Age-dependent prospecting and recruitment to a breeding colony of Short-tailed Shearwaters Puffinus tenuirostris

In a small colony of Short-tailed Shearwaters Puffinus tenuirostris , monitored annually since 1947, many of the young ringed as nestlings returned to their natal colony for one or more years before breeding for the first time. These 'prospectors' were first recorded at 4.1 ± 0.1 (range 2–7) years old; they started breeding at 7.0 ± 0.2 (4–14) years old. Overall, 31% of all prospectors bred on their natal island with no significant annual variation in this proportion. However, 8% of all natal recruits started breeding without having been recorded prospecting and bred first when significantly younger (5.6 ± 0.4 years old) than those first recorded prospecting. Shearwaters which subsequently bred at their natal colony started prospecting when significantly older (4.3 ± 0.1) than those that did not breed at their natal colony (4.0 ± 0.1 years). Non-natal recruits spent significantly less time prospecting (1.0 ± 0.1 years) than prospectors born on the island (1.9 ± 0.1 years). The shearwaters appeared to become progressively more faithful to the colony during their prospecting period.     

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.     

Determinants of local recruitment in a growing colony of Audouin's gull

1. Local recruitment of Audouin's gull ( Larus audouinii Payraudeau) was studied between 1988 and 1997 at the Ebro Delta colony (north-western Mediterranean). Since its establishment in 1981, the colony has dramatically grown to include, in 1997, 65% of the total world population. Several hypotheses were tested, involving the effects of a badger predatory event in 1994, and sex, age and cohort (year of birth) on recruitment.
2. Results supported the prediction that colony size influenced recruitment: the probability for any individual to have previously bred increased throughout the study, together with colony size. At the end of the study, 90% of breeders aged 4 years had already been recruited at age 3, the age of first reproduction by Audouin's gulls. As expected by the dramatic increase of breeding numbers, most local recruitment occurred at very young ages, especially when compared with other Laridae.
3. Neither food availability nor reproductive success affected recruitment. Recruitment was not affected by high nest predation by the badger, although after the event, the proportion of Ebro Delta birds nesting on the nearby Columbretes Islands tripled.
4. Probability of first reproduction depended on age: it was the highest at ages 3 and 4, and then decreased sharply with age to stabilize beyond age 6 to a value depending on the year and cohort but always very low (< 5%). Cohort and sex did not influence local recruitment.
5. Annual resighting rates ranged between 35% and 82%, and were higher for females. This may represent a sex-dependent suspension of breeding, probably as a trade-off between early recruitment and future survival.     

Interrelationships between breeding frequency,timing and outcome in King Penguins Aptenodytes patagonicus: are King Penguins biennial breeders?

King Penguin Aptenodytes patagonicus chick growth is interrupted by a winter fast which extends the length of the breeding cycle (laying to chick independence) to 14–16 months, so that continuous annual successful breeding appears to be impossible. The 3–month laying period imposes further constraints with respect to timing of breeding attempts in relation to the onset of fasting. By the frequent resighting of individually marked birds at Marion Island, we examined the frequency of breeding and the relationships between timing, outcome and frequency in the same and in consecutive years. A total of 3101 adult King Penguins were banded between 1984 and 1991, yielding continuous breeding histories spanning a maximum of 5 years. Most penguins attempted to breed in consecutive years, although the likelihood of taking a year off increased with the number of consecutive attempts. In any one season, about 19% of potentially breeding adults did not breed. Early breeders were more likely to succeed than late breeders and comprised 84% of breeding attempts of known timing. Successful birds in one season usually bred late in the following season, whereas failed attempts were usually followed by an early attempt. Non-breeding was more likely to occur after a successful outcome than a failed one, and after a year off 93% of birds bred early. It is incorrect to refer to the King Penguin as a biennial breeder.     

