THE EFFECT OF AGE ON THE BREEDING BIOLOGY OF THE KITTTWAKE RISSA TRIDACTYLA

1. From 1954 to 1956 inclusive, the biology of individual marked Kittiwakes was studied at North Shields, Northumberland.
2. It was concluded that older Kittiwakes reacted to the breeding stimulus earlier, more intensively and with greater success than younger breeding birds.
3. Birds with previous breeding experience returned to the colony before birds breeding for the first time and these before non-breeders.
4. Before breeding started, birds which had bred previously spent more of their time at the colony than those about to breed for the first time.
5. Birds breeding for at least the second time laid the first egg 7·5 days earlier than those breeding for the first time.
6. Breeding started one day later for every four days the return to the colony was postponed.
7. Older breeding birds showed greater nest-site tenacity, laid larger clutches and had greater breeding success than younger birds.
8. The chicks in broods of two (but not of one) increased in weight more rapidly where the parents had previous experience.
9. Breeding Kittiwakes showed strong colony tenacity, but 24% of the marked non-breeding birds were subsequently seen in other colonies.
10. Over half the birds retained the same mate as in the previous year.     

Movements of Eiders Somateria mollissima on the east coast of Britain

Movement patterns, sex differences in natal dispersal and breeding dispersal, and interchange of birds between colonies were studied in the population of Eiders Somateria mollissima breeding on the east coast of Britain. First-winter Eiders reared at the Sands of Forvie, Grampian, remain at or close to the colony, while most adults move about 100 km south to winter on the Firths of Forth and Tay. A proportion of the Forvie population is sedentary. Eiders which breed in Northumberland either move north to winter on the Tay and Forth estuaries or remain close to their breeding areas. Eiders breeding in Fife are sedentary. Recoveries of British-ringed Eiders in Scandinavia indicate that some British-born males join the Baltic breeding population, probably by pairing with Scandinavian females wintering in Britain. There is extensive natal dispersal of males from Forvie, with more than twice as many Forvie-bred females as males returning to the colony to breed. The breeding dispersal of males is also twice that of females. Dispersal of males from the relatively sedentary Forvie wintering population is less than that from the breeding population. Previous work suggested that at Forvie sedentary birds nesting close to the estuary were genetically isolated from migratory ones nesting along the coast. This situation is less clear cut than had been supposed previously, with many migrants nesting close to the estuary. It is unlikely that the genetic differences between females nesting in different parts of the Forvie colony will remain stable in the long term, due to the natal and breeding dispersal of males.     

ECOLOGY OF THE COMMON BABBLER TURDOIDES CAUDATUS

Some aspects of the ecology and behaviour of the Common Babbler are described. The species resembles most other members of the genus Turdoides in being group territorial and exhibiting cooperative breeding. Females remain with their natal group until 6–9 months old and then disperse. Males, in most cases, remain indefinitely in their natal group so that these essentially constitute male clans. In half of the groups observed only a single pair bred per season, but in some large groups two pairs attempted to breed, one always being unsuccessful, and in other cases one male bred successively with two different females. Overall breeding success was highest for groups in which successive polygyny occurred. Male non-breeders fed the nestlings more often than female non-breeders which were sometimes driven off by the breeding female. A positive correlation was found between the number of non-breeders feeding the nestlings at day 9–11 and the size of the brood. This probably did not indicate a causal relationship. Breeders in large groups continued to moult while breeding, while those in small groups arrested their primary moult. This difference suggests that assistance from non-breeders may have had some effect in reducing the physical strain on the breeding pair. Observed overall survival of adult birds was 63% per annum, but that of the breeding males was 88% per annum. The population declined by 19% during the period of the study, suggesting that observed rates of survival may have been lower than normal.     

Conspecific reproductive success affects age of recruitment in a great cormorant, Phalacrocorax carbo sinensis, colony.

