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.     

Breeding sites and winter site fidelity of Piping Plovers wintering in The Bahamas,a previously unknown major wintering area

Most of the known wintering areas of Piping Plovers (Charadrius melodus) are along the Atlantic and Gulf coasts of the United States and into Mexico, and in the Caribbean. However, 1066 threatened/endangered Piping Plovers were recently found wintering in The Bahamas, an area not previously known to be important for the species. Although representing about 27% of the birds counted during the 2011 International Piping Plover Winter Census, the location of their breeding site(s) was unknown. Thus, our objectives were to determine the location(s) of their breeding site(s) using molecular markers and by tracking banded individuals, identify spring and fall staging sites, and examine site fidelity and survival. We captured and color‐banded 57 birds in January and February 2010 in The Bahamas. Blood samples were also collected for genetic evaluation of the likely subspecies wintering in The Bahamas. Band re‐sightings and DNA analysis revealed that at least 95% of the Piping Plovers wintering in The Bahamas originated on the Atlantic coast of the United States and Canada. Re‐sightings of birds banded in The Bahamas spanned the breeding distribution of the species along the Atlantic coast from Newfoundland to North Carolina. Site fidelity to breeding and wintering sites was high (88–100%). Spring and fall staging sites were located along the Atlantic coast of the United States, with marked birds concentrating in the Carolinas. Our estimate of true survival for the marked birds was 0.71 (95% CI: 0.61–0.80). Our results indicate that more than one third of the Piping Plover population that breeds along the Atlantic coast winters in The Bahamas. By determining the importance of The Bahamas to the Atlantic subspecies of Piping Plovers, future conservation efforts for these populations can be better focused on where they are most needed.     

FACTORS AFFECTING THE AGE OF FIRST BREEDING OF THE KITTIWAKE RISSA TRIDACTYLA

At a Kittiwake colony in Northumberland, 80% of those birds which returned to their natal colony to breed were males and these supplied 52% of all male recruits. More females breed away from their natal colony than males. There was no differences in the proportions of young fledged from sites in the centre or at the edge of the colony, or by parents of different experience, which returned to breed. Kittiwakes breed for the first time at ages from 3 to 8 years, but most at 4 or S years old. Males arrive back at the colony at an earlier age than females and breed for the first time one year earlier. Males obtaining sites at the centre of the colony first breed at an earlier age than those at the edges. Neither the age nor the area of first breeding appear to be transmitted from parent to offspring. Males breeding first aged 4 years or younger produced more young than those which first bred aged 5 years or older, despite their partners laying smaller clutches. This difference was most marked among those males recruited to sites in the centre of the colony. The advantage of this earlier breeding is counteracted by a lower survival rate among those males which start to breed at the younger ages. In all breeding Kittiwakes, annual reproductive output increases with experience while annual survival rates decrease. Once they had started to breed, many birds failed to breed in one subsequent season. Nearly 60% of these cases of intermittent breeding occurred in the year following first breeding. Intermittent breeding was most frequent among young birds and among females. It is suggested that each breeding involves a cost to the individual in terms of reduced survival, and that deferred and intermittent breeding are means of guarding survival. A model is proposed whereby the age at which a bird starts to breed, the nesting site which it obtains, and its subsequent breeding strategy result in each individual producing an optimal number of reproducing offspring in its lifetime, relative to its quality.     

Effects of age, cohort and individual on breeding performance in the Lapwing Vanellus vanellus

The breeding performance of individually colour-ringed Lapwings Vanellus vanellus was studied on marginal grassland in Upper Teesdale, County Durham (UK), from 1992 to 1995. Contrary to many studies of birds, female age had only a minor effect on breeding performance: yearling females produced eggs on average 5% smaller than those of adults. In contrast, the average productivity of male Lapwings was estimated to increase by around 40% between one and two years of age, and by a further 10% between two and three years of age. This was because of a 40% increase in the proportion of males that bred between one and two years of age, and of an increase in mating success with age coupled with a higher breeding success for polygynous males. The likelihood of breeding for male Lapwings was affected by year of hatching. More than half of those males that hatched in 1990, and which were present in both 1993 and 1994, did not breed. It was suggested that this was due to environmental effects experienced by these males as chicks, resulting in reduced success in securing a breeding territory later in life. Individual female Lapwings showed a high degree of consistency between years in both laying date and egg size, and those that raised up to two young in 1993 raised approximately the same number in 1994. However, females producing more than two young in 1993 raised significantly fewer in 1994, perhaps suggesting a cost of reproduction.     

