Territorial intrusions and copulation behaviour in the great skua, Catharacta skua

Birds of many species intrude on to territories in order to obtain extra-pair copulations, and frequent within-pair copulations are thought to be a response to cuckoldry where mate guarding is not possible. Great skuas are colonial birds in which females are left alone in the breeding territory while males forage for the pair. Opportunities for cuckoldry are therefore numerous, and it could be predicted that sperm competition should be intense in this species. We tested several hypotheses that attempt to explain territorial intrusions by female great skuas. Few intrusions coincided with the main fertile period and extra-pair copulations were almost never solicited, and therefore the sperm competition hypothesis was rejected. Only 0.9% of the copulations observed (N=339) were extra-pair. Thus opportunity for cuckoldry is a very poor predictor of the intensity of sperm competition, in spite of the relevance given to this factor in the literature. Of three extra-pair copulations observed, two involved unpaired territorial males. This suggests that genetic benefits were not the aim of unfaithful females. All three were preceded by courtship feeding, while only 26% of within-pair copulations followed successful food begging. This statistically significant difference constitutes evidence for a trade of copulations for food in a monogamous bird. Evidence is presented supporting hypotheses that females intrude on to territories in order to induce males to give away some food, and to become familiarized with potential partners for future breeding seasons.Copyright 1997 The Association for the Study of Animal Behaviour1997The Association for the Study of Animal Behaviour     

Multiple open forms of ribose-binding protein trace the path of its conformational change

Spawning success of males and its correlates were investigated in a natural colony of whitebelly damselfish, A. leucogaster (Pomacentridae), to explore the criteria that females use in choosing mates. The mating success of individual males was variable, with some males acquiring as few as 5000 eggs and others as many as 450 000 eggs during a breeding season. Male spawning success was not correlated with body size, territory size, nest site parameters or parental care behaviour. Egg hatching success was not related to either male size or egg clutch size, and all males were capable of rearing eggs to hatching. The temporal sequence of choices by females indicated non-independent choice by females, such that males chosen by females on the first spawn of the day were also chosen by females that spawned later in the day. Field observations indicated that, in the absence of male courtship, females preferentially visited males that had eggs in their nest site. Males that had recently mated were preferred by females over those males with either late-stage eggs or no eggs in the nest. This female preference did not appear to be related to increased paternal care or egg clutch survival. Given that the mating system is promiscuous and non-resource based, and that there appears to be little difference among males in body size, females may be mating non-independently by mimicking the choice of other females. Copyright 1998 The Association for the Study of Animal Behaviour Copyright 1998 The Association for the Study of Animal Behaviour.     

Prey availability and selective foraging in shorebirds

Accurate measurements of prey availability are vital to our understanding of foraging behaviour, particularly prey selectivity. In the present study, observations of shorebirds foraging for fiddler crabs on intertidal mudflats demonstrate that prey availability depends both on the temporal variation in crab activity and on the crabs' responses to the presence of foraging shorebirds. Our results suggest that measurements of prey availability that do not specifically account for prey activity patterns and their responses to predators are neccessarily inaccurate. Furthermore, our results also show that tests for foraging selectivity are extremely sensitive to the way in which prey availability is measured and can even indicate active prey selectivity when more accurate measures of prey availability show predators to be non-selective. Because inaccurate measures of food resources greatly reduce our ability to detect food preferences, greater care must be taken to account for prey activity patterns and their responses to predators in measurements of prey availability. Copyright 1998 The Association for the Study of Animal Behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.     

The island posterior calf fasciocutaneous flap: an alternative to the gastrocnemius muscle for cover of knee and tibial defects

By repeatedly presenting an alien scent to territory-owning beavers, Castor canadensis, we tested two competing hypotheses about the function of scent marking: scent fence and scent matching. The scent-fence hypothesis predicts that territory owners should respond increasingly strongly over time towards a recurrent alien scent because of the ineffectiveness of previous responses. The scent-matching hypothesis predicts that the intensity of response should be the same or decrease because, without the presence of the intruding signaller coupled with the chemical signal, the presence of the scent itself does not advertise the ownership of a territory. The response level of resident beaver families was stable to strangers' anal gland secretions (AGSs) and decreased to strangers' castoreum during a period of 6 days. These results support the scent-matching hypothesis but not the scent-fence hypothesis. Copyright 1998 The Association for the Study of Animal Behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.     