Few studies have addressed the proximate factors affecting the age at which individuals of long-lived bird species are recruited into the breeding population. We use capture-recapture analysis of resightings of 16 birth cohorts of colour-ringed great cormorants, Phalacrocorax carbo sinensis, in a Danish colony to assess the evidence for two hypotheses: conspecific attraction (earlier recruitment when the colony is large) and conspecific reproductive success (earlier recruitment following years of high breeding success). For both males and females, conspecific reproductive success was the most important covariate explaining the interannual variation in age of recruitment; colony size was also important for females. These covariates explained nearly 60% of the year-to-year variation for both sexes. The age of recruitment increased for cohorts born after 1990, and this increase was correlated with a decline in breeding success in the colony; we interpret this as an indirect and delayed density-dependent effect. Females were recruited earlier than males (mean age of recruitment for cohorts born before 1990: 2.98 years versus 3.53 years); the most plausible reason for this is a skewed sex ratio in favour of males in the adult population. Recruitment of males may thus, to some extent, be constrained by the availability of females. This study provides the first evidence that conspecific reproductive success can affect the age at which individual birds start to breed.     

THE ANNUAL RE-OCCUPATION OF BREEDING SITES BY THE FULMAR

The Fulmar has a long period at the breeding colony prior to egg-laying. The pattern of annual occupation and build-up in numbers has been examined in detail at Marsden, Co. Durham, at a colony in which over 100 eggs are laid annually (Order 3 of Fisher's classification). The re-occupation of the cliff starts in early November with an occasional visit by one or two birds. The main period of activity at the cliff is during the morning and, as the numbers build up, the diurnal period of occupation increases. By mid-December the first birds to arrive in the colony do so before dawn and the last to leave remain well after dark until near midnight. Almost throughout the pre-egg stage, the colony is deserted each night and re-occupied the next day and birds only stay regularly overnight just before egg-laying. A similar pattern of occupation occurs after breeding but in the reverse order. The numbers of birds at the colony in January and February exceed the breeding population and include many non-breeders. The non-breeders progressively decline in numbers until May when only the breeding birds remain with a few non-breeding birds. The daily variation in the numbers of birds at the cliff is influenced by the wind speed. In general, the birds leave the colony under freshening conditions and the number present at the colony can be interpreted in terms of the wind conditions over the last three days. It is suggested that the synchronised departures are primarily feeding trips, the birds using the strong winds to reach feeding areas, except that the departure just before egg-laying is linked to egg development and synchronised laying in the colony. Competition between Fulmars and Kittiwakes for nesting sites usually results in the Kittiwakes gaining the site. This is achieved by the Kittiwakes taking over the Fulmar sites during one of the latter's departures.     

Identification and breeding of yearling Piping Plovers

ABSTRACT Age of first breeding is an important life history trait. Many Piping Plovers (Charadrius melodus) do not breed as yearlings, but little information is available concerning the age of first breeding. From 2000 to 2006, we marked 991 chicks in three areas in Saskatchewan, Canada, and subsequently determined when 102 (49 females and 53 males) first bred. Females bred significantly earlier, on average, than males. More females (68%) bred as yearlings than did males (41%; P= 0.04), with most others first nesting when 2‐yr old (29% of females and 50% of males). As expected from differences between the sexes in age of first breeding, younger females were more likely to pair with older males than were younger males with older females. Chicks that hatched early in the breeding season did not breed at an earlier age than those that hatched late in the year. Unlike older birds, juvenile Piping Plovers do not replace flight feathers during their first winter. As a result, 18 of 27 yearlings (67%) had worn outer primaries, whereas only one of 123 (1%) older birds had worn primaries. In addition, whereas 20 of 24 yearlings (83%) retained a few buff‐tipped median coverts, none of 119 known older birds had such coverts. As a result, we were able to identify all yearlings by their worn primaries, buff‐tipped median wing coverts, or both. Wing lengths of yearling Piping Plovers were 3% shorter than those of older birds, presumably due to wear. Because there is no evidence of differences in adult survival rates between the sexes and breeding habitat is available, we speculate that fewer yearling males than females breed because primary wear may reduce the ability of yearling males to perform aerial breeding displays.     

