Do territorial and non‐breeding Australian Magpies Gymnorhina tibicen influence the local movements of rural birds in New Zealand?

Australian Magpies Gymnorhina tibicen sometimes attack and kill other birds in New Zealand. Here we assess how Australian Magpies influence the local distribution of other birds in New Zealand and identify the members of an Australian Magpie population that display the most agonistic acts. We conducted regular observations on six territorial breeding groups and three non‐breeding flocks of Australian Magpies for 1 year to determine (1) if other birds avoid flying or foraging close to Australian Magpies, (2) the proportion of passing birds that are attacked and (3) which social subunits of the Australian Magpie population are most aggressive. In comparison with adjacent Magpie‐free control areas, significantly fewer birds of a range of species (e.g. Common Blackbird Turdus merula, Skylark Alauda arvensis, Yellowhammer Emberiza citrinella) foraged close (i.e. ≤ 50 m) to both territorial breeding groups and non‐breeding Australian Magpie flocks; fewer birds were also recorded flying near (i.e. ≤ 50 m) territorial breeding groups but not non‐breeding flocks. Excluding Australasian Harriers (Circus approximans: see below), only 8% of birds recorded within 50 m of territorial breeding groups were observed being attacked. Attacks were most frequent when numerous birds occurred near Australian Magpies and species recorded in the highest frequencies were generally attacked most. Territorial breeding groups attacked 39% of passing Australasian Harriers. All attacks consisted of the victim being swooped at or chased; no physical contact was ever observed. Both adult male and female breeding Australian Magpies were seen attacking other birds; juveniles in breeding groups sometimes supported adults but never initiated attacks. Australian Magpies in non‐breeding flocks were not seen attacking other birds, except Australasian Harriers (attacked in 17% of appearances). Our results suggest that some birds avoid foraging and/or flying close to Australian Magpies because they are sometimes chased by breeding adults of both sexes; however, the proportion of passing birds actually attacked was small. The numerous published observations of Australian Magpie attacks are apparently biased heavily towards sensational events that are rare. Possible reasons why Australian Magpies attack other birds are discussed.     

The role of male territories in the mating system of the Yellow-shouldered Widowbird Euplectes macrourus

The territorial system and breeding biology of the Yellow-shouldered Widowbird Euplectes macrourus (Ploceidae) was investigated in western Kenya. Yellow-shouldered Widowbirds had a resource-defence polygynous mating system: males defended large (mean = 0.95 ha) territories and built the coarse framing for the nests in tall grass. Males had up to five females nesting per territory. Females provided nearly all parental care except for a territorial male seen feeding a fledgling: the first observation of paternal care in the wild for this genus. There was considerable variation in territory size, but the cause of this variation remains unknown: territory size was not related to potential indicators of territory quality, such as grass height and abundance, did not relate to male morphology (mass, size and ornament size) or territorial behaviour (boundary displays and singing) and did not affect female preferences. Although resources (territories and nests) were defended by the males, observations that males frequently fed outside their territories and formed communal roosts during the breeding season suggest that this species represents a transitional stage between typical resource-defence polygyny and lek breeding.     

A SOCIO-ECOLOGY OF MAGPIES PICA PICA

The behaviour and ecology of unmarked Magpies Pica pica living in the Cotswolds were investigated over 18 months. The spacing of nests was significantly more regular than random and breeding pairs defended territories throughout the year. In winter pairs occasionally left territories to feed with non-breeding birds which flocked throughout the year.
Magpies in flocks had a higher averiage feeding rate than birds alone or in pairs. Aggression rates in flocks increased as birds fed closer together and as food patches became richer.
Agonistic interactions between Crows Corvus corone and Magpies were common; Crows frequently dominated Magpies in disputes over food, and Crows also ate Magpie eggs and young. Single Magpies were most often chased by Crows while Magpies in flocks were able to feed longer in the presence of Crows.
Differences in fledgling success among breeding pairs were related to the location of nests; Magpie pairs breeding near Crow nests suffered higher rates of agonistic encounters with Crows and also produced fewer fledglings than did Magpies breeding at greater distances from Crow nests. Magpie nests were located nearer human habitations than were Crow nests.     

