Effects of location in territory and distance from neighbours on the use of song repertoires by Carolina wrens

By capturing territorial Carolina wrens (Thryothorus ludovicianus) and housing them in portable cages on their territories, I could control each one's location in its territory and distance from neighbours in experiments on the use of song repertoires. Experiment 1 demonstrated that these wrens sang more songs in the centre of their territories than at the edge, but that they did not use more song types or different song types at the centre than at the edge. In experiment 2, in which I played tape-recorded songs at two distances from wrens caged in the centres of their territories, birds responded more strongly to songs at 25m, simulating an intruder, than to songs at 165 m, simulating a territorial neighbour. Birds also switched more frequently between song types and sang more song types per 100 songs in response to the nearer playback. Experiment 3 compared captive wrens 140 m, 80 m, 20 m, or 0 m apart on adjacent territories. As the distance between neighbours decreased, birds sang less, but also switched more frequently between song types, used more song types per 100 songs, and matched songs with neighbours more frequently. There were no differences in the kinds of song types sung at different distances from neighbours. A comparison of the results from experiments 1 and 3 confirms that the use of song repertoires is influenced by distance from conspecifics and not by location in the territory.     

Cooperation and conflict among dunnocks, Prunella modularis, in a variable mating system

The mating system of a population of 90 breeding dunnocks (or hedge sparrow, Prunella modularis) included monogamy, polygyny, polyandry and polygynandry. Monogamous males guarded females during their fertile period to prevent neighbouring males from copulating. The most intense guarding occurred where two (unrelated) males shared a territory. Here, the alpha male tried to prevent the beta male from copulating with the female. Beta males were seen to copulate in only half the cases. They were more likely to succeed when the alpha male found it difficult to guard the female closely because her range was large, the vegetation was dense or there were other females breeding synchronously on the same territory. Close guarding and chasing by males reduced the female's feeding rate and was correlated with unhatched eggs in the nest. Females attempted to escape the alpha male's attentions and actively encouraged the beta male to mate. Beta males only helped to feed the young if they copulated with the female. Nestlings fed by two males and a female got more food and weighed more than those fed by just one male and a female. Indirect evidence suggested that when beta males failed to copulate, they destroyed eggs or young chicks. Females laid larger clutches when two males mated with them as opposed to one, thus adapting their clutch size to the amount of parental care they expected. The results of natural removal experiments and matched comparisons of the same female in different mating systems support these conclusions. For females, selection favours cooperative polyandry, whereas for males if favours polygyny; the variable mating system may reflect the different outcomes of this sexual conflict.     

Female mate choice in the pupfish, Cyprinodon variegatus

Experimental evidence suggests that female Cyprinodon variegatus do not select between competitively inferior and superior males, but rather, deposit eggs randomly and in proportion to the size of the male's territory. The females do seem to have an habitat preference and probably influence the location where males compete for territorial space.     

Body size as a factor determining dominance in staged agonistic encounters between male brown anoles (Anolis sagrei)

The results of staged agonistic encounters between males indicate that body size is an important determinant of dominance in male brown anoles (Anolis sagrei). Larger males defended their perch sites more successfully than did smaller males, perched higher than smaller males, and were most often the first male to enter the other male's territory. Larger males also exhibited more challenge displays than did smaller males. Head-nods, a display given by subordinate individuals, were observed only in smaller males.     

Some geometric aspects of house sparrow flocks

We analysed 50 movie films of house sparrows (Passer domesticus) foraging on an experimental grid. The location and orientation of each bird was recorded; sex of the individual was determined in about two of every three cases. Results revealed the following: (1) Flock size exhibited a weak inverse relationship to ambient temperature, though aggression was rare in the experimental patch. (2) The frequency of males in foraging groups exceeded the frequency of males in the local population. (3) Within a given flock size, nearest neighbour distances did not differ significantly between male-male and male-female pairs. However, average nearest neighbour distance was inversely related to flock size. (4) Solitaries oriented away from safety and toward a source of disturbance. Orientation of an individual within a larger group was more variable than that of a solitary, and the orientation of nearest neighbours indicated a significant tendency to keep each other in view.     