THE BREEDING OF THE GREATER FLAMINGO AND GREAT WHITE PELICAN IN EAST AFRICA

This paper brings together observations on the breeding of the Greater Flamingo Phoenicopterus ruber and Great White Pelican Pelecanus onocrotalus, mainly at Lake Elmenteita, Kenya, 1951–1971. The Greater Flamingo bred at Lakes Elmenteita and Nakuru in 11/21 observed years and at Lakes Natron and Magadi in 5/12 observed years. On average, it breeds about every second year, but a succession of breeding years is followed by several years in which no breeding occurs. A history of 21 years' breeding at Lakes Nakuru and Elmenteita is given. At Elmenteita three sites have been used, the main site in every breeding year, the others less often. The number of pairs breeding in any year has varied from 500–9,250, but in 1968 flamingos bred three times, involving perhaps 8,500 pairs which made about 15,700 nests, some pairs perhaps laying twice or even thrice in a year. Losses of eggs (38.2% overall) were caused by rising water (16.2%), competition for nest space with Great White Pelicans (6.9%, after 1968 only), human interference (3.5%), Marabou Stork predation (1.8%) and other natural causes (9.8%). Losses among chicks totalled 68.3% overall and were mainly due to Marabou Storks (36.5%), undiagnosed disease (8.6%), and rising water (6.6%). Disease caused serious loss only in 1966, and after 1968 losses from Marabous rose from 2.7% to 76.5%, resulting in an increase in overall mortality from 48.7 to 92%. This was perhaps associated with the establishment of a fish factory at Lake Naivasha. When attacking flamingo colonies Marabous did not actually eat many eggs or chicks, but simply caused wholesale desertion by alarming the flamingos. In 1968 total desertion of a colony of 4,500 pairs was caused between 18 and 26 March by a maximum of 17 Marabous, and similar wholesale desertion was caused in later years. The overall breeding success among Greater Flamingos at Elmenteita was about 19% of eggs laid, but without the excessive post-1968 Marabou predation would have been about 30%. At such a rate Greater Flamingos require at least 24 years of adult life to replace themselves, but if the mortality caused by Marabous since 1968 continues they will require about 58 years, and the population will inevitably decline. Breeding success at Lakes Magadi and Natron has been higher, about 44% of eggs laid; but figures available are much more approximate than at Elmenteita. Some new data on display, nest-site selection, laying dates, clutch-size, hatching and creche behaviour are given for the Greater Flamingo. The Great White Pelican bred at Lake Elmenteita from 1968 to 1971 without a break, some birds laying in every month, but with reduced laying November–December. They bred on the same islands as, and in association with the Greater Flamingo, and caused heavy losses among the latter, not through aggressiveness, but simply because of their superior size and weight. Although food supply must ultimately have controlled the pelican's ability to breed, an adequate food supply was available for 6 years before they did and continued after they had ceased. Their breeding was finally triggered by the Greater Flamingo colonies, with which the pelicans associated. When a flamingo colony was deserted because of Marabou Storks the pelicans, unafraid themselves of the Marabous, also deserted. They also associated with, and wiped out, a colony of Sacred Ibis. From July 1968 to June 1969 about 2,600 pairs of pelicans bred at Elmenteita, rearing about 2,200 young to the flying stage. The breeding colony apparently comprised most of the adults from Lake Nakuru and Lake Naivasha, the main feeding areas. From July 1968 to January 1971 certainly 7,200 and probably 8,000 pairs of pelicans bred at Elmenteita. Some pairs may have bred twice or thrice in this period. Breeding ceased suddenly in January 1971, eggs, and small and large young being alike abandoned for no established reason, although food supply was certainly still plentiful. Additional information on pair formation, incubation and fledging periods, nest-relief, etc. is given. The best available record of the incubation period is 35–36 days. Nest relief takes place on average about once every 48 hrs, and is dependent on thermal activity enabling the pelicans to soar. At Elmenteita large young ate quantities of putrefying matter, including the corpses of other young pelicans. They also ate living young hatching from eggs, and up to 14 days old. Touch probably plays an important part in helping them to locate possible food in opaque water.     

Pair bond and breeding success in Blue Tits Parus caeruleus and Great Tits Parus major

Data from 939 nests of the Blue Tit Parus caeruleus and 1008 nests of the Great Tit P. major from nestboxes provided in superabundance in mixed forest study sites between 1976 and 2001 were analysed to examine the effects of mate retention on breeding success and the relationship between mate fidelity and site fidelity. Most birds retained their former partner (76% in Great Tits and 65% in Blue Tits). The probability of a pair divorcing was affected by male age in Great Tits, divorce being more likely in pairs with first‐year males. Great Tit pairs breeding together for a second season bred earlier, but had no higher breeding success than pairs breeding together for the first time. In Blue Tits laying date and start of incubation tended to be earlier in pairs breeding together for a second season, but hatching and fledging dates were not earlier than in other pairs. Great Tit pairs breeding together for two consecutive seasons bred earlier in the second season than in the first, but breeding success did not differ significantly between years. In both species, breeding performance did not differ between pairs that divorced after a season and pairs that stayed together. Thus breeding success did not determine whether a pair divorced or bred together again. Neither Blue Tits nor Great Tits improved their breeding performance through divorce. Blue Tit females even had fewer fledglings in the year after divorce than in the year before. Mate retention affected breeding site fidelity. Blue Tit females had greater breeding dispersal distances between consecutive years when re‐mating than when breeding again with the same mate. In Great Tits both males and females dispersed more when re‐mating than when retaining the former partner, suggesting that mate retention increased the chance of retaining the breeding site. In both species, breeding dispersal distances did not differ between pairs that divorced and pairs in which one mate disappeared. Because no major advantage of mate retention was evident, we suggest that mate retention evolved under different conditions than those found in study sites with high breeding densities and a superabundance of artificial nesting sites.     