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.     

Prior residence, territory quality and life-history strategies in juvenile Atlantic salmon (Salmo salar L.)

Three groups of juvenile salmon were introduced sequentially into an artificial stream to investigate the effects of prior residence on behaviour and territory choice. Almost half of the first group obtained and defended distinct territories, the other half being constrained to an area approximately the size of one large territory. All of the fish in the subsequent groups, bar one, were also constrained to the same site. Since the fish were of similar size, prior residence alone seemed to influence which individuals obtained territories. However, within the first group, the fish that obtained territories were larger and more aggressive. The territorial fish did not appear to choose the most profitable territories, although they had the greatest opportunity to do so. Since juvenile salmon emerge from their gravel nests fairly synchronously, a time constraint on site sampling is hypothesised: there may be a risk in taking time to sample sites, since these same sites may become occupied with conspecifics. However, fish with territories fed at faster rates than non-territorial fish, possibly because of reduced competition for prey items. Consequently, fish from the first group (containing most of the territorial fish) grew faster than the other two groups. Moreover, most of the territory holders, but only one of the non-territorial fish, reached the threshold size that increases their probability of smolting the following year. This suggests that ability to obtain a defensible territory, primarily through prior residence, influences the age at which juvenile salmon can migrate to sea.     

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 use of supplementary foods by Australian magpies Gymnorhina tibicen: Implications for wildlife feeding in suburban environments

Abstract The provision of foods to wild birds is an extremely common practice among householders throughout the Western world. Nonetheless, concerns over potential impacts of the practice are currently being raised, including the possibility that some species may become reliant on human‐provided food. We compared the foraging and breeding ecology of pairs of fed and unfed Australian magpies Gymnorhina tibicen living in suburban environments in Brisbane, Queensland, Australia. Detailed behavioural observations of both foraging and chick provisioning were made for males and females of both groups throughout the breeding season. Natural foods dominated the diets of both fed and unfed magpies, making up 76% and 92% of all items consumed respectively. During the morning, fed magpies obtained fewer food items during ground foraging than did unfed birds, apparently because they visited suburban feeding stations more often during the early part of the day. At other times, the amount of food items obtained during foraging was similar for both fed and unfed birds. Magpies utilizing suburban feeding stations started all breeding activities significantly earlier than unfed magpies, except during the fledgling phase. Both fed and unfed magpie parents provisioned their chicks predominantly with natural food. Magpies were not reliant or dependent on supplementary food provided by wildlife feeders at any time during the breeding season. Although many magpies did utilize suburban feeding stations extensively, they continued to forage for and provision their chicks with natural food.     

Evidence for long-distance dispersal in a sedentary passerine, Gymnorhina tibicen (Artamidae)

Australian magpies (Gymnorhina tibicen) are group-living birds found across much of mainland Australia. Adults commonly remain in a breeding territory until death. Young of the year either remain on the natal (birth) site or are forced by their parents to disperse. Observational studies in south-eastern Australia suggest that most dispersing juveniles settle within 7 km of their natal territory. Therefore, despite potential for considerable gene flow (via flight), social organization predisposes magpies towards local population structuring. In this study, we measured genetic variation at both nuclear (allozyme) and mitochondrial loci and found evidence of substantial gene flow over very large distances (up to 1599 km). Thus, some juvenile magpies may disperse much greater distances than was previously thought. For mtDNA, geographic and genetic distance were strongly correlated, consistent with a pattern of isolation by distance. Therefore, although female gene flow is substantial it is apparently geographically restricted over large distances, in approximately a stepping-stone fashion. We conclude that a strong relationship between gene flow and geographic distance can develop even over large distances if populations have experienced no major historical disturbances to gene flow.     