THE TIMING OF BIRDS' BREEDING SEASONS

Examination of survival rdtes of nestlings and fledglings of some species show that there is a strong tendency for those young which are hatched earliest in the season to have the greatest chance of surviving to breed. Since natural selection so strongly favours parents who leave many surviving young, the question arises as to why other birds breed later than the date at which they could most successfully raise their young.
It is suggested that the food supply for the breeding females immediately prior to the breeding season may limit their ability to form eggs and the females may thus not be able to lay at the time which would result in young being in the nest at the best time for raising them, but as soon after this time as the female is able to produce her eggs. Not all species are likely to be prevented, by food shortage, from breeding at the best time for raising young and the groups of birds most likely to be affected are discussed.     

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.     

Is Divorce in Young Common Terns,Sterna hirundo,after Recruitment just a Question of Timing?

The probability of divorce in birds has been linked with age, breeding experience, reproductive output and synchrony in return. Here, we investigate the consequences of first breeding attempts in common terns for mating in the subsequent season. Nearly 20% of all first‐time breeders disappeared or skipped at least one season after recruitment. In 84 pairs, which consisted of at least one recruit and of which both partners returned to the colony, the divorce rate was 45%. We compared reproductive success, arrival dates, and asynchrony in arrival dates of pairs of the first breeding season against the second season, for both reunited and divorced pairs and males and females separately. First, in pairs of which both members came back to the colony, we found an increase of reproductive success most pronounced in males. In the second season reproductive success of divorced compared with reunited pairs was higher, as only divorced pairs significantly improved the number of fledglings, and again this relation was stronger in males. Secondly, females of reunited pairs arrived significantly earlier from the first to the second season and by far more days than their males. However, in divorced pairs former mates did not differ in the number of days they advance their arrivals. Finally, divorced males arrived on average 4 d earlier than their former mates, whereas divorced females arrived 5 d later compared with their former mates of the recruitment season. Contradictory to nearly all other divorce studies in birds so far, we found a clear fitness gain in divorced males. We suggest that the improvement in reproductive success of young males stems from a side‐effect of the birds’ quality and ability to reach the breeding site in appropriate time and earlier as potential competitors. In long‐lived bird species the heterogeneity among young individuals in the timing of arrival at the colony seems to explain why former recruit‐pairs reunite or split. For young males we suggest as best explanation of divorce that they profit from ‘pushing for an empty chair’, while females seem to profit from their choosiness and may actively decide between former and other mates.     