Territorial behaviour in the Tasmanian native hen: group and individual performance

We studied the territorial behaviour of the Tasmanian native hen, Gallinula mortierii, a cooperatively breeding gallinule, for three breeding seasons at Maria Island National Park, Tasmania, Australia. Our analyses of 2738 territorial contests among 54 groups revealed that 87% of interactions were won by the group that initiated the interaction, 7% yielded no clear result, and only 6% were won by the opposing group. Asymmetries in the resource-holding potential of competing groups were not involved in deciding the outcomes of territorial contests. Our results suggest that asymmetries in payoff were involved, as in each of the 3 years groups with better quality territories were more likely to start interactions, and as a result, were more likely to win them. Only 6% of territorial contests escalated to involve physical contact and changes in territory boundaries rarely resulted from individual contests. The quality of a group's territory either changed very little between breeding seasons, or markedly increased or decreased. We conclude that major changes in territory quality occurred only under relatively rare conditions where the risks associated with escalated contests were reduced, or where the payoffs from a favourable result were improved. Males put more effort into the defence of the group territory than did females. The extra effort of males was directed more towards extragroup males than towards extragroup females. Females, in contrast, were equally aggressive towards both sexes. This suggests that the agonistic behaviour of male Tasmanian native hens may contain a mate-guarding component as well as functioning in the defence of territory. (c) 1998 The Association for the Study of Animal Behaviour.     

Two cooperatively social populations of the theridiid spider Anelosimus studiosus in a temperate region

The quality of an individual's odour can allow potential mates to discriminate against individuals of low social class, poor health status or unsuitable genotype. Competitive scent marking provides a further mechanism which could allow mates to discriminate between individuals of apparently high quality. The presence or absence of fresh countermarks from competitors within an owner's territory or area marked by a dominant animal provides a reliable indicator of the owner's ability to defend its territory or dominate competitors. This could be used by potential mates to discriminate between individuals advertising their apparently high competitive ability through their scent-marking behaviour and odour quality. We tested this by manipulating scent marks in the neighbouring territories of wild-caught male house mice, Mus domesticus. As predicted, oestrous females used scent marks to select males apparently able to defend exclusive territories over those unable to exclude intruders. Females were more strongly attracted to the odour of owners of exclusively marked territories and showed more sexually related behaviour when interacting with these males. Furthermore, while females preferred a territory containing a better protected nest site regardless of the owner's apparent competitive ability, they still used the presence or absence of intruder countermarks when selecting a potential mate. This suggests that females use scent marks as a reliable signal of the best-quality mate among neighbouring males independently of their nest location. Since assessment depends on both the territory holder's own marks and those of competitor males, countermarking is likely to be an important mechanism of competition for mates between neighbours. Copyright 1998 The Association for the Study of Animal Behaviour     

Yellowhammers get fatter in the presence of a predator

According to predictions from theoretical models, body mass and the daily foraging pattern of small birds in winter should be sensitive to predation risk. With increased risk of predation birds are expected to lower their body mass and shift their foraging to the afternoons. I studied the effect of predator presence on daily mass gain in captive yellowhammers, Emberiza citrinella. The birds were tested in unisex pairs, one bird being dominant over the other. In the first treatment the amount of food was restricted, and the daily pattern of mass gain was recorded. The second treatment was identical to the first, but, in addition, a stuffed sparrowhawk, Accipiter nisus, was displayed to the birds in the morning. Contrary to predictions from some theoretical models and other studies, the birds gained weight on days when the hawk was shown. Furthermore, the increased foraging happened mainly in the first half of the day, which also contradicts predictions from models. These results could be explained as a response to interrupted foraging, decreasing the probability of the birds acquiring the critical evening body mass. Alternatively, the birds increased their body reserves in preparation to leave a dangerous situation. Both dominance classes showed a similar response. The results suggest that the effects of predators on foraging interruptions should be included in models of daily routines of reserve accumulation. Copyright 1998 The Association for the Study of Animal Behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.     

