) than with other non-matching individuals. Our results show that male I. cyreni lizards use scent matching as a mechanism to assess the ownership status of other males, which could contribute to modulate intrasexual aggression, reducing costs of agonistic interactions.     

Intraspecific nest parasitism in the swallow Hirundo rustica: the importance of neighbors

Summary Intraspecific nest parasitism in the swallow Hirundo rustica involves several parasite and anti-parasite tactics. Neighboring swallows breed asynchronously, perhaps because neighboring nests with overlapping egg-laying periods have higher frequency of intraspecific nest parasitims than neighboring nests with no such overlap. Since nest guarding prevents nest parasitism, the effect of nest guarding was removed by putting up old swallow nests. Such experimental nests were parasitized by swallows more often when the nests contained eggs, than when empty, and when the nests were far away from, rather than close to, an active neighboring nest. Close experimental nests with eggs were, however, parasitized less frequently if asynchronous (non-overlapping egg-laying periods), rather than synchronous, to active neighboring nests. Aggressive anti-parasite behavior of nest owners was determined from responses towards female intruders. Responses were more often aggressive before and especially during egg laying than during incubation. Nest owners also behaved aggressively more often when the intruder approached close to as compared to at some distance from their nest. Focal pairs behaved aggressively more often towards closely approaching intruders during the incubation period than towards distantly approaching intruders during the egg-laying period. Swallows reduce the frequency of intraspecific nest parasitism by nesting asynchronously with and close to a neighboring nest. Aggression by neighbors may reduce the success of potential parasitic swallows.     

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.     

Territory ownership and familiarity status affect how much male root voles (Microtus oeconomus) invest in territory defence

Neighbour–stranger discrimination occurs when individuals respond with more aggression to strangers than to territorial neighbours—a phenomenon termed the “dear enemy phenomenon” (DEP). We investigated the DEP with male and female root voles (Microtus oeconomus Pallas 1776) using field dyadic arena tests conducted in enclosures where we could test for the effects of familiarity (familiar versus stranger), ownership (resident versus intruder status) and resource-holding potential (body mass) on territorial behaviours. The results showed that males put more effort into territorial defence than females, and males could discriminate between neighbours and strangers. In males, aggressiveness was influenced by a significant two-way interaction between treatment and ownership. Male residents were more aggressive towards stranger intruders than towards neighbour intruders, while male intruders were less aggressive towards stranger residents than towards neighbour residents. In females, neither treatment nor ownership status had a significant effect on aggressiveness. Familiar males performed more social behaviours but less non-social behaviours than stranger males. Furthermore, there was a clear dominance hierarchy between residents and intruders in stranger dyads, with the male territory holders dominating the intruder in pairwise interactions. To our knowledge, these results demonstrate for the first time DEP in a small mammal with a known pedigree and present the first evidence for “prior resident advantage” in voles. We argue that both ownership status and familiarity status affect how much an individual invests in territory defence. The benefits of neighbour–stranger discrimination for male root voles and the absence of neighbour–stranger discrimination in female root voles are discussed.     

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     

Alternative mating strategies in the water strider Gerris elongatus (Heteroptera,Gerridae)

Summary Males of the water strider Gerris elongatus established territories which included copulation and oviposition sites (small pieces of fallen bamboo). Males were aggressive and competition for territory and females was observed frequently. Male midlegs were more developed than female midlegs and were used as weapons. Reproductive behaviour changed as the breeding season advanced. Early in the season immature females were attracted by male surface wave courtship signals, then copulated white floating on the water surface without ovipositing (type 1). In midseason, males established territories, produced calling signals and attracted females which copulated and oviposited there with male postcopulatory guarding (type 2). In late season, many females oviposited without postcopulatory guarding on pondweed mats near fallen bamboo. Non-territorial males waiter for the arrival of these females and copulated without courtship, but mating success was low (type 3). These alternative mating strategies appeared to depend on differences in male size. Larger males were superior to smaller males in many ways (establishing territory, fighting, mating etc.). The largest males defended territories and had higher mating success than small non-territorial males. Medium sized males used all three strategies according to the number of empty territories and seasonal femald distribution.     