THE BREEDING BIOLOGY OF THE GANNET SULA BASSANA ON THE BASS ROCK, SCOTLAND

The Bass colony is increasing—in 1962 there were 5,350–5,700 breeding pairs; 1,340–1,430 pairs of non-breeders with nests or sites (mainly pairs in their season before first breeding) and 2,000–2,500 “club” birds without nest or site. 15% of nests were occupied by both birds of a pair at the time of the count. Oldest males return to the colony in January, followed by experienced females, considerably later by young adult-plumaged birds, immature birds later still, and the few one year-olds that return usually not until May and June. Mid-cliff sites are the first to be re-colonized each year. Gannets usually breed in their fifth year and there is some evidence that females breed earlier than males. The characteristics of Gannet nests and sites are described. Nests function in raising the egg and young above the morass of the breeding colony, and reach a density of about 2.3 per square metre. Nests are demolished and their positions changed more often than might be suspected. The extremely strong social tendency which causes Gannets to establish their sites amongst or very close to existing breeders probably is the factor ensuring high density and this contributes to synchronization of laying. Egg laying is analysed. Experienced pairs forming an isolated group of 20 nests began laying later and showed less synchronization than two other groups of the same size but from the middle of a dense mass, probably due to the greater social stimulation experienced by the non-isolated groups. The date for first and median eggs was also earlier in larger than smaller groups in the same year. The effect of density as distinct from group size is also discussed. Early eggs are mainly laid on cliff or cliff-edge sites and in large nests. Different groups within the colony produced the median egg within 2–3 days of the end of April each year. In the fullest documented group the mean date was also constant from year to year and closely approached the median, implying a considerable degree of synchronization within the gannetry as a whole. Laying in the observation colony became progressively earlier in successive years, probably due to recovery from previous disturbance. Nevertheless, individual females tended to lay in a fixed position each year with relation to the mean for the group. Increasing age of the female causes earlier egg laying and heavier eggs for up to at least five years. It is suggested that the survival value of seasonally synchronized laying in the Gannet is maximum utilization of a seasonally dependable and abundant food supply for the production of young with the optimal chance of post-fledging survival. The spread of laying acts as an insurance against possible adverse conditions. There is a considerable reserve of unutilized breeding capability within the colony (adult non-breeders, a pre-maturity period longer than physiologically necessary for egg production, and a one-egg clutch when in fact two young can easily be reared). The mean of 393 Gannet eggs was 104.5 gm. (range 81–130). Eggs constitute about 3.4% of adult female weight and lose 9–13% in weight during incubation. Replacement laying of the invariable one-egg clutch takes 6–32 days. The mean incubation period was 43.6 days. Male incubation spells averaged 35.6 hours; female's 32.0 hours. Copulation ceases immediately after egg laying. During three seasons, 82% of eggs laid in the observation colony hatched. Inexperienced pairs hatched 62.6% of eggs laid; experienced pairs hatched 86%. Some of the processes of incubation and chick rearing depend on the maturation of innate abilities and not on experience Inexperienced breeders do not seem inferior to experienced ones in finding enough food for their young. Parental care of the new chick is described; the pterylosis of the chick is figured. A summarized account of plumage development is given. Food fish and chick feeding are described. The average frequency of feeds throughout a continuous two-day watch was 2.7 feeding bouts per chick per day. Adult fishing trips usually took 7–13 hours and the estimated fishing range is over 100 miles, and possibly up to 400 miles, from the breeding colony. Despite this, 15% of daylight hours are spent by the pair together at the nest in addition to the constant guarding of the chick by one or other. Gannet young have a very compressed growth period compared with boobies and fledge at 3,100–4,100 gm., after a steady growth uninterrupted by periods of starvation or arrested development after an average 90 days. 92.3% of all eggs which hatched in 500 nests in the observation colony during three years of the study gave fledged young. Excluding inexperienced birds, there was no difference in the fledging success of eggs laid at different dates in the breeding season, in accordance with the proved abund- ance of food. However, post-fledging survival is probably higher among young fledging at the peak period (first half of September) than later and the few relevant ringing returns tend to support this. Breeding success at the small colony at Bempton was less compared with groups from the Bass for the years 1961–3. Causes of chick loss before and during fledging are discussed. They are unimportant compared with the great mortality in the first year of life after fledging. The adaptive significance of black plumage in the juvenile Gannet probably lies in reduced attack-releasing qualities of such plumage on the male parent. The Gannet alone in the Sulidae produces young which leave the nest with large fat deposits, and which are not fed at all by the parents after fledging. This is possibly another result of adaptation to using a seasonally abundant food supply to the maximum. The present Gannet population increase is discussed in relation to cessation of human predation, the possible impetus given by a temporary but large increase in pelagic fish during the war, but also the overall steady downward trend in fishing returns since the early part of this century. One cannot explain the steady and considerable rise in Gannet numbers only in terms of increased food supply. The fact that, at a time of population expansion and obviously favourable conditions, Gannets are still far from utilizing their full recruiting powers, needs investigating further. It may be partly due to the relative slowness of evolutionary change in a long-lived species with slow population turn-over, if the Gannet has evolved its characteristics in response to an environment different from the present one.     

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.     

Philopatry and colony fidelity of Shags Phalacrocorax aristotelis on the east coast of Britain

Shag Phalacrocorax aristotelis colonies from the Firth of Forth south to the Fame Islands (SE Scotland and NE England) were surveyed in 1982 and 1983 for nesting Shags ringed as chicks or adults in the previous 20 years. On average, 5% of 863 Shags ringed as chicks were breeding away from their natal colony, and only 1% of 401 marked adults had left their breeding colony. The proportion p of chicks that settled at or beyond distance D from the natal colony followed the relationship p = 0.263D-^0.771, implying that more Shags moved 10 km or beyond to breed than expected from diffusion models or a constant-rate dispersal model; adjusted for birds that settled outside the study area, the relationship was p = 0.314D^-0771. There was no effect of colony size on emigration rate, but females showed a greater tendency to settle away from the natal colony than did males. At the natal colony, more males (69%) than females (47%) nested within 300 m of their birthplace. Two mechanisms appear to determine where a Shag settles to breed: a navigational one governing the return to the natal colony, plus the competitive process of nest-site establishment.     