Senescence and carryover effects of reproductive performance influence migration,condition, and breeding propensity in a small shorebird

Breeding propensity, the probability that an animal will attempt to breed each year, is perhaps the least understood demographic process influencing annual fecundity. Breeding propensity is ecologically complex, as associations among a variety of intrinsic and extrinsic factors may interact to affect an animal's breeding decisions. Individuals that opt not to breed can be more difficult to detect than breeders, which can (1) lead to difficulty in estimation of breeding propensity, and (2) bias other demographic parameters. We studied the effects of sex, age, and population reproductive success on the survival and breeding propensity of a migratory shorebird, the piping plover (Charadrius melodus ), nesting on the Missouri River. We used a robust design Barker model to estimate true survival and breeding propensity and found survival decreased as birds aged and did so more quickly for males than females. Monthly survival during the breeding season was lower than during migration or the nonbreeding season. Males were less likely to skip breeding (range: 1–17%) than females (range: 3–26%; β_sex = ?0.21, 95% CI: ?0.38 to ?0.21), and both sexes were less likely to return to the breeding grounds following a year of high reproductive success. Birds that returned in a year following relatively high population‐wide reproductive output were in poorer condition than following a year with lower reproductive output. Younger adult birds and females were more likely to migrate from the breeding area earlier than older birds and males; however, all birds stayed on the breeding grounds longer when nest survival was low, presumably because of renesting attempts. Piping plovers used a variety of environmental and demographic cues to inform their reproduction, employing strategies that could maximize fitness on average. Our results support the “disposable soma” theory of aging and follow with predictions from life history theory, exhibiting the intimate connections among the core ecological concepts of senescence, carryover effects, and life history.     

Age-related patterns of reproductive success in house sparrows Passer domesticus

A common life history pattern in many organisms is that reproductive success increases with age. We report a similar pattern in house sparrows Passer domesticus , older individuals performed better than yearlings for most measures of reproductive success. Older males and females began breeding earlier in a given season and fledged more young than their yearling counterparts. Individual males also fledged more young in their second breeding season than they did in their first, but individual females did not show consistent improvement in reproductive success from year one to two. A path analysis indicated that age in both sexes acted primarily through the timing of breeding; earlier nesters laid more eggs and hence fledged more young but did not have more nesting attempts. We tested whether the increased reproductive success with age arose from high quality individuals surviving to be older (selection hypothesis). In contrast to the main prediction of this hypothesis that reproductive success and survival should be positively related, we found that survival from one year of age to two years of age was negatively related to reproductive success in the first year for males and females combined. Additionally, individuals that survived to breed as two-year-olds did not differ in total young fledged in their first year from those that did not survive to their second season of breeding. Our results indicate that fledgling production increases with age due to improvements in timing of breeding, particularly in females, and not because of the loss of poor breeders or increased output. Mechanisms producing age-related differences in timing of breeding warrant further study.     

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.     

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.     

Space use by Great Lakes Piping Plovers during the breeding season

ABSTRACT.   Identifying and protecting breeding habitat for imperiled species requires an understanding of the spatial and temporal movements of breeding individuals. During the 2003 and 2004 breeding seasons, we examined space use by Piping Plovers ( Charadrius melodus ) in the federally endangered Great Lakes population. We used coordinate geometry to estimate home range sizes of individual birds and examined relationships between home range size and breeding stage (incubation versus chick rearing), year, sex, number of locations, minimum plover age, distance to the nearest nest, and human beach use (high, medium, or low). The mean size of home ranges of Piping Plovers that fledged at least one chick was 2.9 ± 0.5 (SE) ha (range = 0.4–11.2 ha), and the mean linear beach distance traversed was 475 ± 53 m (range = 130–1435 m). Individuals used 3 times more beach area and 1.5 times more shoreline distance in 2003 than in 2004. Females used smaller areas than males overall and during chick rearing. Home ranges were smallest on beaches with low public use, suggesting that human disturbance may cause greater movement by individual plovers and that larger protected areas may be warranted on beaches frequented by the public to minimize disturbance to breeding birds. Our results demonstrate that nesting Great Lakes Piping Plovers occupy relatively small ranges and, therefore, that even relatively small areas of suitable habitat can have a high conservation value for this endangered population. However, the total area of habitat used varied substantially among individuals, and this should be considered when protecting habitat for the species.     