The effect of food supplementation on the reproductive success of Black-billed Magpies Pica pica

The effect of supplementary food on the reproductive success of Black-billed Magpies Pica pica was studied in an urban habitat in the city of Edmonton, Alberta, Canada. We provided supplementary food (pelleted dog food) regularly from August 1986 to June 1988 in a feeder situated outside all known magpie territories. Magpies using the supplementary food showed (i) an advancement of 7 days in the initiation of laying, (ii) no increase in clutch-size, (iii) higher survival of nest contents (eggs and young) during a spring snow storm, (iv) an increase in rate of nestling weight gain, and (v) an increase in fledging success. Supplementary food did not prompt the fed pairs to select nest sites nearer the feeder in 1988 than in 1987. The feeder did not affect the breeding density because the nearest neighbour distances of fed magpies did not differ from unfed magpies in either year, nor did they differ between the two years for either fed or unfed magpies. This study provided evidence that magpies frequently renest even if their initial nests fail after the eggs hatch—a practice that has been reported to be absent or rare in other studies. Although brood reduction was equally frequent in the nests of food-supplemented and control pairs, the actual number of nestlings disappearing from the former (1.56 per nest) was significantly smaller than that from the latter (2.60 per nest), suggesting that the availability of food plays a major role in magpie reproductive success.     

Breeding of the bellbird on the Poor Knights Islands,New Zealand

Abstract

The breeding of the bellbird (Anthornis melanura) was studied intensively over three seasons on Aorangi Island, Poor Knights Islands. Adult males defended territories all year but ventured beyond them to exploit localised food resources and to obtain water; some adults defended the same territory for at least 5 years. Adult females shared a territory with a male only during the breeding season. At other times of the year adult females were joined by juveniles and immatures and formed feeding flocks. The breeding season extended from late September to late December. A few nests were built on the ground but most were in dense vegetation, usually near the canopy. Peak egg-laying extended from mid-October to mid-November and only one clutch of two to four eggs was laid. Nest building and incubation were completed by the female alone but both parents fed nestlings. Fledglings stayed in the vicinity of the nest for several days, and were fed by both parents. Incubation and nestling periods were about 15 and 19 days respectively. Comparisons are made with the breeding biology of bellbirds and other native passerines on mainland New Zealand, and the importance of the predator-free enviomment of the Poor Knights Islands is stressed.     

Harem structure and female territoriality in the dwarf hawkfish <Emphasis Type="Italic">Cirrhitichthys falco</Emphasis> (Cirrhitidae)

We investigated the territoriality and the spatial and mating relationships of the haremic hawkfish, Cirrhitichthys falco, on a reef off Kuchierabu-jima Island in southern Japan. Each individual maintained a territorial home range which was defended against same-sex conspecifics at the boundary of the home range. The territory of each male encompassed the territories of 2–3 females, allowing the male to completely monopolize mating opportunities with those females. Based on our observations, we classified the harem type of C. falco as a territorial female type. Large juveniles maintained independent home ranges outside the female territories. In contrast, small juveniles were allowed to cohabit within the territory of an adult female. Stomach contents analysis revealed that the smallest size class of C. falco fed primarily on copepods. In contrast, all other size classes fed primarily on decapods. Together, these results suggest that female territoriality plays an important role in defending food resources.     

Social organization in the pheasant (Phasianus colchicus): harem formation, mate selection and the role of mate guarding

Harem formation and mate selection were studied in the pheasant in order to determine the advantages of territorial harem defence polygyny to the two sexes. We investigated the factors affecting harem size and the advantage to a female in remaining with one territorial male during breeding.
Female group size declined during late March and early April as females moved from large overlapping ranges into smaller, more widely dispersed breeding ranges. The proportion of female groups accompanied by males increased during this period.
Some males had a disproportionate share of females. Settled females were monogamous but, because a female's nest was generally outside the male's territory, her home range was larger than his territory.
Harem members were usually from the same winter group. Harem size was not related to territory quality in terms of food supply or nesting cover. Females were loyal to one male in more than one year even if his territory position changed. Older, territory-owning males had more females, both adult and immature, than males with newly-established territories. Harem size was not correlated with territory size.
We conclude that the mating system of the pheasant is based on mate guarding which protects females not only from the risk of predation or injury, but also from excessive energy expenditure incurred through being chased by other males. When escorted by a territorial male, females spent three times as much time feeding, one-fifth as much time running, and one-tenth as much time alert, as they did when not guarded.     