Context determines the sex appeal of male zebra finch song

We explored the conditions under which playbacks of male zebra finch, Taeniopygia guttata, song induced reproduction in females. In a laboratory study, a rise in faecal oestrogen levels predicted egg laying. Song playbacks by themselves induced a decrease in oestrogen levels. There was an increase in oestrogen levels, followed by egg laying, when the song was broadcast from inside a male model positioned away from the nest. However, this effect occurred only when a second, silent male model was perched on the rim of the nest. If song was broadcast from inside the model perched on the nest, there was no increase in oestrogen levels. We conclude that tests of song efficacy in female songbirds must respect some contextual rules, which are likely to vary between species. Only then does it become possible to ascertain which sounds are most effective in inducing physiological changes leading to reproduction. Copyright 1998 The Association for the Study of Animal Behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.     

Proteins in urine scent marks of male house mice extend the longevity of olfactory signals

The binding of volatile semiochemicals to lipocalin proteins in many mammalian scent marks may provide a gradual release of volatile ligands, extending the life of airborne odour signals. We tested this by using menadione to displace semiochemical ligands from major urinary proteins (MUPs) in urine streaks obtained from adult male house mice, Mus domesticus, and assessed the responses of other males to these and to intact urine marks as they aged. Dominant male mice scent-mark their territories extensively with urine streaks; MUPs in these marks bind at least two semiochemically active molecules, 2-sec-butyl-4,5-dihydrothiazole (thiazole) and 2,3-dehydro-exo-brevicomin (brevicomin), associated with the males' aggressive status. Wild-caught males (N=24), housed in individual enclosures, were presented with two glass slides, behind mesh to prevent contact, on which 10 μl of both unfamiliar urine and 0.5 mg/ml menadione in ethanol had been streaked. On one slide the urine and menadione solution were mixed to displace ligands; on the other they were separate (intact urine). We carried out tests 0, 0.5, 1 or 24 h after deposition, and matched them to changes in the concentration of thiazole and brevicomin within the intact and displaced marks. Males were hesitant to approach intact urine up to 1 h old but, when ligands were displaced, or were reduced to low levels by natural evaporation from intact urine streaks aged 24 h, their approach was similar to that to water and to menadione controls. Ligands did not appear to cause any longer term avoidance and, after the first approach, investigation increased with the freshness of urine regardless of when the ligands were displaced. This is the first direct demonstration that proteins evince a slow release of olfactory signals from mammalian scent marks. The nature of their response suggests that, from a distance, mice may be unable to tell whether airborne signals emanate from scent marks or from the donor himself and we suggest that this may provide territory owners with a major advantage in defending their territories.Copyright 1998 The Association for the Study of Animal Behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.     

Female dispersion and the evolution of monogamy in the dik-dik

In facultatively monogamous mammals, females are thought to be too widely dispersed for males to defend more than one female range. We tested this hypothesis in a monogamous antelope, Kirk's dik-dik, Madoqua kirkii. Dik-dik territories were compared across three indices of quality to investigate whether males are monogamous because of constraints on the area, or resources, that they are capable of defending. Territories varied substantially in size and quality, with some containing up to five times the resources of others. Moreover, the territories of four temporarily polygynous males were not of higher quality than those of monogamous males. These results are inconsistent with the idea that dik-diks are facultatively monogamous: males can, and often do, defend enough resources and sufficient area to support two or more females. We investigated the relationship between resource dispersion and monogamy further by providing food to territories over a 3-month period. Although provisioned pairs obtained 20% of their daily requirements from the food, there was no change in the territory size of either sex. Female dispersion does not account for monogamy in the dik-dik; instead we argue that monogamy has evolved as a result of male mate guarding.Copyright 1997 The Association for the Study of Animal Behaviour     

Behavioural correlates of testosterone and seasonal changes of steroids in red-winged blackbirds