Measuring immigration and philopatry in seabirds; recruitment to Black-legged Kittiwake colonies

The degree of philopatry in two new Black-legged Kittiwake Rissa tridactyla colonies in northeast England was studied for 36 and 16 successive years, respectively. There was a tendency in both colonies for the number of recruits to the breeding group to increase with the age (and size) of the colony, but by an average of only one additional recruit each year. In the two new colonies, it was 7 and 9 years respectively before the first young birds bred in their natal colony. By that time, over 100 immigrants had recruited and bred in each colony. Thereafter, new philopatric birds were recorded annually. But these formed only an average of 23% of the recruits during 36 years (1955–90) at North Shields, and 4.2% over 16 years (1991–2006) at Coquet Island. In every year at both colonies the number of new immigrants exceeded the number of philopatric recruits. Males formed 82% of philopatric birds. The low degree of philopatry in the Kittiwake will result in appreciable gene flow between colonies and this is consistent with little geographic variation in the species. Each immigrant has to make a choice of which colony to enter, but the criteria for this selection are not understood. It is argued that many measures of philopatry in birds are biased towards high values because of the greater ease in identifying philopatric individuals. The generalization that there is a high degree of philopatry in colonial seabird species is not justified, as there is considerable variation between species, and possibly between colonies. More estimates based on unbiased data are needed.     

Annual variation in breeding biology of gentoo penguins, Pygoscelis papua, at Bird Island, South Georgia

The breeding biology of the gentoo penguin, Pygoscelis papua , was studied over a three-year period (1986–1988) at Bird Island, South Georgia, with particular reference to birds of known age or breeding experience. Laying date varied significantly between all three years, being three weeks later in 1987, when the breeding population decreased markedly. Factors involved in the timing of breeding are discussed. Within years egg-laying was highly synchronous: 95% of clutches were initiated in 14·5 days or less. The incubation period was 35 days and the laying interval, between the two eggs, 3·3–3·4 days. Chicks creched when 25–30 days old, and this varied between years, possibly related to food supply and chick growth. Chicks left the colony for the first time between 75 and 85 days of age. The breeding population at Bird Island decreased by 20% and increased by 84% in successive years during the study period. Breeding success (chicks fledged per egg laid) varied between 0·33 and 0·65 within colonies, but for the whole island was very consistent over the three years: 0·45, 0·51 and 0·47. Overall, colony differences were not correlated between years. Disturbance from Antarctic fur seals, Arctocephalus gazella , is suggested as the cause of consistently lower breeding success at one colony. Mean egg weight varied annually, and with age of the breeding bird, nest location and, in one year, with laying date. Young, first-time breeders laid smaller eggs and had lower breeding success compared to older, experienced birds, similar to other seabirds. However, they differed from other species in laying on average earlier than older birds. The relationship between age, egg weight, laying date and breeding success is discussed in relation to predation and seasonal food supply.     

Two sibling species sympatrically breeding: a new conservation concern for the critically endangered Balearic shearwater

Two Puffinus shearwater species are endemic to the Mediterranean region: the Yelkouan shearwater P.␣yelkouan breeding in the central and eastern, and the critically endangered Balearic shearwater P.␣mauretanicus in the western, exclusively breeding at the Balearic archipelago. All individuals caught in Minorca, a Balearic breeding site, showed phenotypic traits of Yelkouan shearwaters. Ten birds from that colony were sampled and genetically analysed. A fragment of 857 base pairs of the cytochrome b was sequenced and compared to published sequences of both species obtained from GenBank. A Neighbor-Joining (NJ) reconstruction distinguished two clades with high bootstrap support, showing that both species bred sympatrically in that colony. This result was unexpected since shearwaters do not usually form mixed colonies. New conservation concerns arise also from our study: at first, breeding population of Yelkouan shearwaters is even smaller than previously thought; additionally, other potential threats emerge, such as hybridization and ecological competition.     