The social ecology of the Pyrenean desman (Galemys pyrenaicus) (Insectivora: Talpidae), as revealed by radiotelemetry

The social organization, spatial utilization and movement patterns of free-ranging desmans ( Galemys pyrenaicus G.) were studied by radiotracking individuals in the French Pyrenees. During the period of study, May to July, single adult male and female desmans formed a pair and defended their shared territory from neighbouring conspecifics by vigilance and scent-marking behaviour. Other adults did not form pair bonds and remained solitary. In such pairs, the territory of the adult male was always larger than, and completely enclosed, that of the paired female. Neighbouring pairs occupied contiguous territories of similar length. The territories of neighbouring males overlapped to a small extent, but the territory of one male and that of a neighbouring female were never seen to overlap. Juveniles were observed to utilize the territory of resident pairs, prior to their dispersal. Paired adults and juveniles exploited their ranges on a regular daily basis, whilst those of solitary adults, being larger than those of the former, were utilized on a 48–hour basis. Paired males appeared to invest more time in defending the borders of their shared territory than did females which, in contrast, spent most of their time within the centre of the territory. Despite some degree of territorial overlap between neighbouring conspecifics, their mutual avoidance, achieved through a system of temporal range utilization, resulted in few agonistic encounters.     

Responses in spatial organization to manipulations of the food resource in polygynandrous alpine accentors

The alpine accentorPrunella collaris is a territorial and polygynandrous mating species. The breeding unit is a group consisting of about seven members who share a group territory which contains all the resources necessary for living. Each female holds a small territory around her own nest within the larger group territory. Supplemental food in the form of millet seed was provided for a total of 23 group territories to test the hypothesis that the abundance and distribution of food influences the size of a female's territory and her dispersion pattern which, in turn, determine the mating system. Both males and females had regular access to the feeders, but females and groups provided with feeders did not have smaller territories than those without feeders. No groups were divided into two by offering extra food. Provision of extra food resulted in an increase of group size but did not alter the sex ratio. All members, including new settlers, were sexually active and the sexual relationships among the members of a group were still multiple matings. Food addition did not prompt the fed females to choose nest sites nearer the feeder. Nest sites were restricted to particular rocky slopes and non-vegetated areas which were invariably free from snow each year due to the effects of strong winds and the topography. Females showed a nest-site fidelity for successive years. The reason why supplemental food had no effect on the territory size or dispersion of female alpine accentors is discussed.     

The Behavioral Ecology of Libellula luctuosa (Burmeister) (Anisoptera: Libellulidae): I. Temporal Changes in the Population Density and the Effects on Male Territorial Behavior

A single population of a common pond dragonfly, Libellula luctuosa, was studied at a site where the density of males increased dramatically during the breeding season. Early in the summer one active male was found on each territory on the pond. Satellite males were only occasionally found on the territories. Later in the season the number of males per territory increased so that two or more males simultaneously defended on many of the territories, and several satellite males occupied each of the territories. The number and rate of female visitations per day did not change over the summer. These factors resulted in a change in the operational sex ratio with variations in male density. Male behavior was also altered with increasing population density. As male density increased, males were less likely to be seen perching on their territories and more likely to be seen performing aggressive acts such as chasing nearby territorial males and chasing intruders. At high male density, the duration of territorial behaviors was shorter than at low male density. Thus, the percent of a time budget spent in any one activity did not change despite the change in number of males present. Male activity in L. luctuosa is not strictly determined by the opportunity for aggression. Costs of aggression associated with territoriality are minimized by maintaining flexible territorial behaviors.     