I studied the relationship between behaviour and plasma testosterone level (T) and the seasonal changes in T and plasma corticosterone levels (B) in male red-winged blackbirds, Agelaius phoeniceus. I measured T and B using radioimmunoassay, and on the day after taking blood, I observed each male's behaviour for 60 min. The time that males spent conspicuously perched and the number of songs were positively correlated with T, but the proportion of time spent conspicuously perched and the frequency of song were not correlated with T. The frequency of aggressive encounters, sexual chases, epaulet exposure when singing and flights within the territory were positively correlated with T, suggesting a direct role for circulating testosterone influencing male aggressive behaviour. Both T and B increased early in the breeding season, peaked when the first females were receptive, and decreased through the remainder of the breeding season. Late in the season, the presence of a receptive female caused males to have increased T. The peak in T when the first females were receptive, the positive correlation between aggression and T, and the response to a receptive female with increased T support predictions of the challenge hypothesis. T was positively correlated with B, suggesting a cost to the males of maintaining high T. When a receptive female was present on the male's territory, T was negatively correlated with date. Male red-winged blackbirds in Indiana may respond less to receptive females late in the season when benefits associated with protecting paternity and gaining extra-pair fertilizations decrease.Copyright 1998 The Association for the Study of Animal Behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.     

The changing trade-off between food finding and food stealing in juvenile oystercatchers

When juvenile oystercatchers, Haematopus ostralegus, first arrived on the wintering grounds in August and September, they regularly stole mussels, Mytilus edulis, from other, mainly older, oystercatchers. By October, however, juveniles stole far fewer mussels and found almost all their mussels independently for themselves on the mussel bed. Although stealing a mussel was always less profitable than taking a mussel from the mussel bed, a simple rate-maximizing optimality model showed that, in August and September, juveniles increased both their net and gross rates of energy intake by stealing because they were rather inefficient at foraging for themselves. By October, their greater efficiency at finding good quality mussels, combined with the increased resistance of potential victims to kleptoparasitic attacks, resulted in higher intake rates if juveniles stopped stealing mussels and took mussels only from the mussel bed. Copyright 1998 The Association for the Study of Animal Behaviour Copyright 1998 The Association for the Study of Animal Behaviour.     

Feeding interruptions, diurnal mass changes and daily routines of behaviour in the zebra finch

We investigated daily changes in body mass, fat reserves and crop contents, and diurnal organization of behaviour, in the zebra finch, Taeniopygia guttata, in relation to experimental manipulations of food availability. Diurnal mass change and the organization of foraging behaviour during the day were in general agreement with recent theoretical predictions. Foraging intensity, and hence rate of mass gain, was most rapid immediately after dawn and before dusk. The experimental birds did not alter either mean body mass or their diurnal mass trajectory after a period of 2 weeks when food was made unavailable for 2 h a day at unpredictable times. Instead, they changed their allocation of time to different activities during the day, decreasing the mean amount of time spent locomoting and increasing the mean amount of time spent inactive over the day. Thus, contrary to a number of recent studies on different species, zebra finches appear to respond to unpredictable interruptions in food supply by reducing energetically expensive activities rather than adjusting their levels of energy reserves. Copyright 1998 The Association for the Study of Animal Behaviour Copyright 1998 The Association for the Study of Animal Behaviour.     

Expression of a virus-derived cytokine, KSHV vIL-6, in HIV-seronegative Castleman''s disease

Flights from foraging flocks of sanderlings, Calidris alba were observed at Redcar, northeast England, in order to investigate how individual decisions to fly related to the behaviour of other flock members. Flights of sanderlings tended to be either of all birds in the flock on the ground, or of single birds or groups representing only a small proportion of the foraging flock. The latter accounted for the majority of movement events but after weighting by flock size the former accounted for the majority of bird movements. As flock size increased, the number of birds that flew without the whole flock taking flight increased. Thus, as flock size increased, it took more individual departures before the whole flock took flight. When flocks were disturbed by people or dogs, they were more cohesive, with more departures being of whole flocks.Copyright 1997 The Association for the Study of Animal BehaviourCopyright 1997The Association for the Study of Animal Behaviour.     