To breed or not to breed: causes and implications of non-breeding habit in the willow tit Parus montanus

Causes and consequences of non-breeding in willow tits were studied in northern Finland during 1986–1992. The breeding status was sex and age biased; males and yearling birds were in excess among the non-reproducers. Due to sex bias in the population it appeared detrimental for males to lose a mate, especially shortly before breeding. Lack of a mate was a important factor for males not reproducing (37% of non-breeding males) than for females (14%). Most of the non-breeding birds maintained a pair bond which only rarely broke up for the next breeding season (divorce rate 5.5%). This implies that parental incompatibility is not a possible explanation for pairs not reproducing. Males that did not breed tended to survive better than reproducing ones, whereas such a relationship was not found for females. It is possible that this sex-related difference in survival cost is attributable to quality differences among non-breeding individuals. It was especially low-quality yearling females, with low survival prospects, that were responsible for the discrepancy. The proportion of non-breeding females in the population correlated highly with clutch size and subsequent juvenile survival. It is therefore suggested that for most of these females non-breeding is a phenotypic response to low offspring value in the prevailing circumstances (inter-generational tradeoff). However, it is uncertain whether willow tits in a northern population can use breeding density as an indicator of changing survival prospects of their descendants, as suggested by Ekman and Askenmo (1986) for southern Sweden.     

Survival,site fidelity,and the population dynamics of Piping Plovers in Saskatchewan

ABSTRACT To conserve threatened species, managers require predictions about the effects of natural and anthropogenic factors on population growth that in turn require accurate estimates of survival, birth, and dispersal rates, and their correlation with natural and anthropogenic factors. For Piping Plovers (Charadrius melodus), fledging rate is often more amenable to management than adult survival, and population models can be used to estimate the productivity (young produced per breeding female) necessary to maintain or increase populations for given levels of survival. We estimated true survival and site fidelity of adult and subadult (from fledging to second year) Piping Plovers breeding in Saskatchewan using mark‐resight data from 2002 to 2009. By estimating true survival rather than apparent survival (which is confounded with permanent emigration), we were able to provide more accurate projections of population trends. Average adult and subadult survival rates during our study were 0.80 and 0.57, respectively. Adult survival declined over time, possibly due in part to the loss of one breeding site to flooding. Average adult and subadult site fidelity were 0.86 and 0.46, respectively. Adult site fidelity declined during our study at two study sites, most strongly at the flooded site. Male and female Piping Plovers had similar survival rates, but males had greater site fidelity than females in some years. Based on our survival estimates, productivity needed for a stationary population was 0.75, a benchmark used for plover management on the Atlantic Coast, but not previously estimated for Prairie Canada. In stochastic simulations incorporating literature‐based variation in survival rates, productivity needed for a stationary population increased to 0.86, still lower than that previously estimated for western populations. Mean productivity for our study sites ranged from 0.87 to 0.96 fledged young per pair. Our results suggest that fledging rates of Piping Plovers in Saskatchewan were sufficient to ensure a stationary or increasing population during our study period. However, large‐scale habitat changes such as drought or anthropogenic flooding may lead to dispersal of breeding adults and possibly mortality that will increase the fledging rate needed for a stationary population.     

Seasonal abundance of nonbreeding Piping Plovers on a Georgia barrier island

ABSTRACT.   Although breeding populations of Piping Plovers are well studied, their winter distribution is less clear. We studied the seasonal abundance of nonbreeding Piping Plovers ( Charadrius melodus ) during the winters of 2003–2004 and 2004–2005 on Little St. Simons Island (LSSI), Georgia. Our objectives were to determine the relative abundance of individuals from three breeding populations at LSSI, and identify possible differences among populations in arrival time, winter movements, or departure time. We observed up to 100 Piping Plovers during peak migration, and approximately 40 plovers wintered at LSSI. From July 2004 to May 2005, approximately 20% of the Great Lakes breeding population used LSSI. Plovers were not present at LSSI during June. All breeding populations of Piping Plovers had similar patterns of temporal occurrence on LSSI, suggesting no need for population-specific management plans at this site. Our results suggest that LSSI is among the most important wintering sites on the Atlantic coast for Piping Plovers, especially for individuals from the endangered Great Lakes population.     

