Population dynamics and reproduction of Northern Lapwings Vanellus vanellus in a meadow restoration area in central Sweden

To investigate the effectiveness of a meadow restoration programme, we studied the relationships between population changes and environmental changes, return rates and hatching success in a population of Northern Lapwings Vanellus vanellus. The study was performed on mixed farmland (59 km²) in central Sweden, an area that underwent extensive meadow restoration between 1985 and 1994. The study included more than 2600 nests, supplemented with observations of 127 individually colour-ringed Northern Lapwings. The breeding population varied (2.7–5.3 pairs/km²), but showed no significant trend with time. The population increased in years with high spring flooding levels. Population size was unrelated to demographic factors (e.g. hatching success the previous year (14–50%), and return rate). Lapwings moved considerably between years and their nest site fidelity was unrelated to previous hatching success or other factors, suggesting that changes in habitat quality and migration between populations were important in regulating population size. Recent extensive meadow restoration did not seem to aid the Northern Lapwing population; birds continued nesting on tillage even though most nests were destroyed by farming activities. A relatively high relaying frequency improved hatching success, which was still lowest in the most preferred habitats (spring sown crops, total hatching success c. 30%). The few Northern Lapwings breeding in the least preferred habitats (meadows and cultivated grassland) had a better hatching success (> 70% total hatching success), suggesting that habitat selection was not determined by hatching success. From these results, recommendations are made for the management of Northern Lapwing (and associated species) populations on farmland.     

The breeding biology of the Dalmatian Pelican Pelecanus crispus

The breeding biology of the Dalmatian Pelican Pelecanus crispus , a vulnerable bird species in the Palaearctic, has been studied for 7 years at Lake Mikri Prespa, a continental wetland and Tsoukalio lagoon, a coastal wetland within the Amvrakikos Gulf. The two colonies showed stability, with fluctuations in the number of breeding attempts each year varying by about 30% over a period of 7 years. The date of arrival at the breeding sites and the date of laying of the first egg varied between mid-February and mid-March. The egglaying period (including renesting) varied between 18 and 114 days between years. The annual average clutch size varied between 1.60 and 1.94 eggs per nest. The incubation period averaged 31.4 ± 1.4 days. The overall hatching success varied from 36% to 70%. Egg losses were mainly due to nest abandonment, infertile eggs and eggs rolled from the nests. The Dalmatian Pelican chicks took 11–12 weeks to fledge. Nestling mortality was very low in contrast to other species of pelican. Such a low nestling mortality explains why the breeding success each year is largely explained by the hatching success.     

Spatial differences in breeding success in the pied avocet Recurvirostra avosetta: effects of habitat on hatching success and chick survival

I studied the breeding biology of pied avocets Recurvirostra avosetta in natural habitats (alkaline lakes), and in semi-natural sites (dry fishpond, reconstructed wetlands) in Hungary to relate habitat selection patterns to spatial and temporal variation in breeding success. Colonies were initiated earlier in semi-natural sites than in natural habitats, and earlier on islands than on the mainland. Hatching success was higher on islands than on the mainland, in semi-natural sites than in natural habitats, in colonies of at least 15 pairs than in smaller colonies, and for nests initiated earlier than later within a colony. Fledging success was higher in the wet years (1999–2000) than in the dry year (1998), decreased strongly by season in both habitats and increased with average daily temperature in the first week post-hatch in 1999–2000. Most pairs hatching young in semi-natural sites attempted to lead their chicks to feeding areas in natural habitats, whereas no such movement occurred in the opposite direction. Chick mortality due to predation was high during brood movements and only 23% of the pairs moving their young produced fledglings, compared to 43% for pairs remaining in semi-natural sites and 68% for pairs hatching and rearing young in natural habitats (total n=192 broods). These results suggest that semi-natural sites were more suitable for nesting, whereas natural habitats were more suitable for chick-rearing. The opposing trends in habitat-related breeding success between nesting and chick-rearing suggest sub-optimal habitat selection by Pied Avocets due to an incorrect assessment of the potential for successful reproduction of semi-natural sites, which may thus function as ecological traps.     