Demography of Bouteloua gracilis in a mixed prairie: analysis of genets and individuals

1 We analysed a data set containing the locations and sizes of individual plants from five 1-m² permanent quadrats that had been mapped annually from 1934 to 1972. We digitized each map and used geographical information software to evaluate annual recruitment and mortality, age structure, survivorship and longevity of genets and individuals of Bouteloua gracilis .
2 Ninety-one per cent of the years between 1938 and 1972 had at least one seedling recruited. An average of 10.3 genets died annually, although the variability among years was high (CV = 70%). The number of seedlings recruited each year was significantly correlated with the number of genets dying. Most genets died before age 10 and the average life span was 3.7 years, although some genets were much longer lived (> 38 years).
3 The average number of individuals per genet was 1.3. Excluding seedlings, 56% of the genets were represented by a single individual. The annual number of individuals in the 5 m² ranged from 29 to 278. Average mortality of individuals was 25.8 per year and the variability among years was high (CV = 91%).
4 The plant community appeared to be relatively stable and openings created by mortality were recolonized by recruitment. Climatic variability did not have a large influence on demography.     

Body size and determinants of laying date variation in the Snow Petrel Pagodroma nivea

We studied several determinants of laying date variation and the relationship between laying date and reproductive success in the Snow Petrel Pagodroma nivea . The effects of female body size and condition, year, individual laying period, colony size, mate fidelity, previous reproductive success, and duration of the pre-laying exodus on laying date, were investigated during a 3-year study. The average laying date was 4 December. The laying period was compressed into 10–16 days and was very constant from year to year, both for the population as a whole and for individual females. The laying period of individual females varied from year to year on average by less than one day. Body size explained 24% of the variation in laying dates, with large females laying their egg later than small ones. Laying in large colonies occurred c. 2 days later than in small colonies, mainly because a higher proportion of large females bred in large colonies. There was no effect of mate fidelity, age, body condition and previous reproductive success on laying date, but the duration of the pre-laying exodus was strongly correlated with laying date. Smaller females had shorter pre-laying exodus (c. 17.7 days) than larger ones (c. 20.4 days). During the three years of the study reproductive success either increased, decreased or did not vary with laying date. Although body size is not maintained by selection on laying date alone, the genetic body size component of this species suggests that balancing selection on body size may act through laying date.     

THE FOUNDING OF A NEW POPULATION OF DARWIN'S FINCHES

We report the natural colonization of the small Galápagos island Daphne Major by the large ground finch (Geospiza magnirostris). Immigrants of this species were present in every year of a 22-yr study, 1973–1994. Typically they arrived after a breeding season and left at the beginning of the next one. Geospiza magnirostris bred on the island for the first time in the exceptionally wet El Niño year of 1982–1983, and bred in all subsequent years except drought years. In agreement with theoretical expectations the frequency of inbreeding was unusually high. Pronounced fluctuating asymmetry in tarsus length, together with slightly reduced breeding success of inbreeding pairs, suggests a low level of inbreeding depression. Despite this, the population increased from 5 breeding individuals in 1983 to 20 breeding individuals in 1992, and probably more than twice that number in 1993, largely through recruitment of locally born birds.     

A 13-YEAR POPULATION STUDY OF THE BLACK EAGLES IN THEMATOPOS,RHODESIA, 1964–1976

Gargett, V. 1977. A 13-year population study of the Black Eagles in the Matopos, Rhodesia, 1964–1976. Ostrich 48:17-27.

The resident black Eagle Aquila verreauxi population in approximately 620 km² of the Matopos, Rhodesia, underwent changes from 1971 and appeared less stable than in the first five years. Six new territories were established in the National Park; two territories in Tribal Trust Land were abandoned; one territory in the National Park vacant for five years and one in Tribal Land vacant for at least 11 years were reoccupied. breeding data covering 13 years confirm the five-year findings. In 652 pair-years 442 breeding attempts were made with 339 young eagles f edging: a reproductive success rate of 0,52 young per pair per year. On average 68 % of the population bred every year, each pair attempted breeding in two years out of three, and one attempt in four was unsuccessful. Annual and individual variations in breeding performance were considerable, from 46% to 89% of the population breeding, and from two successes in six attempts for one pair to 12 successes in 12 attempts for two pairs. Over 13 years the percentage of the population breeding annually followed the form of a sine curve with a ten year period.