The timing, duration and pattern of moult and its relationship to breeding in a population of the European Greenfinch Carduelis chloris

Moult was studied in 1 year among Greenfinches trapped in a garden in east‐central England. Over the period June–December 2003, 333 captures of 179 individual adults provided information on breeding condition, moult, body weight, sex and age (yearling or older adult, equivalent to birds in their second or later calendar years, respectively). About 95% of all birds (sex and age groups combined) started primary feather moult from 2 July to 14 August, and finished from 10 October to 22 November. The mean date of moult onset in the population as a whole was 24 July. On average, males began 8 days before females, and yearlings began 6 days before older birds. The mean duration of moult was 100 days, whether the figure was calculated for the population as a whole or just for the 36 individual birds that were caught more than once during moult. However, moult rate was slightly slower, and moult duration slightly longer, in yearlings than in older adults of both sexes. No evidence was found for any systematic relationship between moult onset date and rate (duration). Breeding and moult overlapped by up to 5 weeks or more in individual birds, and some birds probably started to moult as early as the incubation stage of their last clutch of the season. The cloacal protuberance (taken as indicative of breeding condition) had regressed in all males by the time the fifth primary was shed, and the brood patch had regressed and re‐feathered in all females by the time the fourth primary was shed. The bulk of feather replacement in the secondary, tail and body tracts occurred in the second half of primary moult, and after cloacal protuberances and brood patches were completely regressed. In all birds examined near the end of primary moult the secondaries were still growing, and would have continued growth for up to another 19 days or more, extending the end of the moulting season into December. Body mass during moult was affected significantly by sex and age, as well as by time of day, amount of food in gullet, reproductive condition and date. No firm evidence emerged that body mass was affected by moult stage, after allowing for effects of date and other variables (although there was a non‐significant negative relationship between moult stage and body mass in males). In the population as a whole, the breeding season (from first egg‐laying to independence of last young) was spread over 21 weeks and moult over 24 weeks. With an overlap between the two events at the population level of up to 9 weeks, the two processes together took up to 36 weeks, some 69% of the year.     

State uncertainty models and mark‐resight models for understanding non‐breeding site use by Piping Plovers

Conservation of beach‐nesting medium‐distance migrants has focused on breeding areas because protection of nests is more tractable than protection of non‐breeding habitat. As breeding ground management has encountered diminishing returns, interest in understanding threats in non‐breeding areas has increased. However, robust estimates of non‐breeding demographic rates and abundance are generally lacking, hindering the study of limiting factors. Estimating such rates is made more difficult by complex population dynamics at non‐breeding sites. In South Carolina, endangered Piping Plovers Charadrius melodus start arriving in July and some depart prior to December (the autumn‐only population) while others remain through at least March (the wintering population). State uncertainty capture‐mark‐recapture models provide a means for estimating vital rates for such co‐occurring populations. We estimated the proportion of the population entering the study area per survey (entry probability) and proportion remaining per survey (persistence rate) for both populations during autumn, and abundance of the wintering population, at four sites in South Carolina in 2006/7 and 2007/8, taking advantage of birds previously colour‐ringed on the breeding grounds. We made fairly precise estimates of entry and persistence rates with small sample sizes. Cumulative entry probability was ~50% by the end of July and reached 95% for both populations by October. Estimated stopover duration for birds in the autumn‐only population was 35 days in year 1 and 42 days in year 2. We estimated a wintering super‐population size of 71 ± 16 se birds in the first year and 75 ± 16 in the second. If ringing programmes on the breeding grounds continue, standardized resighting surveys in the non‐breeding period and mark‐recapture models can provide robust estimates of entry and persistence rates and abundance. Habitat protection intended to benefit non‐breeding Piping Plovers at our coastal sites should be in effect by late summer, as many birds are resident from July to the end of winter.     