Reproductive performance of the Bald Ibis Geronticus calvus in relation to rainfall and grass-burning

Reproductive performance of the Bald Ibis Geronticus calvus was studied during three consecutive nesting seasons (1978–80) at a single large breeding colony in Natal, South Africa. A focal sample of 22 nest sites (out of a total of 60 in the colony) was inspected at frequent intervals to determine clutch sizes, egg survival, hatching success, nestling mortality and fledgling production in each year. Mean clutch size was slightly lower in 1980, compared to the two previous years, although survival and hatching success of eggs were similar in all three years. However, nestling mortality was significantly higher in 1980, largely as a result of parental nest desertion and nestling starvation, so that the proportion of breeding pairs producing at least one fledged offspring, and the average brood size at fledging, were lower compared to 1978 and 1979.
The 1978 and 1979breeding seasons were both preceded by 12 months of above average rainfall, and Bald Ibises fed extensively on small invertebrates obtained in burnt grassland in the study area during the nesting period. By contrast, the onset of nesting in 1980 followed 11 consecutive months of sub-normal rainfall. As a result of the drought, and the consequent lack of controlled burning by landowners in the study area in 1980, very little burnt grassland was available during the Bald Ibis' breeding season, and the birds fed primarily in cultivated pastures during the customary burning period. The lower reproductive performance of Bald Ibises at the study site in 1980 is discussed in relation to the possible effects of drought on the abundance of invertebrate prey and the extent of controlled burning in the species' grassland biotope.     

Breeding biology and success of the Bearded Vulture Gypaetus barbatus in the eastern Pyrenees

We present data from an extensive study of Bearded Vulture Gypaetus barbatus breeding biology in the Pyrenees from 1992 to 1999. Average laying date was 6 January (range 11 December to 12 February, n  = 69) with no significant differences between years. Eighty per cent of clutches were of two eggs ( n  = 20) and average incubation was 54 days (range 52–56, n  = 14). Hatching occurred on average between 21 February and 3 March (range 5 February–7 April) and the first and last chicks fledged in 21–27 May and 20 July, respectively. The average chick age at fledging was 123 days (range 103–133, n  = 20). Bearded Vulture density increased significantly during the study period. Breeding success and productivity declined apparently as a consequence of the increase in the percentage of breeding failures during incubation and chick rearing, most during the hatching period. The factors that may determine breeding failure and the decline in breeding performance are analysed and management recommendations for more effective conservation measures are discussed.     

Interannual variation in the breeding biology of the Antarctic prion Pachyptila desolata at Bird Island, South Georgia

Interannual variation in aspects of the breeding biology of Antarctic prions was studied for three summers (1989–1992) at Bird island, South Georgia. Egg size, mass and incubation period remained constant. Laying, hatching and fledging were significantly delayed and less synchronous in 1991/92 (range of laying dates 51 days compared to 10–15 days in the two other seasons). This was due to an unusually cold and protracted winter, with ice blocking burrows into the spring, restricting availability of nest sites. Brooding lasted longer in 1991/92 but the overall fledging period was unchanged. Skeletal growth rates did not vary amongst years; growth in mass was slower in 1989/90 but fledging mass was similar in all three years. In 1989/90 and 1991/ 92 later hatched chicks grew (in mass) faster. The survival of chicks from hatching to fledging did not vary amongst years or with hatching date. Feeding frequency was similar between years, once allowance had been made for starlit nights. Thus late and asynchronous breeding in 1991/92 did not result in reduced breeding success either through predation or starvation.
Crustaceans formed 98–99% of the mass of the identifiable portion of regurgitated food samples. Significant annual variation was found within these crustaceans with the presence of krill (least in 1990/91) being inversely related to that of amphipods and copepods. There was no relationship between diet composition and chick growth or survival. Other seabird species, lacking the morphological specialization for feeding on copepods and amphipods, had very low breeding success in 1990/91, when krill was scarce.     

Effects of microhabitat preferences on kelp gull Larus dominicanus breeding performance

We examined microhabitat preferences and their adaptiveness for the kelp gull Larus dominicanus in one of its largest colonies in Patagonia. We quantified 13 habitat variables at 104 and 92 nest-sites and at 70 and 60 random points in 1998 and 1999, respectively. We recorded egg laying dates, hatching success, number of chicks fledged and breeding success at 92 of these nests in each of the study seasons. Compared to random points, nest sites had more vegetation cover and were closer to the nearest available bush, but they were located further from the nearest clearing for flight take-off, all variables included in the "vegetation" principal component. Nests were built on substrates with lower slopes and higher percentages of silt-clay, but with lower percentages of rock. In both years, hatching success, number of chicks fledged and breeding success were related positively to the vegetation principal component. Substrate slope and composition contributed to explain the variation in hatching and breeding success in 1998 and 1999, respectively. Our study shows that some kelp gull breeding parameters are affected by nest-site habitat characteristics. These associations were in most cases influenced by the timing of breeding, with earlier breeders being more successful. Vegetation effects and substrate characteristics were important variables in distinguishing random sites from nest sites, and in explaining variation in breeding performance, showing that their habitat preferences are adaptive and allowing us to detect both the pattern and process in kelp gull habitat selection.     