Fourteen factors that might affect annual and/or individual performance are considered. Below average rainfall years were followed by above average breeding; and generally fewer clutches were laid early after heavy rainfall in the three months preceding laying. Clutches laid late were less successful than those laid early. Breeding was affected by the proximity and intervisibility of nests, the previous year's performance and intraspecific disturbance. Appropriation of nests by other species prevented breeding and no new pair bred while establishing or re-establishing its territory. Building a new nest was followed by laying in the same season only if the nest was completed by mid-May, the peak laying period, building or partly building two nests in one season did not prevent breeding in the following year. At least 76% of clutches were c/2. The causes of two eaglets' deaths were ascertained and the remains of two adults were found. Pairs with territories in Tribal Trust Land had a significantly lower breeding performance than pairs in the protected National Park. Only traces of pesticide residues were found in four eggs. Observers' visits did not affect reproductive success.     

Nestling obesity in procellariiform seabirds: temporal and stochastic variation in provisioning and growth of short-tailed shearwaters Puffinus tenuirostris

Procellariiform seabirds such as short-tailed shearwaters Puffinus tenuirostris accumulate large quantities of lipid during the nestling period. The functional significance of this pattern of development remains unclear, but has been related both to temporal variation in feeding conditions around the colony and to stochastic variation in the foraging success of individual parents. This paper examines temporal and age-specific variation in the pattern of food delivery to nestling short-tailed shearwaters, which have one of the lowest provisioning rates of any procellariiforms and are known to experience occasional long intervals between feeds. We assess whether variation in the provisioning rates of chicks was associated primarily with temporal variation in food delivery at the level of the colony or with stochastic variation in food delivery at the level of the individual. We then discuss this variability in the context of nestling obesity. For all but the youngest chick age-classes, individual meals delivered by adults averaged 141 g, which was 25% of adult body mass. The proportion of chicks fed each night was low (49%) and highly variable (coefficient of variation = 82%), which means that occasional long intervals between feeds would be expected to arise simply by chance. In keeping with this, intervals between feeding events for individual chicks followed a negative exponential distribution with a mean of 2 nights and a maximum interval of 13 nights. There was significant temporal variation in food delivery, but deviations from expected values for both feeding frequency and meal size were restricted to a small number of nights, included values both higher and lower than expected and did not persist for more than 2 nights in succession. These data suggest that even among those species with very low feeding frequencies and occasional long intervals between feeds, nestling obesity in Procellariiformes should be regarded as a response to chronic stochastic variability in food delivery at the level of the individual chick rather than as insurance against sporadic temporal variation at the level of the colony. Received: 3 March 1997 / Accepted: 10 May 1997     

BREEDING PERFORMANCE OF PUFFINS FRATERCULA ARCTICA IN RELATION TO NEST DENSITY, LAYING DATE AND YEAR

The paper presents data on the breeding and predation of Puffins in two areas of different nest density within a single colony on Dun, St Kilda group, Outer Hebrides in 1973-78.
Within a season birds laying early had a slightly higher nesting success than birds laying late, but laying date had little influence on the peak and fledging weights of young. The main disadvantage in late laying was a reduced chance of relaying if the first egg was lost.
Breeding success and chick weights varied from year to year. The 1974 season was the least successful with the lowest nesting success, lowest frequency of feeds, lowest calorific value of feeds, lightest chicks and slowest growth. Overall breeding performance was not related to the annual mean laying dates.
In all years pairs nesting in the area of high nest density did better than pairs nesting at low density. The effect is attributed to differential predation and disturbance by predatory gulls. At least 4.2% of adult Puffins breeding in the area of low burrow density were killed by gulls each breeding season; this is higher than the total annual mortality found in three other studies. Only 0.9% of adults from the high density area were found killed. The subpopulation in the low density area cannot survive without much immigration, yet there is no evidence that this happens.     

