Social class,socially-induced loss,recruitment and breeding of red grouse

Summary Individually marked red grouse (Lagopus lagopus scoticus) could be distinguished as territorial or non-territorial from November till the next spring. Territorial birds survived well and many reared young, but numerous nonterritorial birds died or disappeared, and none paired up or reared young. Differences in spring sex ratio between years had already been set in the previous autumn's territorial stock. Many old birds which had been territorial in earlier seasons lost territories after an annual re-distribution of territory each October–November. Thereafter very few of them were seen alive on the area over the winter, and many were found dead, whereas nearly all old birds that kept their territories were still there in spring. The number of birds in spring and the fate of individuals over winter were predicted reasonably accurately from observations of behaviour in the previous November–December. Thus, differences of social class in autumn were critical to a bird's survival to breed in the next summer and to its chances of contesting for territory and breeding in subsequent years.     

Family-based winter territoriality in western bluebirds, Sialia mexicana: the structure and dynamics of winter groups

Winter residency is characteristic of the majority of cooperatively breeding birds, but the composition and dynamics of winter groups have been examined in relatively few. In 1996-1998, we examined winter territoriality in the western bluebird, a year-round resident that shows a limited degree of helping behaviour in central coastal California, U.S.A. In spring, most western bluebirds breed as socially monogamous pairs, but a small proportion of pairs (3-16%) have additional breeding-age males helping at the nest, usually assisting parents or brothers. We found that year-round residents commonly wintered in family groups that defended territories similar to those used in spring. Winter groups had an even sex ratio and formed early in the autumn, when hatch-year birds dispersed. More females than males left their natal groups to be replaced by an influx of immigrant hatch-year birds. Winter groups typically consisted of breeders and one or two sons from the prior breeding season along with one or more immigrant females. A second period of dispersal occurred in spring when winter groups broke up and most birds other than the breeding pair left the winter territory. When they bred, yearling males and females often bred with unrelated individuals from their winter groups. Sons were more likely to remain on the study area as yearlings when they wintered with both parents than when they wintered with just one parent. We suggest that young males stay the winter due to benefits of remaining in family groups on mistletoe-based winter territories. Subsequent localized dispersal of sons then leads to opportunistic kin-based interactions later in life. Copyright 2001 The Association for the Study of Animal Behaviour.     

Testosterone and autumn territorial behavior in male red grouse Lagopus lagopus scoticus

In many bird species, males exhibit territorial aggression outside the breeding season, when testosterone concentrations are low and may not regulate territorial behaviors. The hormonal regulation of aggression at this time of year has only been studied in passerine birds. Here, we investigated the role of testosterone in the regulation of aggression in a non-passerine bird, the red grouse Lagopus lagopus scoticus. Male red grouse are aggressive in early spring when breeding starts, in autumn when they establish territories, and sporadically through much of the winter. We first describe seasonal variations in plasma testosterone concentrations and in the size of males' sexual ornaments, their red combs, which relates to aggressiveness. Testosterone concentrations and comb size were correlated. Both increased in autumn to a peak in October, and then increased again in spring, to a greater peak in early April. Secondly, we experimentally investigated the effects of testosterone, and of an anti-androgen (flutamide) used in combination with an aromatase inhibitor (ATD), on autumn territorial behavior. Males were treated with either empty implants, as controls (C-males), testosterone implants (T-males), or with flutamide and ATD implants (FA-males). One month after implanting, both T- and FA-males had higher concentrations of testosterone than C-males. Comb size, aggressive call rate, and response to playbacks of territorial call all significantly increased in T-males. However, the increase in testosterone in FA-males did not increase comb size or aggressive behavior. In the following spring, after the content of implants was used, FA-males had significantly lower testosterone than C-males, and had a reduced seasonal increase in comb size. The results suggest that testosterone plays a significant role in regulating red grouse aggressive behavior in autumn. However, the observation that flutamide and ATD treatment did not reduce territorial behavior, suggests that estradiol may also be involved in the regulation of non-breeding aggression.     

