Breeding territory fidelity in a partial migrant, the American dipper Cinclus mexicanus

American dipper Cinclus mexicanus populations are frequently composed of resident individuals that occupy permanent territories year round and migratory individuals that overwinter with residents but migrate to breeding territories on higher elevation creeks each spring. Between 1999 and 2004 we examined how migratory strategy (resident/migratory) and sex differences influence breeding territory fidelity of American dippers occupying the Chilliwack River watershed, British Columbia, Canada. Counter to expectation we found that the migratory strategy of American dippers did not influence whether birds breeding in one year were found on their former breeding territory in the next. Migratory strategy also did not affect the probability that known surviving dippers occupied the same breeding territory in the following year. Males and females were equally likely to be found on their former territory in the following year (females 43%, males 41%) and known survivors had similar levels of breeding territory fidelity (females 74%, males 68%). However, breeding territory fidelity of males and females varied in response to different factors. Surviving female dippers were more likely to be found on their former breeding territory in the subsequent year following a successful breeding attempt than an unsuccessful breeding attempt. Prior reproductive performance did not influence whether surviving male dippers were found on their former breeding territory. Male dippers were more likely to be found on their former territory and, if they survived, have higher breeding territory fidelity when their mate also returned to that same territory. Mate retention also influenced whether females were found on their former territory in the following year but had no effect on the breeding territory fidelity of known survivors. We argue that sex‐specific dispersal decision rules in American dippers are driven by sex differences in the predictability of breeding performance between years and sex differences in how mate retention influences subsequent reproductive success.     

Turnover and dispersal in a Peregrine Falco peregrinus population

In south Scotland, most Peregrines returned to the same territories to breed in successive years, though a few females changed territory from one year to the next.
Annual mortality among breeding birds was at most 9% among females (or 11% in both sexes combined). There may have been considerable annual variation, however, and excluding one exceptional year out of five reduced the estimate for females to 7%. These estimates are maxima, but are still considerably lower than those obtained from ring recoveries of dead birds reported by members of the public.
Among trapped birds, four males first bred at age two years, one at three and another at four or five; two females first bred at one year, 13 at two years old and one at three. Five other females which were seen to be in first-year plumage but were not trapped, also laid eggs, and 12 other such paired females held territory but did not lay. Only one paired male held territory in first-year plumage.
In their movements between natal and breeding territories, some females moved further than males, with median distances of 83 and 58 km respectively. In addition, of birds trapped breeding in the study area, a greater proportion of the males than of the females had been born locally, despite an equal sex ratio among fledglings; this was also consistent with a greater dispersal of females. In general, Peregrines made much longer movements in their first year of life than subsequently. Movements were in any direction.     

Turnover and recruitment in a tundra population of Peregrine Falcons Falco peregrinus

The population dynamics of tundra-nesting Peregrine Falcons Falco peregrinus tundrius was studied over 7 years on a 450 km² study area in the Keewatin District of Canada's Northwest Territories. Peregrines showed strong fidelity to nest sites; none of 25 males ringed changed territories, while five of 38 females ringed were recorded changing territories. Such changes usually occurred after nesting failure. Annual turnover of territorial adults was estimated to be 22% (15% for males and 26% for females). Annual mortality of adults was estimated to be 17% (15% for males and 19% for females). If we assumed that territory vacancies, in addition to mate replacements, were indicative of mortality, then maximum annual mortality was estimated at 24% for each sex. Territories were held only by adult Peregrines. The oldest male on territory was at least 7 years old, the youngest was 2. The oldest female on territory was at least 7 years old, the youngest was 3. Territories were held by individuals of each sex for at least 6 years. One pair remained together for at least 4 years. Less than 4% of all young Peregrines produced on the study area in the first 5 years of the study were recruited into the breeding population. More male than female young were recruited despite an even sex ratio among nestlings. Peregrines did poorly in their first breeding attempts. The single young female recruited into the study population dispersed more than three times the median dispersal distance of six recruited males, suggesting that other females probably dispersed beyond the boundaries of the study area.     