Patterns of territory size and defence level in rural and urban tawny owl (Strix aluco) populations

The effect of different variables on the territory size and defence level of tawny owl ( Strix aluco L.) populations was studied in two habitats: mixed farmland (1976–1978) and town (1986–1988). Territory size was determined by plotting the position of territorial challenges (hoots) of males which were individually recognizable in most cases. No differences were found in size, defence, number of competitors and biomass between territories in farmland and in town, but territories in town were more fragmented and showed a larger nearest neighbour distance. Size of rural territories was adequately predicted both by some structural habitat features and food supplies, while size of urban territories was affected only by habitat structure and by years of territory occupancy. No effect on territory size by competing neighbours was found in either habitat, thus confirming that behavioural interactions between the owners of territories and potential settlers prevent a decrease of territory size.
Defence level of urban territories was affected by 'social'variables (nearest neighbour distance and years of occupancy), structural variables (percentage of poplar grove and diversity), and food supply, while in rural territories only structural variables (fragmentation or natural vegetation) and food supplies affected defence level.
This was due to different settlement tactics in the two areas, depending on the extent and quality of suitable habitat.     

Frogs Have Rules: Selective Attention Algorithms Regulate Chorusing in Physalaemus pustulosus (Leptodactylidae)

In various acoustic insects and frogs, females preferentially orient towards the leading of two or more males' advertisement signals that occur closely in time. Such preferences in receivers have apparently selected for timing mechanisms whereby male signallers actively refrain from calling immediately following the onset of a neighbour's call and thereby increase their production of leading calls. However, indiscriminate application of this inhibitory mechanism to all neighbours might severely reduce a male's calling rate, particularly in high density. Consequently, mechanisms of selective attention to only a subset of signalling neighbours are expected.
Female Túngara frogs ( Physalaemus pustulosus ) exhibit strong preferences for leading male calls, and males refrain from calling immediately following a neighbour. Four-loudspeaker playback experiments demonstrated that males do selectively apply this inhibitory mechanism to only a subset of close signalling neighbours. Selective attention is regulated by a combination of sliding threshold and fixed number rules: (i) Attend to the loudest (nearest) conspecific neighbour and those additional ones whose calls are within 6–8 dB of the loudest one; (ii) attend to only two neighbours in total when the calls are weak or the second one is much farther than the first; (iii) attend to three neighbours when the calls are loud or all neighbours are approximately equidistant. The means by which such plasticity may be achieved and its potential adaptiveness are discussed.     

Territorial intrusions by pikas (Ochotona princeps) as a function of occupant activity

Pikas (Ochotona princeps) defend individual territories on talus. We tested the short-term stability of pika territories and found that although animals may intrude on the territories of conspecifics at any time, they were much more likely to do so if the occupant was inactive. Pikas usually waited until occupants were unavailable to defend their territory before intruding, apparently to avoid detection and repulsion by the occupant. Intruder avoidance of active occupants was most pronounced among same-sex dyads and non-nearest neighbours. Instances of territory intrusion were most frequent among nearest neighbour heterosexual dyads. Most cases of intrusion appeared to be related to eventual relocation of territories, deterring settlement of unfamiliar conspecifics on nearby vacant territories, and/or increasing familiarity with nearest neighbours of the opposite sex.     

Responses to oral gland scents from different males in Columbian ground squirrels

Acrylic cubes carrying oral gland scents from strange, neighbouring, and resident male Columbian ground squirrels (Spermophilus columbianus), plus a cube with no scent, were presented near the home burrows of squirrels in the field. Adult males and adult females sniffed scented cubes longer than unscented cubes, sniffed cubes with scent from other (strange and neighbouring) males longer than cubes with the resident male's own scent, and sniffed cubes with scent from strange males longer than cubes with scent from neighbouring males. Males also scent-marked the cubes on a number of occasions; differences in the frequency with which the cubes were marked followed the same pattern. It thus appears that adult Columbian ground squirrels can recognize familiar as opposed to unfamiliar males, and may also be able to determine individual identity, through oral gland scent. An ability to distinguish familiar from unfamiliar males by scent could be advantageous in facilitating differential treatment of wandering strange males versus established neighbours.     