THE SOCIAL BEHAVIOUR OF THE WHITE WAGTAIL MOTACILLA ALBA ALBA WINTERING IN ISRAEL

White Wagtails Motacilla alba wintering in Israel are partly territorial, mostly around human habitations, and partly live in flocks around temporary food sources. Individual birds may spend part of the season (or the day) in the territory and the other part with a flock. Experiments with artificial distribution of food, in a natural habitat, brought about a change from flocking to territorial behaviour. Preliminary observations suggest that in the natural situation the pattern of food distribution may be the proximate factor which regulates the birds' behaviour, by determining whether they have to fight for their food. Pairs are formed on many territories, and may last for long or short periods. Pair formation is initiated by females, who when seeking food appease the territorial males and are able to stay with them on their territories. Females also manifest territorial behaviour. Although pairing in winter territories is similar, in the behaviour involved, to sexual pairing, it is very unlikely that winter pairing continues into, or influences, pairing for breeding. It is suggested that the function of winter pair formation is that it allows two birds to exploit one territory, and that the main advantage is to the female which is the subordinate bird of the pair. This kind of pair-formation may be analogous to non-breeding group territories reported in some other birds.     

Winter territoriality and its implications for the breeding ecology of White-throated Dippers Cinclus cinclus

Capsule: Pairs of White-throated Dippers Cinclus cinclus which defended winter territories bred earlier than non-territorial individuals, but there was no difference in reproductive success.

Aims: The effect of winter territoriality on breeding ecology has rarely been studied in resident birds. We carried out a preliminary investigation of whether winter territorial behaviour and territory size affect the timing of reproduction, breeding territory size and reproductive success in a riverine bird, the White-throated Dipper.

Methods: We monitored an individually marked population of White-throated Dippers in the UK. Wintering individuals were classified as either territorial or ‘floaters’ according to their patterns of occurrence and behaviour, and their nesting attempts were closely monitored in the subsequent months. Winter and breeding territory sizes were measured by gently ‘pushing’ birds along the river and recording the point at which they turned back.

Results: All birds defending winter territories did so in pairs, but some individuals changed partners before breeding. Territorial pairs that were together throughout the study laid eggs significantly earlier than pairs containing floaters and those comprising territorial birds that changed partners. However, there were no significant differences in clutch size, nestling mass or the number of chicks fledged. There was no relationship between winter territory length and lay date or any measure of reproductive success, although sample sizes were small. Winter territories were found to be significantly shorter than breeding territories.

Conclusion: Winter territoriality may be advantageous because breeding earlier increases the likelihood that pairs will raise a second brood, but further study is needed. Territories are shorter in winter as altitudinal migrants from upland streams increase population density on rivers, but this may also reflect seasonal changes in nutritional and energetic demands.     

The wintering of migrant Dunnocks Prunella modularis in two Mediterranean habitats after fire

Migrant Dunnocks Prunella modularis are common winter visitors to the Mediterranean area. In a burnt forest, Dunnocks were present from October to April. They occupied definite individual home ranges of 0.18 ha on average (n = 83), at a density of 30 birds/10 ha in the fifth winter and 18 birds/10 ha in the sixth winter following a fire. Birds used visible posts and uttered ‘tseep’ calls often. Counter-calling was particularly frequent in the winter with the highest population density. In a burnt maquis, Dunnocks showed return rates (15%) and median recapture distances (62.5 m) not significantly different from those of territorial wintering Robins Erithacus rubecula. We suggest that Dunnock ranges can be regarded as ‘feeding territories’, promoted by an abundant, predictible and renewable food supply of rock-rose Cistus spp. seeds, characteristic of early Mediterranean successions. Ownership might be proclaimed by ‘tseep’ calls, recalling the behaviour of female Dunnocks during prebreeding territorial conflicts. Further research using colour-ringed birds may confirm Dunnock winter territoriality, exceptional for a migrant and mostly granivorous passerine.     