Chemical rival recognition decreases aggression levels in male Iberian wall lizards, <Emphasis Type="Italic">Podarcis hispanica</Emphasis>

In an agonistic interaction, the assessment of the probable outcome of future encounters with the same individuals may be the best way of decreasing costs of fighting, but this may only be accomplished if animals are able to recognize individual conspecifics. We staged encounters between male lizards, Podarcis hispanica, to examine whether odoriferous cues are involved in rival recognition during agonistic interactions. We experimentally manipulated the odour of intruding males, creating familiar males with their own odour or bearing odours of unfamiliar males, and unfamiliar males bearing unfamiliar odours or odours of familiar males. The results showed that when familiar males were impregnated with scents from unfamiliar males, they elicited an aggressive response by resident unmanipulated males similar to that observed for a new unfamiliar male with unfamiliar odour. This suggests that resident males were unable to recognize familiar males when their own scents were removed. In contrast, responding males were less aggressive towards familiar males impregnated with their own odour and towards unfamiliar males impregnated with scents of familiar males, suggesting that when two males have already interacted, their scents become familiar for both males, and that the detection in successive encounters of the familiar scent suffices to reduce the aggressive response of territorial males. Therefore, recognition mechanisms based on chemical cues during agonistic encounters may contribute to reducing the intensity and the costs of fighting in P. hispanica and may play an important role in the organization of their social system.     

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.     

Female aggressive response and hormonal correlates—an intrusion experiment in a free-living passerine

The optimal mating system is rarely the same for males and females—whereas males usually benefit from attracting additional females to the territory, this could incur costs for the resident female. Females should therefore prevent prospecting females from settling on the territory. We studied the male and female behavioral and hormonal responses to simulated female territorial intrusions in free-living bluethroats during the pre-laying period. In the study population, polygyny occurs with potential fitness costs for the resident female. We recorded different aspects of aggressive behavior before and after presentation of a live female decoy and playback of female song. These behaviors were compared with a set of intrusions using a male decoy. At the end of a trial, the birds were captured, and blood samples were analyzed for androstenedione, testosterone, estradiol and corticosterone. During the pre-intrusion period, none of the females were observed. Females generally responded strongly to the female decoy by increased flight rate, vocalizations, and by conspicuous perching. Nearly half of the males displayed to the female decoy but never while the resident female was present. We suggest that resident female aggression in bluethroats prevents courtship by her mate and signals her mating status to the female intruder. Female aggression could therefore prevent additional females to settle on the territory and shape the mating system. Females that responded with song had higher levels of estradiol. These findings suggest that estradiol may support aggression in breeding female birds.     

Feeding and reproductive behaviour and their effect on catchability of the spanner crab Ranina ranina

The behaviour of 16 adult (8 male and 8 female) spanner crabs (Ranina ranina), collected off southern Queensland, Australia, was monitored continuously in captivity by closed-circuit television for fifteen months from September 1982. Spanner crabs spent most of the time buried in the substrate, emerging mainly only when food became available. They remained emerged for twice as long on feeding days as on days without food. Females responded significantly faster than males to the presence of food. Response was slowest in October and November. There was no correlation between temperature and response time. The average feeding time was 2.0 min (SE=0.12), with no significant difference between males and females. Crabs without food were aggressive towards crabs with food, which sometimes led to fighting and wounding. These interactions also could result in food being transferred from one individual to another. When males interacted, food was transferred more often than when either females, or males and females interacted. Around moulting, male crabs did not feed for 52 d (SE=9.0) and females for 22 d (SE=2.2). This habit would reduce the frequency with which newly moulted crabs are caught in the baited tangle nets used by commercial fishermen. In mating interactions, copulation was always initiated by the males. Males dug up other crabs but, apparently unable to distinguish the sex of these individuals, attempted copulation with either sex. The majority of copulations occurred between midday and midnight and in the period August to December. The frequency of copulations with a female increased 10 d before she extruded eggs, after which it dropped to zero for the following 41 to 50 d. Eggs were carried for 39 to 44 d in the period September to November. The females remained emerged for long periods before extruding their eggs, but the period shortened immediately afterwards. This behaviour would lead to low catchability of ovigerous females. Because females respond more rapidly than males to a food stimulus, they may be more catchable in baited nets than are males. It is concluded that seasonal changes in the behaviour of spanner crabs could affect the number caught by baited tangle nets and may influence the sex ratio in catches.     