The disperser dilemma in cooperatively breeding birds

In most cooperatively breeding birds, individuals do not breed with their natal group members. In order to breed, they have either to disperse into another group or wait for an opposite-sex individual to join their group. In most of these species, females disperse more than males. We develop a dynamic game-theoretic model to account for this asymmetry. When males are physically larger/heavier than females, this allows them to effectively welcome female immigrants into their natal group and overcome the local females' opposition more than vice versa. The model further assumes that the dispersal decision is not confined to a restricted time window, but is rather based on acquired information and responsive to opportunities. The model predicts that (i) females disperse more than males, and (ii) females are willing to tolerate more risks in dispersal than do males. The latter prediction is supported inter alia by the fact that in many cooperatively breeding birds, females disperse at a younger age, and further away from their natal group as compared to dispersing males.     

THE EVOLUTION OF FEMALE BODY SIZE IN RED-WINGED BLACKBIRDS: THE EFFECTS OF TIMING OF BREEDING,SOCIAL COMPETITION,AND REPRODUCTIVE ENERGETICS

We examined opposing selective forces on female body size in the sexually dimorphic red-winged blackbird: social competition favoring larger females, and energetic advantages favoring smaller females. Downhower proposed that selection might drive female birds to be smaller than the optimum for survival, if smaller females were able to exceed their energetic requirements for self-maintenance earlier in the season and therefore breed earlier. Since in most birds the earliest breeders fledge the most young, this could favor the evolution of smaller female size, and therefore contribute to the magnitude of sexual size dimorphism in these birds. We tested this hypothesis in 1987 and 1988 by comparing the size and breeding date of female red-winged blackbirds. Consistent with our preditions, early-nesting females had much higher nesting success, but contrary to prediction, larger females bred earlier. We then examined the effects of female size on competition. If large females have an advantage in social competition, and if competition influences breeding date and reproductive success, then larger females might breed earlier. Primary females, the first females to arrive and nest on a territory, were more aggressive than lower ranked females; more aggressive females settled on better territories and laid earlier than less aggressive females; and larger females were more aggressive. Social competition between females may therefore favor large females. Finally, we tested the prediction that selection favoring large females might be limited by energetic constraints on large females. We found that large females had less fat than small females during breeding, and that the levels of fat that females of a given size carried affected breeding date and egg size. Therefore, social competition may favor large females, but reproductive energetics favoring smaller females may constrain selection for large female body size.     

Breeding site fidelity and natal philopatry in the Redshank Tringa totanus

This paper presents the results of a study carried out on breeding Redshank in the Ribble Marshes, Lancashire, England.
Redshank tend to return to the same breeding area year after year. There was no detectable sex bias in return rates. Experienced birds were more site faithful than inexperienced birds, with previously successful birds exhibiting the highest degree of breeding site fidelity. Older, more experienced birds were more successful at hatching eggs than inexperienced birds.
Breeding dispersal was the same both within and between years. Faithful pairs and males nesting with a new mate dispersed significantly shorter distances than females nesting with a new partner. Dispersal distances in female Redshank were affected by breeding success: unsuccessful females, nesting with a new mate, dispersed significantly farther than successful females. A pair's breeding success influenced the following year's mate fidelity. However, other factors such as overwintering survival and date of return may also have influenced mate fidelity.
Redshank were highly faithful to their natal area; a high proportion of birds that survived post-fledging mortality returned to breed in their area of birth. No sex bias in natal dispersal was detected. Approximately 50% of Redshank breed in their first year of life.     