Individual breeding decisions and long-term reproductive strategy in the King Penguin Aptenodytes patagonicus

Although studied for 35 years, knowledge of the reproductive biology of the King Penguin Aptenodytes patagonicus remains incomplete. The chick requires more than 12 months of care, which extends the breeding cycle, including moult, to more than one year, i.e. the King Penguin is neither annual nor biennial. In an attempt to resolve ambiguities in the literature and to elucidate the long-term breeding strategy of the species, we studied breeciing frequency at the individual level, considering the decision to breed in relation to breeding history over the previous few years. Although adult birds attempted to breed annually (0.83 breeding attempts per year), successful rearing occurred, at best, every two years only (maximum of 0.41 fledged chick per pair). Comparing successive years, the number of breeding birds in the colony was stable but the number of fledged chicks varied from 29 to 278 over eight years. These results suggest that King Penguins adopt (as individuals) an opportunistic reproductive strategy, in that they usually lay an egg every year, even when failure is certain. Nevertheless, the decision to breed was not entirely blind, and we identified groups of birds that invested differentially in breeding attempts. The decision to breed was related to the previous breeding frequency, i.e. 81% of the birds that had bred continuously in the past started a new breeding attempt, but only 67% of birds that had missed a year did so. In intermittent breeders, birds that had bred frequently, more often started a new breeding attempt than birds that had largely missed years (71% versus 57%, respectively). Classes of breeders could correspond to age classes, to birds of different breeding quality or to alternative breeding strategies coexisting in the species. Testing the hypothesis that reproductive effort increases with age should be possible in future.     

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.     

Intraspecific nest parasitism in the Spotless Starling Sturnus unicolor

Intraspecific nest parasitism in two colonies of Spotless Starling Sturnus unicolor breeding in nestboxes was studied in central Spain from 1991 to 1994. Nests were monitored regularly and three criteria were used to detect nest parasitism: the appearance of more than one egg per day during the laying period of the host; the appearance of an egg after the start of incubation; eggs with unusual shape or pigmentation. The proportion of parasitized nests in first clutches (37%) was twice that of intermediate (19%) or second (20%) clutches in colony B, whereas parasitism occurred in first (35%) and intermediate (12%) but not in second clutches in colony A. Most clutches (52–70%) were parasitized during the host's laying period and received one parasitic egg. In 10% of the parasitized clutches in colony B, one of the host's eggs disappeared on the day the parasitic egg was added, suggesting that the parasitic female removed this egg. Although parasitism increased clutch size significantly, it led to a decrease in host breeding success, mainly through the removal of eggs and the loss of host nestlings and the survival of parasitic chicks. Observations suggested that parasitic females were young individuals without their own nests and/or those whose breeding attempt had been disrupted while laying in their own nest.     

Sex-specific fitness correlates of dispersal in a house sparrow metapopulation

1.  Dispersal affects many important ecological and evolutionary processes. Still, little is known about the fitness of dispersing individuals.
2.  Here, we use data from a long-term study of a house sparrow Passer domesticus metapopulation to compare lifetime reproductive success (LRS) of resident and immigrant individuals, all with known origin.
3.  Lifetime production of recruits by immigrant males was much lower than for resident males, because of shorter life span and lower annual mating success. In contrast, lifetime production of recruits did not differ significantly between immigrant and resident females.
4.  Over their lifetime, dispersers contributed fewer recruits to the local population than residents. This shows that immigrant house sparrows have different, sex specific, demographic effects on the population dynamics than residents.     

Recruitment processes in long-lived species with delayed maturity: estimating key demographic parameters

Long-lived species show delayed maturity and generally skip breeding sites on a given year, causing difficulties in estimating demographic parameters. A novel multi-state capture – recapture model (model G for general) is proposed to estimate survival and recruitment. Model G considers long and short periods of non-attendance at the breeding site. Model G is compared against a reduced model (model R) to test if pre-breeders skip the studied site a given year, potentially prospecting other breeding habitats.
This model is applied to the vulnerable Cory's shearwaters seabird species. As for many species, natal philopatry, pre-breeding attendance and prospecting play an important role for recruitment into the breeding population. Model G shows that all Cory's shearwaters come back to their natal colony, and attend the colony before recruiting into the breeding population. Moreover, model G is preferred against model R, highlighting that prospecting plays a potentially role in the recruitment processes. Return and recruitment probabilities show similar time variation, suggesting an environmental forcing. The maximum probability of the first breeding is reached at the age of seven years. During the first year the annual survival rate is 0.52 (IC_95%=[0.40, 0.64]) and pre-breeders and adult annual survival rates do not differ, being respectively 0.88, IC_95%=[0.83, 0.92] and 0.89, IC_95%=[0.88, 0.9].
Model G provides an improvement to estimate demographic parameters for long-lived species life cycle and an adequate framework to test the influence of covariates. Model G is structured by age and breeding status categories, allowing easy further population models.