Variation of territory size and defense behavior in breeding pairs of the endemic Lake Tanganyika cichlid fish Variabilichromis moorii

Lake Tanganyika harbors the ecologically, morphologically, and behaviorally most diverse species flock of cichlid fishes. It is comprised by substrate breeding and mouthbrooding species, most of which live in littoral habitats. Species communities are characterized by complex behavioral and trophic interactions, resulting in a dense pattern of partially overlapping territories, depending on the degree of ecological distinctness. We studied territorial behavior of breeding pairs in a substrate breeding species, with respect to territory size and defense behavior. The study species Variabilichromis moorii belongs to the tribe Lamprologini, the most species rich tribe of cichlids in Lake Tanganyika. Our study shows that breeding pairs of V. moorii can have highly complex territories, in which both parents hold separate sub-territories which are shifted slightly according to the movements of the fry, but the outer borders are conjointly defended. The size of the total defended territory varied from <1 to almost 4 m², averaging at about 2 m². Depending on presence of competitors or fry-predators evoking agonistic interactions, the territory size varied quite substantially over the day. Attack rates and size of the defended area decreased with water depth. Agonistic behavior was observed toward heterospecifics as well as conspecifics, with heterospecific attacks mostly concerning territorial neighbors and potential fry-predators in about equal frequencies. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Guest editors: T. Wilke, R. V?in?l? & F. Riedel Patterns and Processes of Speciation in Ancient Lakes: Proceedings of the Fourth Symposium on Speciation in Ancient Lakes, Berlin, Germany, September 4–8, 2006     

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.     

Territory quality affects the relative importance of habitat heterogeneity and interference competition in a long‐lived territorial songbird

Density‐dependent reproduction is commonly explained by either the habitat heterogeneity (HHH) or individual adjustment (IAH) hypothesis. Under the HHH, high quality territories are assumed to be occupied first. At higher density, occupation of low‐quality territories increases due to lower availability of high‐quality territories, which reduces mean reproductive success. Alternatively, the IAH assumes that increased competition at higher densities reduces reproductive success in all territories. For birds of prey, HHH plays an important role in territorial species, and IAH in socially breeding species. To test the generality of this hypothesis, we studied the mechanism behind density dependence in raven Corvus corax, a long‐lived passerine bird, using long‐term population data from a large number of territories. Population density decreased reproduction, which was explained by increased usage of low quality territories at higher density, supporting the HHH. Density reduced reproduction in low quality territories, but not in high and intermediate quality territories. We additionally compared the explanatory power of different models describing brood size, representing IAH, HHH, or a combination of both. The best model represented a combination of both hypotheses, in which the effect of density depended on territory quality. Our conclusion that both IAH and HHH are supported can be explained by the biology of ravens, where territorial adults not only experience interference competition with other territorial adults, but also with social groups of juveniles and floaters. We conclude that the relative importance of IAH and HHH may depend on variation in territory quality and social structure.     

Seasonal variation of territory size with the Little Owl (Athene noctua)

Summary In a population of Little Owls (Athene noctua) at the Lower Rhine (FRG), the factors were studied that influence the size and shape of territories of this non-migrating, all-year territorial owl species. These studies were carried out between September 1984 and June 1987. The birds were individually marked with transmitter packages so that their locomotional movements could be monitored. Using a standard protable stimulus (a dummy with an attached loudspeaker) the territorial boundaries of 19 male Little Owls were surveilled during the different seasons of the year. Male Little Owls defended their territories in all seasons, but distinct seasonal changes of intraspecific aggressiveness and territory size were observed. The seasonal variations of territory size followed a common pattern in all studied Little Owls, in spite of individual differences in the size of the defended areas. The largest territories were defended in March/April during the courtship season . In May/June all male Little Owls reduced the size of their territories . This corresponds to the breeding and nestling period. The yearly minimum of territory size was reached in the summer months July and August when the fledglings were still fed in the parental territory. When the first-year birds started to disperse in autumn (September/October), the size of the defended areas of the adult tenants again increased . In winter (November to February), a further increase in territory size was observed for periods of warm weather . Male Little Owls, however, were less aggressive during winter than in the following courtship season. On cold winter days with a ground cover of snow territorial aggressiveness ceased. Pastures and meadows offered a continuous food supply almost throughout the year. In some seasons they were overproportionally represented in the territories of Little Owls compared to the general surroundings. In reaction to changing accessibility of food, the hunting ranges within the home ranges were shifted much faster than the boundaries of the defended territories. The significance of various factors contributing to the variability of territory size in Little Owls are discussed (e.g. availability of food, seasonal and individual differences in aggressiveness, experience in the occupied area, population density).