Temporal and spatial variation in dispersal in the greater flamingo (Phoenicopterus ruber roseus)

We studied movements of individually marked greater flamingos (Phoenicopterus ruber roseus) born in the Camargue, southern France, between their two most important breeding colonies in the western Mediterranean (Camargue and Fuente de Piedra, Spain) from 1986 to 1992. The two sites differ in the frequency with which they offer suitable conditions for breeding. Flamingos have bred each year in the Camargue since 1974, but in only 12 of the past 22 years at Fuente de Piedra. Higher colony fidelity is thus expected in the less variable environment (Camargue), but if dispersal occurs competition might be an important factor causing this dispersal. Following years during which breeding birds in the Camrgue were disturbed (1988 and 1990) a higher proportion of adults changed colonies between breeding attempts (= breeding dispersal, 12.4%), while only 0.4% of flamingos breeding in the Camargue dispersed in the other years. As expected, flamingos breeding at Fuente de Piedra showed a higher rate of breeding dispersal (8.14%). No differences were observed between males and females. The importance of breeding failure as a factor causing breeding dispersal in flamingos was also confirmed by the movements of individual birds. The proportion of young flamingos that moved from their natal colony to start breeding at Fuente de Piedra (= natal dispersal) was independent of sex and age, but increased when breeding access to the Camargue colony was more difficult. However, natal dispersal was also higher in 1988 and 1990 (40.5%) than in the remaining years (1.2%), as was breeding dispersal. We discuss possible ways in which the increased natal dispersal among inexperienced birds could be linked with the increased breeding dispersal of adults in the same year.     

Sexual competition in an inbreeding social spider,Stegodyphus dumicola (Araneae: Eresidae)

Summary Sexual competition is shown to occur in the social spiderStegodyphus dumicola (Eresidae). While the secondary sex ratio inS. dumicola was female-biased, the overall operational sex ratio (numbers of breeding males to breeding females over the season) showed no strong female bias. Males matured before females and had a shorter lifespan than the females. Mating took place in the natal colony. Males fought over access to the few mature females available early in the reproductive season, but females appeared to control the duration of mating. Later in the season, some adults of both sexes dispersed alone to breed elsewhere. We conclude that different rates of maturation between the sexes within a colony provide the opportunity for females that mature early in the season to be choosy in selecting a mate and this forces males to compete. Early reproduction may be beneficial for both females and males, because the offspring of females that reproduce early may have a competitive advantage over later (and smaller) offspring in the colony.     

Sex, age, experience and condition as factors affecting arrival date in prospecting common terns, Sterna hirundo

Several studies have shown that seabird colonies consist to a large extent of young nonbreeders (prospectors). These individuals appear at the colony later in the season than established breeders. The reasons for this late arrival have remained unclear in most cases, mainly because of technical difficulties in collecting sufficient data from nonbreeding individuals. We used a novel transponder system to identify remotely the members of a common tern colony, including nonbreeders, during eight breeding seasons and we combined the system with automatic balances. Ninety-two per cent of prospectors returned for the first time when 2 years old and 88.9% of recruits to the breeding population had spent at least one previous season at the colony as prospectors. In both sexes, most individuals prospected for one season, but more males than females prospected for more than one season, although a higher proportion of females started breeding without a previous prospecting phase. Terns arrived earlier in the season the older they were and the more experience of the colony they had, but experience proved to be more important than age. Prospectors gained about 3 weeks with a previous prospecting season whereas an additional year of age allowed birds to arrive only about 6 days earlier. Prospectors returning later in the season arrived with lower body masses. Males on average arrived earlier at the colony than females. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

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.     

Hatching date influences age at first reproduction in the black-headed gull

In long-lived colonial birds, age at recruitment is an important life-history character. Variation in this parameter may reflect differences in several factors, including competitive ability and breeding strategies. Further, these differences may be due to timing of hatching (for instance through differences in competitive ability). We investigated the age of first-time breeders in relation to hatching date in a black-headed gull Larus ridibundus colony situated in central France, from 1979 to 1993. Age at first breeding was estimated for four groups of individuals (total n=550) according to their hatching date, using a recent capture-recapture methodology which allowed us to estimate recruitment rate without the limiting assumptions of methods relying on simple return rates. The age at first breeding was negatively correlated with the hatching date of individuals: individuals hatched earlier in the season started breeding at a younger age than individuals born later. Proportionally more 2-year-old late-hatched individuals were seen breeding on small peripheral colonies than young early-hatched individuals. This difference disappeared after age 3 years. These results strongly suggest that individuals hatched late in the season start to breed on peripheral colonies before recruiting to their natal colony. A difference of few weeks in hatching date has consequences which can last for several years.