Territorial behavior of both sexes in the water striderMetrocoris histrio (Hemiptera: Gerridae) during the mating season

Territorial behavior of overwintered individuals of Metrocoris histriowas observed in an upstream area. Adults of both sexes held territories, but male territories were larger than those of females. Severe competition occurred among males for territories which give them access to receptive females. The effects of male body length and midleg length on establishment of territories were not significant. The effect of female midleg length on activity of females entering preferred foraging sites was equally not significant. Instead, territorial behavior increased with male age and males stayed longer at prime sites. Females of intermediate age were likely to occupy prime sites. Females had longer territory residence time than males. The sexes were dimorphic with respect to midleg length, and dimorphism in M. histriomay be related to a difference in life history, in that sexual selection may be relaxed due to asynchronous adult emergence patterns.     

Winter territoriality of Mute Swans Cygnus olor

There was considerable variation in winter territoriality among Mute Swan pairs around Oxford, though most pairs maintained their territories for at least half the winter. Variation was associated with temperature and with territory quality pairs left their territories most commonly during the coldest months of the winter, and pairs on good territories, with abundant vegetation and accessible pasture. stayed longer on territory than those on poor territories with sparse vegetation and little pasture. As in other species, exclusive territorial defence was maintained at intermediate levels of resource availability, and flocks were able to settle on a few particularly good territories. Circumstantial evidence suggested that winter territories may be maintained not only to provide an adequate food supply for the owners but also to ensure access to a brerding site by preventing other individuals from taking over the area.     

Long‐term phenological shifts and intra‐specific differences in migratory change in the willow warbler Phylloscopus trochilus

Climate change can influence many aspects of avian phenology and especially migratory shifts and changes in breeding onset receive much research interest in this context. However, changes in these different life‐cycle events in birds are often investigated separately and by means of ringing records of mixed populations. In this long‐term study on the willow warbler Phylloscopus trochilus, we investigated timing of spring and autumn migration in conjunction with timing of breeding. We made distinction among individuals with regard to age, sex, juvenile origin and migratory phase. The data set comprised 22‐yr of ringing records and two temporally separated data sets of egg‐laying dates and arrival of the breeding population close to the ringing site. The results reveal an overall advancement consistent in most, but not all, phenological events. During spring migration, early and median passage of males and females became earlier by between 4.4 to 6.3 d and median egg‐laying dates became earlier by 5 d. Male arrival advanced more, which may lead to an increase in the degree of protandry in the future. Among breeding individuals, only female arrival advanced in timing. In autumn, adults and locally hatched juvenile females did not advanced median passage, but locally hatched juvenile males appeared 4.2 d earlier. Migrating juvenile males and females advanced passage both in early and median migratory phase by between 8.4 to 10.1 d. The dissimilarities in the response between birds of different age, sex and migratory phase emphasize that environmental change may elicit intra‐specific selection pressures. The overall consistency of the phenological change in spring, autumn and egg‐laying, coupled with the unchanged number of days between median spring and autumn migration in adults, indicate that the breeding area residence has advanced seasonally but remained temporally constant.     

Temporal shifts in the pair-bond dynamics of New Zealand robins (

Winter is a challenging time for temperate insectivorous songbirds, due to colder temperatures, reduced prey activity and shorter diurnal foraging times. For species that are non-migratory, territorial and monogamous, winter conditions may result in within-pair competition. However, little is known about how monogamous pairs coexist on their winter territories. We investigated temporal patterns in male?female interactions of the New Zealand robin (Petroica australis to better understand mechanisms of coexistence during winter. Previous work has shown that male robins are physically dominant over females and maintain priority access to food year-round. We quantified female behaviour throughout the 2008 non-breeding season to better understand how females coexist with physically dominant males on winter territories. Results showed that pairs rarely forage in close proximity in autumn and winter, suggesting females avoid males at this time of year. Males and females begin to spend more time foraging together as winter turns to spring. During this winter?spring transitional period, females steal large amounts of food hoarded by males. These results indicate that male and female New Zealand robins use different behavioural mechanisms to coexist on their winter territories. While males are dominant physically, females show a seasonally variable strategy where they avoid males in autumn and winter, and then steal male-made caches from early spring until the onset of inter-pair cooperation and the breeding season.     