Interspecific assistance: fiddler crabs help heterospecific neighbours in territory defence

Theory predicts that territory owners will help established neighbours to repel intruders, when doing so is less costly than renegotiating boundaries with successful usurpers of neighbouring territories. Here, we show for the first time, to our knowledge, cooperative territory defence between heterospecific male neighbours in the fiddler crabs Uca elegans and Uca mjoebergi. We show experimentally that resident U. elegans were equally likely to help a smaller U. mjoebergi or U. elegans neighbour during simulated intrusions by intermediate sized U. elegans males (50% of cases for both). Helping was, however, significantly less likely to occur when the intruder was a U. mjoebergi male (only 15% of cases).     

When are neighbours ‘dear enemies’ and when are they not? The responses of territorial male variegated pupfish, Cyprinodon variegatus, to neighbours, strangers and heterospecifics

Dear enemy recognition reduces the costs of territorial defence in some species, but not others, when a neighbour is more threatening to a resident's fitness than an intruder. I asked whether dear enemy effects were fixed in a particular species, or if the reduced aggression between a resident and neighbour was disrupted by the presence of potential mating opportunities. Observing variegated pupfish, Cyprinodon variegatus, in the field and in the laboratory, I examined the effects of a female's presence in a male's territory on residents' aggressive responses to conspecific neighbours and strangers as well as heterospecific opponents. Although reduced aggression consistent with dear enemy recognition was seen between conspecific neighbours in the absence of females, the presence of a female in a male's territory instigated comparably greater aggression between the neighbours. No reduction in aggression was seen between pupfish males and heterospecific opponents. These findings suggest that dear enemy recognition may be a flexible, rather than a fixed, feature of the relationship of neighbouring conspecific males. Despite the disruption in dear enemy recognition caused by a female, residents in the laboratory faced with neighbours spent more time associating with the female than residents faced with strangers. This allowed the residents to secure as many spawns as did males who had been faced with no competitor. Residents faced with any other type of opponent had reduced reproductive success, suggesting that the dear enemy relationship between residents and neighbours is more complex than simply a reduction in aggression. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.       

Song Trait Similarity in Great Tits Varies with Social Structure

For many animals, long-range signalling is essential to maintain contact with conspecifics. In territorial species, individuals often have to balance signalling towards unfamiliar potential competitors (to solely broadcast territory ownership) with signalling towards familiar immediate neighbours (to also maintain so-called “dear enemy” relations). Hence, to understand how signals evolve due to these multilevel relationships, it is important to understand how general signal traits vary in relation to the overall social environment. For many territorial songbirds dawn is a key signalling period, with several neighbouring individuals singing simultaneously without immediate conflict. In this study we tested whether sharing a territory boundary, rather than spatial proximity, is related to similarity in dawn song traits between territorial great tits (Parus major) in a wild personality-typed population. We collected a large dataset of automatized dawn song recordings from 72 unique male great tits, during the fertile period of their mate, and compared specific song traits between neighbours and non-neighbours. We show here that both song rate and start time of dawn song were repeatable song traits. Moreover, neighbours were significantly more dissimilar in song rate compared to non-neighbours, while there was no effect of proximity on song rate similarity. Additionally, similarity in start time of dawn song was unrelated to sharing a territory boundary, but birds were significantly more similar in start time of dawn song when they were breeding in close proximity of each other. We suggest that the dissimilarity in dawn song rate between neighbours is either the result of neighbouring great tits actively avoiding similar song rates to possibly prevent interference, or a passive consequence of territory settlement preferences relative to the types of neighbours. Neighbourhood structuring is therefore likely to be a relevant selection pressure shaping variation in territorial birdsong.     