Breeding performance and survival in the peregrine falcon Falco peregrinus support an age‐related competence improvement hypothesis mediated via an age threshold

We studied the effects of age on breeding performance and survival probability in a peregrine falcon population, using data from a long term monitoring programme (carried out over 16 yr), in which we were able to identify individual birds. We compared the breeding performance and survival of yearling breeders, first‐time adult breeders and adult breeders. We found significant differences in breeding performance but not in survival. Yearling breeders had lower breeding success than older individuals but the breeding performance of inexperienced adults did not differ from that of experienced adults. We did not find changes in terminal breeding success since peregrines in their last year of life sustained the performance levels shown in previous years although with increasing variability. We found no evidence that attempting to breed affected survival probability in any age group. We argue that differences in breeding performance are related to age, not to breeding experience, and that there is an age threshold, coincident with the development of adult plumage, after which breeding performance is not affected either by age or experience. Peregrines that start breeding as yearlings are likely to have greater lifetime reproductive success than birds entering the breeding pool as adults. Consequently, such birds may represent a set of high quality individuals. Our results support the age‐related competence improvement hypothesis as being the relevant explanation for the increase in breeding performance with age.     

Prolonged and flexible primary moult overlaps extensively with breeding in beach‐nesting Hooded Plovers Thinornis rubricollis

We present the first report of complete overlap of breeding and moult in a shorebird. In southeastern Australia, Hooded Plovers Thinornis rubricollis spend their entire lives on oceanic beaches, where they exhibit biparental care. Population moult encompassed the 6‐month breeding season. Moult timing was estimated using the Underhill–Zucchini method for Type 2 data with a power transformation to accommodate sexual differences in rates of moult progression in the early and late stages of moult. Average moult durations were long in females (170.3 ± 14.2 days), and even longer in males (210.3 ± 13.5 days). Breeding status was known for most birds in our samples, and many active breeders (especially males) were also growing primaries. Females delayed the onset of primary moult but were able to increase the speed of moult and continue breeding, completing moult at about the same time as males. The mechanism by which this was achieved appeared to be flexibility in moult sequence. All moult formulae fell on one of two linked moult sequences, one faster than the other. The slower sequence had fewer feathers growing concurrently and also had formulae indicating suspended moults. Switching between sequences via common formulae is possible at many points during the moult cycle, and three of 12 recaptures were confirmed to have switched sequences in the same moult season. Hooded Plovers thus have a prolonged primary moult with the flexibility to change their rate of moult; this may facilitate high levels of replacement clutches that are associated with passive nest defence and low reproductive success.     

Age at first breeding, philopatry and breeding site-fidelity in the Lapwing Vanellus vanellus

Breeding Lapwings Vanellus vanellus were studied in the Eden Valley (Cumbria) and in Teesdale (County Durham) between 1990 and 1992. A total of 300 adult Lapwings and 801 near-fledged young were uniquely colour-ringed. Breeding adults were highly site-faithful, almost always nesting in the same or an adjacent field in successive years. Second-year birds were less site-faithful, with more birds nesting in adjacent and other fields and fewer in the same field in successive years. In Teesdale, 74% of colour-ringed young Lapwings returned in their first or second year of life to within 5 km from where they hatched. In contrast, in the Eden Valley only 37% of young birds in their first or second year of life returned to within 5 km from where they hatched. From an analysis of British ringing recoveries in April and May, 61% of Lapwings were recovered within 10 km from where they were ringed as chicks. A further 11% were recovered more than 100 km from where they were ringed. Young Lapwings were highly philopatric, with 45% of males and 52% of females breeding in the same field or a field adjacent to where they hatched. The majority of Lapwings (67%) began breeding at 1 year old. Of the remaining birds, 27% bred for the first time when 2 years old and 6% for the first time in their third year of life. There was no difference between the sexes. Chicks hatching and subsequently fledging late in the season returned less frequently to the study areas in subsequent years than did chicks hatching earlier in the season.     