POPULATION DYNAMICS OF THE FLIGHTLESS CORMORANT NANNOPTERUM HARRISI

A small population of Flightless Cormorants was followed from 1970 to 1975 inclusive. The birds were extremely sedentary, most never moving more than 2 km from where hatched. Many birds bred several times within a year, almost always with different mates. After successful breeding the mean interval to the next attempt among females was significantly shorter than among males, probably because the male continued to attend the juvenile for longer than did the female. There was an annual peak of nesting in April to November, when sea temperatures were lowest; some nesting occurred in other months but these nests were less successful. About 73% of juveniles survived at least three months after going to sea. Adult females had a significantly higher rate of annual survival (91%) than did males (82%). The mean annual survival of both sexes combined over a 13 year period was 87%. The mean age of first breeding was about 30 months for both males and females. In 1972 breeding success (0·14 young fledged per pair) was much lower than in other years (0·60 young per pair), a lower proportion of juveniles survived, no birds bred for the first time and probably many fewer pairs nested. Adult survival was not affected. This reduced breeding output was associated with an influx of anomalously warm sea water to the area (El Niño). The availability of food is probably both the ultimate and the proximate factor controlling the timing of breeding.     

Patterns of territory settlement and consequences for breeding success in the Northern Wheatear Oenanthe oenanthe

We examined the pattern of territory settlement and its consequences for breeding success in the Northern Wheatear Oenanthe oenanthe on Bardsey Island, Wales, during the breeding seasons of 1991-93. Males returned earlier than females, and older males returned earlier than first-year breeders. Although their boundaries shifted between years, the general location of territories was consistent during the three-year study. There was a high degree of fidelity to area and territory between years for both sexes. The order of territory settlement, from which a territory rank was calculated, was highly consistent for males between years irrespective of individual settlement patterns and territory fidelity. Patterns of territory settlement were less predictable for females, although there was a significant correlation between the mean territory ranks of paired males and females. There was a male-biased sex ratio in each year, and between 5% and 26% of males remained unpaired throughout the breeding season. Male mating status and breeding success were dependent on arrival date, territory rank and breeding density. Early-arriving, usually older, males were able to settle on territories first and were more likely to pair, while later-arriving individuals were more likely to remain unmated. These effects were consistent between years, and consequently territories could be classified as either preferred (accounting for proportionately more breeding attempts) or non-preferred. Territory quality as opposed to individual quality appeared to explain much of the variation in breeding success, and both sexes benefited by breeding on preferred territories through enhanced breeding success and an increased probability that their offspring would be recruited to the population.     

Effects of breeding success, mate fidelity and senescence on breeding dispersal of male and female blue-footed boobies

1. Understanding the effects of individual and population factors on variation in breeding dispersal (the movement of individuals between successive breeding sites) is key to identifying the strategies behind breeders' movements. Dispersal is often influenced by multiple factors and these can be confounded with each other. We used 13 years of data on the locations, mates, breeding success and ages of individuals to tease apart the factors influencing breeding dispersal in a colonially breeding long-lived seabird, the blue-footed booby Sula nebouxii. 2. Breeding dispersal varied among and within years. Males dispersed further in years of higher population density, and late breeding males and females dispersed further than early breeders. This temporal variation related to changes in competition for territory was taken into account in all tests of individual factors influencing breeding dispersal. 3. Individuals that retained their mates from the previous year dispersed shorter distances than those that changed their mates. 4. The effect of previous breeding success depended on mate fidelity. Unsuccessful breeding induced greater dispersal in birds that changed their mates but not in birds that retained their mates, indicating that breeders who change mates may take their own previous breeding experience into account during habitat selection. Faithful individuals may have to stay close to their previous sites to encounter their mates. 5. Male divorcees dispersed over shorter distances than their former mates, possibly because males contribute more than females to establishing territories. 6. Dispersal of males and females declined with increasing age over the first 10-11 years of life, then increased in old age, possibly due to senescent decay in the ability to compete for mates and territories.     