Breeding biology of the Barn Owl Tyto alba in central Mali

Data were obtained on 178 clutches of African Barn Owls in central Mali from four breeding seasons during 1979–1983. Significantly more clutches were laid in 1979–1980 and significantly fewer in 1980– 1981 than the average for the 4 years and there were significantly more clutches laid in the middle period of the annual breeding season. The egg volume was significantly smaller at the beginning of the breeding season and significantly larger in the middle than the overall mean with eggs of second clutches being larger than those of first clutches. The clutch size was 605 eggs of which 479 hatched. The number of young fledged per successful nest was 319 and was 1 83 for all nesting attempts. The month was the only variable shown to affect significantly the clutch size, eggs hatched and fledging rate, the highest success rates being associated with the middle of the breeding period. The average interval between the hatching of eggs was 2–31 days. Survival rates (47'1%) to fledging were significantly affected by year (1981–1982 being the least) and month (mid-season birds the best). The order of hatching significantly affected age at death or disappearance, the first-hatched birds surviving the longest. The year significantly affected age at fledging, the young from the year in which most clutches were laid leaving the nest at the youngest age and those associated with the year having the least number of clutches remaining in the nest the longest. The month of hatching also affected fledging age, birds at the extremes of the breeding season fledging at older ages. The discussion compares these data with those from elsewhere.     

Drivers of the breeding success of American Oystercatchers (Haematopus palliatus frazari) at a critical site in Sinaloa,Mexico

Understanding breeding phenology and success can elucidate population dynamics, which is especially important for species in need of conservation. We describe the factors affecting the breeding biology of American Oystercatchers (Haematopus palliatus frazari) at El Rancho Island, a critical site that contains ~ 7% of the total estimated population, on the coast of Sinaloa, Mexico. We monitored 192 nests over four years (2016–2019). The breeding season lasted from March to June and mean laying dates differed among years, with the mean laying date in 2019 an average of 20 days earlier than in 2016. Clutch sizes decreased as the breeding season progressed. Both breeding success and productivity differed among years, with the lowest values in 2016 (30% hatching success and 0.6 chicks/nest) and the highest in 2019 (66% hatching success and 1.2 chicks/nest). Hatching success was affected by year, laying date, type of habitat, and distance to the high tide line. American Oystercatchers that laid eggs earlier in the season, used mixed marsh and dune habitat, and with nests relatively close to the waterline (< 50 m) had greater breeding success. Overall, however, the breeding success of American Oystercatchers was low and influenced by a combination of several intrinsic and extrinsic factors. Management measures may be required to increase breeding success and ensure the conservation of this subspecies.     

The effect of maternal age and experience on egg-size and hatching success in Wandering Albatrosses Diomedea exulans

The roles of maternal age and experience, on the one hand, and individual, year and random effects on the other, in influencing avian egg-size and hatching success have been much debated but seldom studied comprehensively. We investigated these topics with Wandering Albatrosses Diomedea exulans of known age (7–30 years) and experience (1–8 breeding attempts) over a 10-year period. Older and more experienced birds laid larger eggs. After allowing for year and controlling for experience, significant age effects remained; after controlling for age, no detectable experience effects remained. However, age accounted for only 6% of the overall egg-size variation. Egg-size varied significantly between years and has increased over the last decade. Individuals laid eggs of consistent sizes; 55% of the random variation in egg-weight was due to such effects. Egg- and hatchling-weight were very closely linked; larger eggs also had higher hatching success. The latter was influenced significantly by age and experience but neither remained significant after controlling for the other. Year effects were also detectable.
That there are significant effects of age, experience, year and individual on egg-weight (and hatching success) is probably typical of seabirds generally, though with different balances between factors depending on species and situation; however, insufficient data exist to examine this critically. Our finding that age was a more important influence than breeding experience does not support recent suggestions that hatching success is mainly influenced by experience and that experience will have a greater effect on reproductive success in long-lived species with high mate-fidelity. However, Wandering Albatrosses may have acquired much relevant experience before even starting to breed.     