THE BIOLOGY OF THE WAVED ALBATROSS DIOMEDEA IRRORATA OF HOOD ISLAND, GALAPAGOS

As a nesting species, the Waved Albatross Diomedea irrorata is restricted to Hood Island in the Galapagos archipelago where 12,000 pairs bred in 1971. Outside the islands the species occurs over the northern parts of the Humboldt Current. Two colonies were studied in detail (1970–1971). At the start of a season, males returned first to the colonies and defended a small territory. Copulation occurred without any elaborate ceremony and the female spent little time on land before laying. There was no fixed nest-site, even within a season, and birds moved their eggs considerable distances. This resulted in heavy egg losses. Younger birds bred later than older birds and laid longer but narrower eggs. The average incubation spell varied from four to five days at the extremes of the incubation period to 19 days in the middle. The average incubation and fledging periods were 60 and 167 days respectively. Pairs which lost an egg sometimes adopted the abandoned egg of another bird and successfully reared the chick. Most pairs nested in both seasons. Nesting success was extremely variable, both between years and between colonies. Between 1961 and 1971 at Punta Suarez, virtually no young were reared in four seasons. Even in 1970–71, where nesting success was good, some groups of birds deserted their eggs en masse whereas in neighbouring areas up to 80% of the pairs reared young. The main foods of the young were squid and fish. Birds did not moult wing and tail feathers at the breeding colonies, and about 50% retained some primaries for more than one season, suggesting that successful pairs had difficulty in fitting in a complete moult between breeding attempts. Old feathers were normally found among the inner primaries and at the next moult were preferentially replaced, though adjacent newer feathers were sometimes retained for another season. Some birds bred in their fourth years, but most not until a year or two older. Immatures were present at the colonies late in the breeding cycle, the youngest returning latest and remaining until the last young fledged. Survival of adults and young averaged at least 95% and 93% per annum over many years. Adults and young ringed in 1961 survived equally well. The significance of the timing of the return of immatures and of the large-scale desertion of eggs, apparently not due to food shortage or disturbance, is discussed.     

Sources of variation in breeding-ground fidelity of mallards (Anas platyrhynchos)

Generalizations used to support hypotheses about the evolutionof fidelity to breeding areas in birds include the tendencyfor fidelity to be greater in adult birds than in yearlings.In ducks, in contrast to most bird species, fidelity is thoughtto be greater among females than males. Researchers have suggestedthat fidelity in ducks is positively correlated with pond availability.However, most estimates of fidelity on which these inferences have been based represent functions of survival and recapture—resighting probabilities in addition to fidelity. We applied the modelingapproach developed by Burnham to recapture and band recoverydata of mallard ducks to test the above hypotheses about fidelity.We found little evidence of sex differences in adult philopatry,with females being slightly more philopatric than males inone study area, but not in a second study area. However, yearlingfemales were more philopatric than yearling males in both study areas. We found that adults were generally more philopatricthan yearlings. We could find no relationship between fidelityand pond availability. Our results, while partially supportingcurrent theory concerning sex and age differences in philopatry,suggest that adult male mallards are more philopatric thanonce thought, and we recommend that other generalizations aboutphilopatry be revisited with proper estimation techniques.     

Pre-breeding energy requirements: thyroid hormone, metabolism and the timing of reproduction in house sparrows Passer domesticus

We measured thyroid hormone (T3) levels and energy expenditure of pre-breeding house sparrows Passer domesticus in relation to the timing of breeding and reproductive success. The onset of reproduction was synchronised in two waves, separated by a three-week interval. On an annual basis, early breeders (birds that bred for the first time during the first wave) made significantly more breeding attempts, laid significantly more egg and raised 2.3 times more chicks to fledging than late breeders (birds that bred for the first time during the second wave). By the end of March, about one month before the first egg was laid in the population, plasma titres of testosterone in males and estradiol in females were still low and did not differ between early and late breeders. However, birds that subsequently bred early had higher titres of plasma triiodothyronine (T3) than birds that started to breed late. We show for the first time in a free-living bird population that Basal Metabolic Rate (BMR) is positively correlated with the plasma titre of T3. Differences in plasma T3 accounted for 48% of the inter-individual variation in BMR. Elevated T3 levels indicate that energy requirements increase prior to breeding. Although early breeding appears to be advantageous in terms of the number of offspring raised on an annual basis, the increased energy requirements prior to breeding are thought to delay the onset of reproduction in those birds that cannot afford the additional energy expenditure early in the season.