OBSERVATIONS OF GROUP BEHAVIOUR AND BREEDING BIOLOGY OF THE YELLOW-BILLED SHRIKE CORVINELLA CORVINA

Yellow-billed Shrikes were found to live in groups throughout the year. Within the group, each member helped to defend the group's territory, warn against predators and feed the breeding female, nestlings and fledglings.
During the study there was little change in the location of the boundaries and in the areas of the territories occupied by the majority of the groups. The densities of the larger groups were in general two to three times that of smaller groups. Numbers within one group varied by ±24% of the average (12) during a period of three years.
Progeny remained in a group for some years before dispersing, sometimes in parties of the same sex. Both sexes exchanged groups, the females moving on average further than males. During successive periods in the history of a group the representation of the sexes varied from a surplus of females to a surplus of males. In the population as a whole the sex ratio was probably parity.
Only one female bred in a group at a particular time and she alone incubated. Eggs were laid on consecutive days. Breeding started at the height of the dry season; the first peak in egg laying occurred at the beginning of the rains; laying continued through the wet season and ceased usually in August. The most frequent clutch size was four, and varied little within a breeding season or between seasons. The incubation period ranged from 15 to 18 days, the most frequently recorded being 17 days. The nestling period was 19 days. The percentage of total eggs laid that produced fledglings was 25% and yearlings 11%.
Young shrikes were independent in the seventh week, participated in group displays in their tenth week and fed fledglings in their fourteenth week.
The age of first breeding was not discovered. Two females in their sixth year were still helpers in a group at the end of the study.     

A Polygamous Social System of the Fan-tailed Warbler Cisticola juncidis

The social system of an individually-marked population of the fan-tailed warbler Cisticola juncidis was studied in Japan, over six breeding seasons from 1978 to 1983. More than 127 males established territories, some 50–70% were polygynists each year. Territorial males were replaced frequently within seasons. Females were less faithful than males to their first breeding sites. Perennial or seasonal pair bonds were rare, maintained over two successive breeding attempts, by only 13.6% of females. Half the females left the area after one breeding attempt. Frequent divorce, rapid and multiple remating of females, multiple breedings, and female movement over a wide area all combine to skew breeding sex ratio from unity and favor polygyny.     

Why are Male Columbian Ground Squirrels Territorial?

Male territorial defence is a component of many vertebrate mating systems and is often regarded as a tactic for acquiring mates. Traditionally considered within the context of overt site‐specific defence, territoriality actually may have several components which encompass a variety of behavioural tactics (e.g. post‐copulatory mate‐guarding, defence of resources that females need, defence of area around females) that underlie a mating system. The purpose of our study was to evaluate such influences on the territorial behaviour of male Columbian ground squirrels in southwestern Alberta, Canada. Males were dominant and territorial if they defended a minimum convex polygon activity range by chasing other males more within the activity range than they were chased. Subordinate males had no territory and were chased throughout their ranges, but they competed for mates by increasing chases in their activity range when nearby females were oestrous. Dominant males exhibited conditional breeding tactics, tending to chase other dominant males from their territory when nearby females were oestrous, but travelling outside their activity ranges to chase subordinate males when females were not oestrous. Although females mated first with a dominant male on whose territory they resided (and in order from oldest to youngest if several territories overlapped), mating pairs were not exclusive, as females usually mated with additional males. Males also guarded females after copulation and defended females directly just before oestrus, rather than defending territory per se during those times. Thus, males possess a repertoire of behaviours that complement site‐specific territoriality, and territory ownership serves to facilitate a first mating with females that live on the territory.     

Breeding performance of a captive colony of rhesus macaques (Macaca mulatta)

For most of the 18 years recorded, fewer than 50% of the adult females gave birth in any one year. The colony, of 6 social groups, showed a clear-cut breeding season. Female parity and dominance had no effect on breeding rate, though 1st infants were born earlier in the year than 2nd-born ones. Only when females gave birth in successive years were the months of giving birth correlated. Mothers and daughters may tend to give birth closer in time within a breeding season than do other females.     

Successive nest building and polygyny of Fan-tailed Warblers Cisticola juncidis

The polygynous mating system of the Fan-tailed Warbler Cisticola juqcidis was investigated between 1978 and 1981. The male warbler builds many nests unaided; however, he has no more than one active vacant nest for courting at any time. Nest building lasted from April until August. After completing the outer fabric of a nest, the male advertised it and led a female to the nest site by a unique invitation flight. On average a male built 6.5 nests during one breeding season and three of them were accepted by females. The most successful male completed 18 nests, and mated with 11 females. Out of a total of 111 males which established a territory, 30 had no mate, 14 were monogamous, and the rest were polygynous. About 50 to 70% males were polygynous over the four years. The sex ratio varied from 1.41:1 to 2.17:1 (females: male) in the breeding population. It was partly caused by the presence of 'floating males'. After the completion of the outer fabric of the nest, the male warbler did not take any further role in nesting and caring for young.
The polygynous mating system of the Fan-tailed Warbler is characterized by successive nest building. Its extreme development results from the long breeding period and the male having no role in parental care.     