Pairing strategies and mate choice in female robins Erithacus rubecula

Female European robins (Erithacus rubecula) adopted three different pairing strategies. These were, in order of frequency: moving on to a male's territory; fusing her territory with that of a neighbouring male; and being joined by a male on her territory. Although females were not free to choose between strategies during a particular season, individuals frequently changed strategy between different years. Their behaviour suggested that they were exercising mate choice, although the scope for such choice was limited by a strong tendency for a female to pair in areas where she had lived previously (her ‘Familiar Areas’). Males with large territories were more likely to pair and tended to pair earlier than those defending small territories. Female pairing strategy was also correlated with the area of their own territory. Although it is considered likely that some unidentified confounding variable was responsible for at least part of these relationships, possible causal explanations involving territory size are discussed. These include mate choice and a random settlement model. However the simplest explanation for the advantage enjoyed by males on large territories is that their territories were more likely to overlap with females' Familiar Areas.     

Familiarity and flexible mating strategies of a solitary rodent, Dipodomys ingens

Because mating is a product of individual reproductive strategies that may vary with changing conditions, we predicted variable mating behaviour in an arid-adapted, territorial rodent, the giant kangaroo rat, Dipodomys ingens. We also predicted that familiarity would facilitate nonaggressive courtship and mating in this solitary rodent. Through direct observations in the field, we found that mating varied from exclusive to multiple partners. Where densities were low, and on nights when multiple females were in oestrus, each animal mated with one member of the opposite sex. In conditions where the operational sex ratio was skewed towards multiple males, males footdrummed and competed for females. Males were able to mate with one or two females in adjacent territories, and they successfully competed for these same females throughout the breeding season. Females that mated exclusively with one male had more pups emerge from the burrow compared with females that experienced male competition. Females allowed nearest neighbour males to enter their burrows, and they engaged in more nonaggressive contact with close neighbours than with other males. Paired encounters in the field showed less aggression towards neighbours than strangers. In laboratory tests, females were less aggressive towards and allowed more contact with familiar than unfamiliar males. These results show that D. ingens can alter mating strategies as conditions change. Familiarity is an important factor in nonaggressive interactions between males and females and may be important to mate preferences in females during reproduction. The less aggressive behaviour to neighbours than to strangers (‘dear enemy’ phenomenon) is consistent with other solitary animals that defend multipurpose territories. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Tests of spatial and temporal factors influencing extra-pair paternity in red-winged blackbirds

Extra-pair paternity (EPP) is a widespread and highly variable reproductive phenomenon in birds. We tested the effects of habitat, spatial factors, and timing of breeding on the occurrence of EPP in red-winged blackbirds (Agelaius phoeniceus). We used PCR-amplified microsatellites to assess the paternity of 1479 nestlings from 537 broods on 235 territories over four breeding seasons. Over 4 years, 40% of nestlings were extra-pair. At least 27% of actual sires were non-neighbours, suggesting that males or females interacted over longer distances than in other populations of red-winged blackbirds. The level of EPP was significantly clumped within broods and males but not within females across broods. EPP was negatively related to the area of a male's territory. The spatial proximity of a female's nest to the territory boundary had no effect on total EPP, but tended to increase the probability of an EPP by a nearby male. We found no influence on EPP of the type of habitat on the territory or the level of nesting activity nearby. The time in the season when a nest was started and the synchrony of breeding also had no significant effect on the level of EPP. The age of the male, the age of his neighbours, and the interaction between the two had no effect on total EPP. However, older males were less likely to have an offspring sired by a neighbour on their territory. Males with older neighbours were also less likely to have offspring sired by a neighbour, particularly if they were new territory owners. The high variability in who gained and lost paternity, and the limited impact of spatial and temporal factors influencing it, have some interesting implications for theories seeking to explain mating patterns.     

The effect of local dominance and reciprocal tolerance on feeding aggregations of ocellated antbirds

We studied ocellated antbirds (Phaenostictus mcleannani) to test the hypothesis that reciprocal tolerance between dominant individuals can favour feeding in aggregations. Mated pairs hold large non-exclusive feeding ranges, but roost and nest in a small portion of this range (‘roosting area’); adjacent roosting neighbours are unrelated. Ocellated antbirds congregate to feed on arthropods fleeing from nomadic swarms of army ants that move across the ranges of many pairs. We used playback experiments to simulate acoustic challenges, and results showed that males responded aggressively to other males only in their roosting areas. Responses to adjacent neighbours were less aggressive than to non-neighbours (i.e. the ‘dear enemy’ effect). Prey intake rates were higher when birds fed in their own roosting area or in that of adjacent neighbours compared with more distant sites. Males tolerated adjacent neighbours at swarm fronts where prey are most dense, but more distant neighbours were displaced. Despite small samples for some analyses, our results suggest that reciprocal tolerance between adjacent unrelated neighbours can ameliorate intraspecific competition within ephemeral feeding aggregations.     