Territorial status and survival in red grouse Lagopus lagopus scoticus: hope for the doomed surplus?

A previous study of survival in territorial and non-territorial red grouse Lagopus lagopus scoticus conducted between 1957 and 1967 found that territorial status in the autumn pre-determined over-winter survival. A very high proportion of territorial birds survived and virtually all non-territorial birds died or emigrated. We tested the hypothesis that over-winter survival was dependent on territorial status within four grouse populations in Scotland between 1986 and 1993. In contrast to the previous study, 66% of non-territorial birds survived over winter compared to approximately 70% of territorial birds. There was no significant effect of territorial status on the survival estimates. Moreover, some of the birds considered to be non-territorial during autumn went on to successfully raise a brood. We suggest that on our study sites, territory ownership in autumn did not greatly influence over-winter survival, and territorial behaviour did not determine breeding density as previously supposed. We postulate differences with other studies may reflect variations in scale and predation pressure.     

Family-based territoriality vs flocking in wintering common cranes Grus grus

We describe winter territoriality in common cranes, Grus grus , a long-lived migrant species with long-lasting pair bonds and parental care extending throughout the winter. Cranes are territorial in the breeding season, and usually gregarious during migration and wintering. Only 2% of the families present in our study area were territorial, all other families foraged in flocks with immatures and adult pairs. Territorial pairs defended the same winter territory year after year, but only when they had offspring. They were gregarious otherwise. The average breeding success measured throughout several years was higher in territorial pairs. Winter territories were small (0.7 km² on average), but included a higher diversity of habitats than the areas visited by gregarious birds (11.7 km² on average). Adults of territorial families showed longer vigilance times, and lower food intake rates than did adults in flocks, which were compensated with a longer time spent foraging per day. The accumulated daily food intake did not differ between adults in flocks and in families. We suggest that winter territoriality is a facultative strategy, conditioned by parental experience and habitat availability.     

The utilization of heather moorland by territorial Red Grouse Lagopus lagopus scoticus

Although the breeding density of Red Grouse Lagopus lagopus scoticus is related to the density of burned patches of heather Calluna vulgaris , territories appear to offer surplus food, and other factors, especially the risk of predation, may be important. In order to determine how Red Grouse make use of heather moorland, an individually marked population was studied on a moor in northeast Scotland from 1992 to 1994. Territorial birds preferred stands of tall heather during the daytime in autumn and winter, but in spring they increased their use of heterogeneous heather/grass mixtures. Furthermore, in spring, females showed a very strong affinity for edges between heather stands of different ages, where they had easy access to nutritious young heather and good cover. When near the vegetation edge, territorial birds spent most time on the side of the edge providing greater cover. Broods occupied ranges in areas with rather more old heather and with more edge than was available over the entire study site. Within their ranges though, broods clearly preferred heterogeneous areas during the first month after hatching, and showed a further preference for vegetation edge. Their behaviour changed during the second month after hatching, when they utilized tall heather to a greater extent, but edge remained important.     