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 life cycle of the Upland Goose Chloëphaga picta in the Falkland Islands

Upland Geese Chloëphaga picta were studied between 1977 and 1980, primarily around Darwin, East Falkland, in order to describe their breeding biology, moulting and adult survival. The population of breeding birds in a valley reached a peak from mid-September to late November when nesting took place. The average territory length was 240 m in five valleys. Breeding adults generally returned to breed in the same territory each year and with the same mate. Nests were on the ground, usually amongst whitegrass Cortaderiapilosa. The mean clutch size was 6-1, brood size was 5-1 and fledged family 3–9. Incubation took 30 days and the fledgling period was about 70 days. Most broods were raised in the nesting territory. Growth of goslings is described. The breeding success between laying and fledging was 0–34 (in 1977) and 0–29(in 1978), giving an annual production of 21 and 1–8 young per breeding pair. Fledglings remained in family parties through the autumn and winter and were evicted by their parents in early spring. Some siblings stayed together for short periods and then joined other non-breeders. Females started pairing at ten months of age and most were paired at 17–18 months. Some bred for the first time at 23 months. Males started pairing at about 20 months of age. Flightless moult (shedding) took place at ponds or in sheltered inlets of the sea, in flocks of up to several hundred birds. Flightless birds were found between 14 November and 11 February, though 50% were flightless between 26 November and 2 January. Individuals were flightless for 36 days. First-year birds were more synchronized in shedding than adults. The percentage of first-year males (in the male component) varied from 16-5 to 45-9% in shedding flocks, and significantly more males were present in some flocks. The flocks were composed of first-year and second-year birds too young to breed and failed breeders. The percentage of a shedding population which returned to the same site in successive years was 25-3 and 15-1% at two localities. The moult of other feather tracts is described. The annual survival rate of breeding adults was 82%. A model of the population dynamics is presented. The current level of culling to control the goose population is less than the number which must die each year to maintain a stable population.     

Pair territoriality in the longnose filefish,oxymonacanthus longirostris

The longnose filefish,Oxymonacanthus longirostris, usually lives in heterosexual pairs, the male and female swimming together and sharing the same territory. Pair territoriality in the species was examined in detail in relation to sexual differences in territorial defense activities. Rigorous pair territoriality was maintained only during the breeding season, although pairs used their home ranges exclusively to a certain extent, during the non-breeding season. The frequency of aggression against other conspecific pairs in the breeding season was higher than in the non-breeding season. Agonistic interactions appear to be over both mates and food resources, the strict pair territoriality in the breeding season possibly being due to mutual mate guarding. In intraspecific aggressive interactions, males usually led their partner females when attacking intruders. The feeding frequency of males was much lower than that of females in the breeding season. Mate removal experiments indicated that females could not defend their original territories solitarily and their feeding frequency decreased. Conversely, males could defend territories solitarily without a decrease in feeding frequency. These results suggest that males contribute most to the defense of the pair territory, with females benefiting from territorial pair-swimming with their partner males.     