Parental investment, reproductive success and polygyny in the lapwing, Vanellus vanellus

We studied the consequences of monogamy and polygyny for male and female lapwings at a site in northern England between 1993 and 1995. Males and females differed in breeding behaviour, and thus the pattern of reproductive investment: males contributed less time than females to the care of their offspring and more time to mating behaviour. We argue that this has resulted from sexual selection. Reproductive behaviour was similar in monogamous and polygynous individuals of both sexes. Male mating success was related to territory size, with males on the largest territories gaining more females. Polygynous male lapwings reared on average between 58 and 100% more chicks each year than monogamous males because of fewer complete breeding failures; between-year return rates of males to the area were similar. This would result in a strong advantage in terms of lifetime reproductive success for polygynous male lapwings. The seasonal breeding success of polygynous females was marginally, but not significantly, lower than that of monogamous females. Between-year return rates of monogamous and polygynous females were similar. Copyright 1998 The Association for the Study of Animal Behaviour.     

The influence of tail autotomy on agonistic behaviour in a territorial salamander

Assessment of potentially asymmetrical characters (such as fighting ability and resident advantage) is often important in determining the outcome of agonistic interactions. Loss of body parts, a predator defence mechanism used by many animals, may lead to a reduction in fighting ability and may be easily assessed by competitors. We investigated the influence of tail loss on the expression of agonistic behaviour in the territorial red-backed salamander, Plethodon cinereus. Residents and intruders were matched for body size, and pairs were tested in all combinations of tailed or tailless residents with tailed or tailless intruders. Neither residents nor intruders altered their behaviour based on their own tail condition, but they did alter their behaviour based on the tail condition of their opponents. Intruders showed more aggression or less submission towards tailless residents than towards tailed residents. When contests were between residents and intruders of the same tail condition (both tailed or tailless), intruders were more aggressive towards residents when both were tailless than when both were tailed, indicating that tail loss does not directly hamper aggressive displays. In contests where the asymmetry between residents and intruders was small (based on tail condition and residency status), intruders showed more aggression and less submission than in contests where the asymmetry was large. Residents did not differ in their behaviour for most comparisons. Thus, for intruders, the tail condition of residents is an important determinant of agonistic behaviour displayed in territorial contests. For residents, factors other than tail condition (such as resource value) may be more likely to influence their behaviour. Copyright 1998 The Association for the Study of Animal Behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.     

Cooperative signalling between opponents in fish fights

Cichlids of the species Nannacara anomala employ several colour displays during fights which do not seem to signal either fighting ability or motivation. How should these colour displays be interpreted when winning is reliably predicted by weight asymmetries? Medial Line colour displays were associated with, and predicted, tail-beating, while Vertical Bar colour displays were associated with mouth-wrestling. I suggest that these colour displays are used to facilitate the transmission of assessment information within a fight, and that they are an example of cooperative signalling between opponents. The results support the idea that the structure of fights contains strong cooperative aspects.Copyright 1997 The Association for the Study of Animal Behaviour1997The Association for the Study of Animal Behaviour     

Song learning and vocal tradition in Nuttall's white-crowned sparrows

We investigated song sharing and dispersion of song types in the wild in a colour-marked population of the non-migratory Nuttall's white-crowned sparrow, Zonotrichia leucophrys nuttalli. The songs of fathers, their male progeny (sons), and the neighbours of the sons at recruitment sites were analysed spectrographically and compared qualitatively and quantitatively. To determine whether a son's song more closely matched that of his father or a neighbour at the site settled, we subjected frequency and temporal characteristics of songs within each father-son-neighbour triad to multivariate cluster analysis. The songs of 14 of 16 sons clustered with their neighbours' rather than their fathers' songs, confirming that song matching of neighbours is an integral component of territory settlement by juveniles. Principal components analysis of frequency and temporal measurements of song within a dialectal area show that songs group into neighbourhoods and are non-randomly distributed. Multivariate analysis suggests that sons may entrain on frequency and temporal characteristics of a neighbour's song without matching phrases or complex syllables. Implications for models of instructive versus selective learning are discussed. The timing of closure of the sensitive phase, the length of the silent interval between the sensory phase and plastic song stage, and the time to song crystallization remain open questions in song ontogeny. Copyright 1998 The Association for the Study of Animal Behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.     

Multiple mechanisms may affect birth sex ratio in domestic pigs

No abstract. Copyright 1998 The Association for the Study of Animal Behaviour.     

Mechanisms of sex-ratio adjustment in domestic swine: reply to James

No abstract. Copyright 1998 The Association for the Study of Animal Behaviour.