Possible relationships between morphology,territory quality,and skin color of American Kestrels1

ABSTRACT Carotenoid‐based coloration of skin and plumage has been found to be correlated with individual quality in many species of birds during the breeding season. However, less is known about the possible role of these signals during the nonbreeding season, particularly among nonpasserines that defend winter territories. American Kestrels (Falco sparverius) are sexually dimorphic raptors that defend winter territories and possess carotenoid‐based morphological features known to be correlated with individual quality. Much is known about winter territory use and habitat segregation by male and female kestrels, but possible relationships among morphological features, individual quality, and habitat quality have not been examined. Our objective was to examine possible relationships between morphology, territory quality, and skin color of American Kestrels. Male kestrels had brighter skin than females, and the skin color of male kestrels was positively correlated with size (wing chord and tail length) and territorial quality (hunting territories with less canopy and more grass cover). No such relationships were found for female kestrels. Skin color appears to be an honest indicator of quality for male American Kestrels and may serve both intersexual (territory acquisition) and intersexual (mate choice) functions during the breeding and nonbreeding seasons.     

Turnover, age and sex ratios of kestrels (Falco tinnunculus) in south Scotland

This paper examines the composition and turnover of a population of European kestrels Falco tinnunculus L. in the Southern Uplands of Scotland. Turnover was generally high, with most birds staying in the area for only one summer or winter. The rate at which birds arrived or left was highest in autumn and spring, and was independent of population size. During winter, breeding males were more likely to stay on their territories than females, and males that did so were more likely to retain their partners the following year. Breeding success was associated with a greater likelihood of return the following year, except in first-year females which were unlikely to return anyway. The sex ratio was biased toward males in winter, especially in poor vole years when there were also few first-year birds present. The study confirmed the high rate of turnover reported elsewhere in small raptors, and further indicated that the composition and turnover of the population were related to food supply and seasonal patterns of migration.     

Testosterone is correlated with courtship but not aggression in the tropical buff-banded rail,Gallirallus philippensis

Past studies into the roles of testosterone in birds have focused on species that occur in temperate regions. In such species, plasma testosterone levels are high during the establishment of territories in the spring and are associated with increased aggression. In contrast to most temperate species, tropical birds frequently defend territories year-round, during which time territoriality often occurs in a nonsexual context. The few studies that have been carried out on tropical birds show lower levels of circulating testosterone than occur in their temperate counterparts. In some year-round territorial tropical species, testosterone and aggression are dissociated, while in other species testosterone still plays a role in regulating aggression. This study examined the relationship between aggression and plasma testosterone levels in a year-round territorial, subtropical population of the buff-banded rail with characteristics typical of tropical species. Peak testosterone levels were substantially lower than those found in temperate species. Males displayed a seasonal peak in plasma testosterone level when their partners were most likely to be fertile. At other times, testosterone levels were mostly undetectable, despite year-round territoriality. We found that T levels increased with courtship behavior but showed no relationship with aggression, supporting the hypothesis that dissociation between testosterone and territoriality may be widespread among tropical avian taxa.     

Strained energy budget of winter floaters in the Willow Tit as indicated by ptilochronology

Most Willow Tits Parus montanus live within flock territories during the non-breeding season, but many juveniles roam around or switch between several flocks. Such floaters behave submissively and spend time watching other flock members in addition to scanning for predators. During September–December 1989–2000, I studied individually colour-ringed Willow Tits living in a subalpine forest area in central Norway. When among flock members, the floaters foraged less in favourable parts of trees and were more vigilant than the lowest-ranked juvenile flock members. The nutritional condition of the birds was assessed by examining the growth bar widths of their tail feathers. Birds in better condition have wider growth bars than those in poorer condition. The daily growth bars of induced feathers of male and female floaters, laid down under winter conditions, were significantly narrower than those of the lowest ranked flock members. These results suggest that juvenile floater Willow Tits maintain a poorer nutritional status than low-ranking juvenile flock members during winter.     