BREEDING OF SACRED IBIS THRESKIORNIS AETHIOPICA AT LAKE SHALA, ETHIOPIA

Data are based on more than 200 h of observation at Ethiopia's Lake Shala from 1966 to 1972. Except for differences in size of bill, there are no useful field characters separating male and female Sacred Ibis. The breeding plumage is described; vivid blood-red colour underneath the wings and the ornamental plumes are especially obvious when nesting commences. Physical and biological features of Lake Shala, Ethiopia, and its nesting islands are described; the species of birds nesting on the Shala islands are given. Ibises nest at Shah from March to August; no nesting has been recorded from September to February during the last months of the ‘big’ rains through the main dry season. Nesting normally begins in the ‘small’ rains (between 14 March-24 April), although instances were recorded as early as 1 March and as late as 20 August. The ibises normally nest once per year, although it is possible that occasionally a second nesting may occur after an unsuccessful first attempt. The ibises at Shala nest in discrete groups; several nesting groups may form on any or all of the islands; the number of groups attempting to nest varied from year to year. Nesting activity begins when males arrive and establish pairing territories, usually in a small tree but sometimes on the ground. When females and other males arrive at the pairing territories, pair formation ensues. At this time males perform forward threat, modified forward threat, pursuit flight, supplanting attack and modified snap displays, while both sexes perform stretch and bow displays. Once established, the pair abandons the pairing territory and moves to the nesting area, usually near but always distinct from the pairing territory, and establishes a nest-site territory. Most members of the nesting group move to the nesting area on the same day. Copulation then takes place, and is followed by collection of nest material, usually by the male. Nests are built close together. The average area of 10 nests measured was 0.09 m². Nests are usually less than 20 cm thick and are made of many small branches and sticks. The average clutch in 34 nests was 2.24 eggs; the average size of 34 eggs was 63.4×43.5 mm. Incubation probably begins when the clutch is complete. Both sexes incubate, and the incubation period probably lasts 28–29 days. The development of the young is described. The young leave the nest-site territory when 14–21 days old. Although they are capable of some flight when 35–40 days old, the young do not leave the colony until they are 44–48 days old. In the colony, both parents care for the young. Usually only one parent at a time is with the young. The parents recognize their own young and are usually recognized by them. The behavioural interactions between young and parents are described. Fledging success in 1968 was 1.06 young per pair. The number of pairs successfully rearing young varied annually from none to 81%, on average over six years (1966–70, 1972) 35%. Predation at the breeding colonies is minimal. The food of one one-month old chick consisted of beetle larvae, lepidopteran larvae and beetles. Feeding areas, although undetermined, must be widespread. Inter-specific competition between Sacred Ibis and other nesting birds at Shala is discussed. Among possible factors stimulating nesting at Shala one, fairly heavy rainfall, seems to be especially important. It is also suggested that especially heavy rain-storms cause ibises to abandon the colonies, and result in poor breeding success.     

Does competition for nests affect genetic monogamy in Cory's shearwater Calonectris diomedea?

Most bird species are socially monogamous. However, extra‐pair copulations (EPCs), resulting in extra‐pair paternity (EPP), commonly occur. EPCs should allow females to adjust social mate choice and allow males that fail to obtain a nest a chance to avoid missing a breeding season, especially when poor nest supply constrains social mate choice. Procellariiformes (albatrosses and petrels) are socially monogamous seabirds which seldom divorce, even when nest availability constrains social mate choice. In Cory's shearwater Calonectris diomedea, a burrow‐nesting petrel, two studies conducted in the Mediterranean, where competition for nests is weak, detected no EPP. EPP remains to be investigated at localities where competition for nests is much stronger, such as Vila islet, Azores archipelago, Atlantic Ocean. We conducted a genetic (microsatellites) study over two successive years on Vila, involving the breeding pairs of the same 65 nests each year and their single chick. EPPs occurred each year, the overall rate being 11.6%. Coupling genetic analyses to a 7‐year demographic survey provided additional data on pair bonds and competition for nests. Overall, cuckoldry was unrelated to divorce, nest density and inbreeding avoidance, but was more frequent when the social male was small. Nest changes were more costly for males than for females, and some apparently unpaired males attempted to dislodge social males during within‐pair copulations. These results are compatible with the existence of a link between poor nest availability and EPP and confirm that even species considered strongly monogamous can adopt flexible mating strategies.     

The Cooperative Breeding System of the Red-cockaded Woodpecker

The breeding system of the red-cockaded woodpecker is described based on data collected over six years from a population of 500 marked individuals in the Sandhills of North Carolina. Male-female pairs were the most common social unit (59%), but 30% of social units contained one or more adult helpers, and 11% consisted of solitary males. Helpers were almost exclusively male: 27% of males remained in their natal group as helpers for at least one year, whereas only four (1%) females did. Most breeding females remained as breeders in the same group from one year to the next (56%), but a surprising number (12%) moved to another group. Many movements were related to incest avoidance or mate death, but 39% involved deserting a mate, usually following successful reproduction. We suggest that females sometimes are forced from groups by immigrants or other group members. The median distance of movements by adult females was only 1.3 km. In contrast to females, no breeding males switched groups. Survival of both breeding (76%) and helper (80%) males was higher than that of breeding females (69%). Males exhibited two distinct life-history strategies. Some remained as helpers on their natal territory for one or more years, and became breeders by inheriting breeding status in the natal group (17% per year) or by replacing a deceased breeder in a nearby group (13% per year, median distance moved 1.0 km). Other males dispersed from their natal group permanently during their first year. Some of these males were floaters at age one year, others were solitary, and a few became helpers in a non-natal group, but many were breeders. In contrast to males that first functioned as helpers, those that dispersed after fledging moved long distances (median dispersal distance 4.5 km), longer even than dispersing female fledglings moved (median distance 3.2 km). The habitat saturation model of the evolution of cooperative breeding is based on selection between the two life-history strategies exhibited by male red-cockaded woodpeckers. The model therefore may be tested directly with this species. Another indication that this model is appropriate for this species is the existence of a resource (cavity trees) that might provide an ecological basis for habitat saturation.     