Territorial responses of male blue tits, Cyanistes caeruleus , to UV-manipulated neighbours

Multiple phenotypic traits can affect the outcome of interactions among territorial animals. Individuals may use current and previously acquired information on phenotypic traits to assess the competitive ability of opponents and adjust the strength of their response depending on the threat the opponent poses. In birds, colourful plumage and song are widespread phenotypic traits. Recent work has shown that ultraviolet (UV) plumage reflectance may be used by males in assessing an opponent’s strength and by females in mate choice. In the present study, we investigated whether and how territorial male blue tits, Cyanistes caeruleus, use previously acquired information from UV reflectance of the crown feathers of neighbours to adjust their response to playback of song of these neighbours simulated to intrude their territory. We compared responses to neighbours with those to unfamiliar strangers with unknown plumage features. We found that subject males with UV-enhanced neighbours responded more strongly to these neighbours than to strangers, i.e. showed more flights, used songs without trill and tended to overlap more songs. Subject males with UV-reduced neighbours gave a lower or similar response to neighbours compared to strangers. This indicates that male blue tits combine previously acquired information about an intruder’s plumage with familiarity of its song, and that their response depends on the perceived quality of the neighbour. This study provides evidence that familiarity in combination with multiple signals of quality may influence territorial relations among neighbours.     

Interrelationships of dominance and territorial behaviour in the pupfish,Cyprinodon variegatus

Under laboratory conditions, Cyprinodon variegatus establishes either a dominance or territorial mating system. In the dominance system, one male usually has complete access to receptive females, while in the territorial system, several males have equal access. Four factors were observed to enhance a subdominant's ability to establish a territory and improve his reproductive success; increasing the number of conspecific intruders, increasing the available area, the presence of partial barrier, and the presence of a receptive female.     

Everybody Needs Good Neighbours: Coalition Formation Influences Floater Fight Choice

In territorial species, it is sometimes less costly to help a neighbour fight off an intruder than to re‐establish territory boundaries with a new, potentially stronger neighbour. In fiddler crabs, a male resident will only help his neighbour if he is larger than the intruder who, in turn, is larger than the challenged neighbour. Does this influence with whom a territory‐seeking male decides to fight? I show that territory‐seeking males appear to choose opponents based partly on the size of the resident’s nearest neighbour. By avoiding challenging resident males with larger neighbours, territory‐seeking males can reduce the likelihood of initiating a fight with a resident who might gain help from his neighbour that decreases the likelihood that the intruder will win the fight.     

The effect of neighbours and females on dawn and daytime singing behaviours by male chipping sparrows

Bird song may play an important role for communication among territorial neighbours, but the effect of neighbours on song use is still not well known. My previous field observations suggested that male chipping sparrows, Spizella passerina, use the dawn chorus for interactions among neighbouring males, and use day song for female attraction. To determine how these social factors may influence dawn and daytime singing behaviour, I conducted a series of experiments in which I removed the male neighbours or the female mate of territorial males during 1998-2000. Following removal of all neighbouring males, the solitary male either stopped or reduced his dawn chorus (N=9), but did not change his daytime singing behaviour. After one of the neighbouring males was returned to his territory, the focal male resumed and increased his dawn bout, accompanied with close-range countersinging. Following the removal of a territorial male's mate, the widowed male did not change the dawn chorus, but significantly increased his day song. This study thus revealed that, in chipping sparrows, the presence or absence of neighbouring males has a significant effect on the dawn chorus singing behaviour of territorial males. The presence or absence of a male's mate, in contrast, has a strong influence on a male's daytime singing behaviour. This study also supports the hypothesis that the dawn chorus and daytime song have different functions.