The smell of desperadoes? Beavers distinguish between dominant and subordinate intruders

Olfactory signals can contain information, such as age and social status, and play a vital role in competitor assessment. In many territorial species, subordinates must leave their natal colony to obtain their own territory and mate. These individuals could be aggressive opponents in agonistic encounters, as they will have little to lose (the desperado effect). In this study, we tested the hypothesis that dominance and age are coded in the anal gland secretion (AGS) of the monogamous and highly territorial Eurasian beaver (Castor fiber), and if this information is used by conspecifics to evaluate the potential threat posed by an intruder. Territorial intrusions were simulated by presenting residents with a two-way choice test of AGS from an unknown male territory owner (i.e., dominant) and his son (i.e., subordinate; either 1 or ≥2 years old). Residents spent more time investigating AGS from subordinates than their fathers and responded more aggressively to subordinates than their fathers when subordinates were ≥2 years old. Chemical analyses gas chromatography and multivariate data analysis supported our behavioral findings and revealed differences between chemical profiles of territory owners and subordinates, as well as between the subordinates in different age classes. This study reveals that information about age and social status is coded in AGS of beavers and that this information is used to determine the level of an eventual future response to the signaler.     

How do weaponless male fiddler crabs avoid aggression?

Mimicry of females enables weaker males in many species to avoid intrasexual aggression. In fiddler crabs (Uca annulipes), males use their major claw in aggressive interactions to acquire and defend a territory. Males that have autotomised their major claw will be disadvantaged in fighting, but might use their temporary resemblance to females to avoid costly aggressive encounters with other males. We investigated whether: (1) courting males mistake clawless male fiddler crabs for females; (2) clawless males are able to acquire, defend and retain territories as successfully as intact males; and (3) clawless males are more cautious than intact males. Clawless and intact males differed in burrow acquisition methods and fighting behaviour, but were equally successful at acquiring and retaining burrows. While courting males treated clawless males as female, we found no evidence that clawless males mimic the behaviour of females, or that they exploit the advantage of their mistaken identity. Clawless males further appear to avoid male aggression by altering their territorial strategies to minimise the potential for conflict.     

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.     

Changes of movement patterns from early dispersal to settlement

Moving and spatial learning are two intertwined processes: (a) changes in movement behavior determine the learning of the spatial environment, and (b) information plays a crucial role in several animal decision-making processes like movement decisions. A useful way to explore the interactions between movement decisions and learning of the spatial environment is by comparing individual behaviors during the different phases of natal dispersal (when individuals move across more or less unknown habitats) with movements and choices of breeders (who repeatedly move within fixed home ranges), that is, by comparing behaviors between individuals who are still acquiring information vs. individuals with a more complete knowledge of their surroundings. When analyzing movement patterns of eagle owls, Bubo bubo, belonging to three status classes (floaters wandering across unknown environments, floaters already settled in temporary settlement areas, and territory owners with a well-established home range), we found that: (1) wandering individuals move faster than when established in a more stable or fixed settlement area, traveling larger and straighter paths with longer move steps; and (2) when floaters settle in a permanent area, then they show movement behavior similar to territory owners. Thus, movement patterns show a transition from exploratory strategies, when animals have incomplete environmental information, to a more familiar way to exploit their activity areas as they get to know the environment better.     

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     

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.     

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.