Annual variation in the timing of reproduction in Antarctic fur seals, Arctocephalus gazella, at Bird Island, South Georgia

The arrival of Antarctic fur seals at a breeding beach on Bird Island, South Georgia, was studied over five consecutive breeding seasons, 1983 to 1987. Experienced bulls arrived first and established breeding territories on the beaches in anticipation of the arrival of the cows. Male arrival, which is less synchronous within years than female arrival, was significantly later in 1987 than in any other year. Female arrival, estimated by pup birth date where necessary, was highly synchronous; it usually started when 80% or more of potential territory sites were occupied by males. Cows arrived significantly later in 1984 and 1987 than in 1983, 1985 or 1986. The late arrival of both males and females in 1987 is attributed to unusually severe climatic conditions during the preceding winter. The late arrival and reduced fecundity of females in 1984 is attributed to markedly reduced food availability during the austral winter and summer of 1983. Males were not affected in 1984 because they could move away from the area of reduced food availability earlier than females and because they have a more varied diet. Factors influencing the winter distribution, the timing and pattern of arrival and the breeding of male and female Antarctic fur seals are discussed.     

Group structure,within-group conflict and reproductive tactics in cooperatively breeding Galápagos mockingbirds,Nesomimus parvulus

Reproductive conflict within groups can be an important feature of cooperative breeding systems, especially when more than one individual of a sex breeds within a social group. Relationships between group structure, dominance, within-group conflict and reproductive tactics of cooperatively breeding Galápagos mockingbirds were examined on Isla Genovesa. Territorial groups of 2–24 adults included up to three breeding females, with 42% of the groups containing more than one (plural groups); females in most plural groups nested separately. Territory size increased with group size, but the area available per pair in plural groups was smaller than in singular groups (groups with only one breeding pair). Most pairings were monogamous, and males usually outnumbered females; high-ranking males obtained mates more frequently than subordinate males. In 3 relatively dry years, but not in a wet El Niño year, subordinate pairs in plural groups fledged fewer young than dominant pairs or pairs breeding in singular groups. Interference by dominant breeders, often leading to abandonment of nests by subordinate pairs, appears to account for these differences: through nest disruption in drier years, dominant individuals may reduce the cost of sharing their territories and increase the chances of recruiting helpers. Dominant males in plural groups may also father young through extra-pair copulations with subordinate females. Despite costs imposed by within-group conflict, subordinate breeders have higher long-term reproductive success than birds that defer breeding. Plural group structure is maintained because unpredictable climatic variation favours opportunistic breeding by subordinates.     

Sex-specific patterns in body condition and testosterone level changes in a territorial migratory bird: the Bluethroat Luscinia svecica

The Bluethroat Luscinia svecica is a migratory passerine that exhibits a socially monogamous pair bond and a high level of parental care. Males are territorial both when wintering and breeding whereas females are territorial only in winter. We investigated changes in body condition and testosterone levels during successive life-history stages and determined their relationships. Sex-specific patterns were observed in the variation in body condition and testosterone level. Male body condition varied mainly during the winter. It peaked at the onset of the prenuptial moult and then decreased, whereas it remained stable throughout breeding. In contrast, female body condition varied mainly during the breeding season. It increased during the prelaying stage and then abruptly decreased until fledgling provisioning. As in other monogamous and territorial passerines, testosterone levels in Bluethroat males were low during winter, increased in late winter, peaked during the prelaying stage and then decreased when provisioning young. In wintering females, territorial competition caused testosterone levels to rise. These females were able to produce territorial vocalizations and exhibit aggressive postures. Females showed higher mean testosterone levels than males when wintering whereas the opposite was observed when breeding. Our data from wintering female Bluethroats support the 'challenge hypothesis' under which high testosterone levels are associated with periods of social instability, and testosterone can regulate female territorial behaviour during this period.