Seasonal change in Red-backed Shrike Lanius collurio territory quality - the role of nest predation

This paper examines the relationships between territory preference, reproductive performance and nest predation in a Red-backed Shrike Lanius collurio population breeding in rapidly changing farmland. Territory preference was measured by (a) the order of laying dates in territories within years and (b) territory occupation frequency over a five-year period. Seven environmental variables were tested but only the density of Sloe (Blackthorn) Prunus spinosa in territories was correlated positively with territory occupation frequency. Within years, Red-backed Shrikes started to breed earlier in territories with more Sloe, but reproductive performance was not related consistently to territory differences. A high proportion of Sloe-rich territories was vacant each year. Thus, the observed negative association between settlement order and amount of Sloe may reflect a seasonal change in habitat preference. Early breeders prefer Sloe-rich territories whereas late birds prefer Sloe-poor territories. Supporting evidence is that the proportion of occupied territories that was Sloe-rich was higher for early than for average or late first breeding attempts. Moreover, in contrast to early in the season, the probability of a replacement clutch following nest failure was not associated with the amount of Sloe, but increased with increasing densities of Juniper Juniperus communis. Late in the nesting season, pairs breeding in Sloe-rich territories suffered higher nest predation than those breeding in Sloe-poor territories. In early nesting attempts, however, the relationship was reversed. Nest predation is probably an important selective agent on habitat selection since it was the principal factor explaining variation in the numbers of fledglings produced among territories. Thus, the Red-backed Shrike may shift territorial preferences adaptively as the season progresses to match within-season changes in predation pressure in different types of territories.     

Survival in a long-lived territorial migrant: effects of life-history traits and ecological conditions in wintering and breeding areas

Despite its key role in population dynamics and evolutionary ecology, little is known about factors shaping survival in long‐lived territorial species. Here, we assessed several hypotheses that might explain variability in survival in a migratory Spanish population of a long‐lived territorial species, the Egyptian vulture Neophron percnopterus, using a 16‐year monitoring period and live‐encounter histories of 835 individually marked birds. Cormack‐Jolly‐Seber capture–recapture models showed no evidence for effects of sex or nestling body condition on survival. However, the normalized difference vegetation index (NDVI; an indicator of primary productivity) of natal territories had positive effects on juvenile survival, indicating that environmental conditions experienced early in life can determine survival prospects. Survival increased with age (0.73±0.02 in the first 2 years to 0.78±0.03 in years 3 and 4) to later decrease when birds were five years old (0.60±0.05), the age at which they acquire the adult plumage, abandon the communal lifestyle of juveniles, and may look for a breeding territory. At older ages, survival was higher for non‐breeding (0.75±0.02) and breeding adults (0.83±0.02). Among the latter, birds that recruited into better territories had higher survival prospects. Age‐specific variation in survival in this species may be related to behavioural changes linked to dispersal and recruitment into the breeding population, while survival prospects of adult birds strongly depend on breeding territory selection. These results suggest a tradeoff between recruiting soon, and thus reducing mortality costs of a long and extensive dispersal period, and trying to recruit into a good quality territory. Finally, annual survival rates for birds of all age classes were positively related with the NDVI in their African wintering grounds. Although this relationship was probably mediated by food availability, further research is needed to properly identify the limiting factors that are affecting trans‐Saharan migrants, especially in light of global climate change.     

Territory strategy of the migratory Brown Shrike Lanius cristatus

Brown Shrikes Lanius cristatus are both transients and winter residents in Taiwan. Large numbers of birds crowd into the available habitat during the transient period, and the cost of maintaining territories changes greatly, depending on the number of shrikes present. This study examined how wintering Brown Shrikes adapt to the changing level of competition in territory establishment. About 24% of autumn transients arrived before any wintering birds, while 73% of wintering birds arrived during autumn transient time. The frequency of aggression recorded was significantly related to the number of birds seen along the transect line that day. Among ringed birds that returned during transient time, 80% returned directly to their previous territories, but some waited for the departure of transient birds before establishing territories. Territory compression occurred at high population densities. Departure from and arrival at the study area were frequent during the winter, but no birds moved territories within the study area. On average, 25% of ringed wintering shrikes returned to the study site in the subsequent years, while no transients returned. This low site tenacity may be. in part, a result of a high persecution rate along the migration route.