Restoration and management for cliff‐nesting birds in Mediterranean mining sites: the Sand Martin case study

Habitat enhancement for birds is frequently implemented during mine site restoration. Cliff‐nesting birds often colonize anthropogenic environments such as mining areas (aggregate sites and quarries for aggregate and cement production). Mining activity can compromise breeding success, causing cliff‐nesting birds to depend on the management and restoration of mining areas. The objective of our study is to assess the importance of mine site habitats for Sand Martin conservation and reconcile mining activity with breeding success in Mediterranean environments. We studied Sand Martin breeding habitat preferences in mining areas at three spatial scales. At the mining site scale, we compared 10 mining sites with Sand Martin burrows with 19 mining sites without burrows. At the colony scale (vertical structures with colonies), we evaluated the relationships between the number of breeding pairs, number of burrows, and colony characteristics within 30 distinct Sand Martin colonies. At the burrow scale, we compared the characteristics of the available vertical structure with the areas used by Sand Martins. At the mining site scale, Sand Martins preferred more surface of water bodies, shorter distances to flowing water, older sites, and mining sites which produce aggregates instead of cement. At the colony scale, Sand Martins preferred southwest orientations and stockpiles to vertical extraction faces. At the burrow scale, birds preferred the most vertical areas of the face. Our results support the need for effective habitat restoration and improved management for more effective Sand Martin conservation within mining areas. Simple interventions can enhance habitat quality and conservation of cliff‐nesting birds.     

The role of habitat composition in determining breeding site occupancy in a declining Ring Ouzel Turdus torquatus population

Ring Ouzel Turdus torquatus breeding site occupancy in the Moorfoot Hills, southeast Scotland, was determined during the periods 1952–85 and 1998–2000, and sites were classed as either occupied or deserted during the latter period. Site occupancy during 1998–2000 was related to habitat data derived from a supervised classification of a Landsat 7 satellite image. Breeding sites were more likely to have remained occupied if they were at higher altitudes and had more heather cover within radii of both 200 and 450 m, although these effects became non-significant when spatial autocorrelation was accounted for. Using only topographical variables, potential breeding sites were predicted and the habitat surrounding them was compared with actual breeding sites. Actual breeding sites were more likely to have heather or grass–heather mosaic within 100 m than potential breeding sites, but there were no further habitat differences between actual and potential breeding sites within radii of 200 or 450 m. These results indicate that breeding sites at higher altitudes and with a higher cover of heather were less likely to have been deserted, and that lack of heather may prevent otherwise suitable breeding sites from becoming occupied by Ring Ouzels. More detailed field studies are needed to understand the mechanisms underlying these associations.     

Breeding ecology of the Fulmar Fulmarus glacialis and the Kittiwake Rissa tridactyla in high-arctic northeastern Greenland, 1993

The breeding ecology of the Fulmar Fulmarus glacialis and the Kittiwake Rissa tridactyla in the high Arctic was studied in relation to the occurrence of the northeast water polynya in northeasternmost Greenland (80̀N). Mean laying dates were 31 May in the Fulmar and 18 June in the Kittiwake; the total nesting season for the Fulmar just matched the time window of the polynya opening period. Fulmar colony attendance fluctuated within a period of 11.6 days because of variation in nonbreeding prospectors but showed no clear diurnal variation. Fulmar incubation shifts, on average, lasted 6.1 days (range 1–13 days), which is significantly longer than elsewhere, and the average chick-guard period of 10.9 days (range 1–17 days) was significantly shorter than in other studies. Egg neglect occurred in 18% of Fulmar nests or 0.7% of nests per day. Overall breeding success (chicks fledged per egg laid) was 0.56 in the Fulmar and 0.67 in the Kittiwake; the latter produced 1.4 young per active nest or 1.2 per completed nest. Mean Kittiwake clutch size was 2.03; larger clutches were laid early. Nest site characteristics (presumably reflecting nest predation risk) and breeding behaviour affected breeding success. in the Fulmar, hatching success was negatively correlated with laying date and the proportion of egg neglect, while overall breeding success was correlated negatively with distance to nearest neighbouring site and positively with the length of the chick-guard period. Kittiwake breeding success was negatively correlated with laying date. Using seabirds as indicators of marine food supply, breeding success in both species suggested moderate to good food supply in the northeast water polynya in 1993, although at least in the Fulmar the high reproductive output appeared partly maintained by behavioural buffering; long incubation shifts, egg neglect and short chick-guard periods were symptoms of foraging constraints.     

Intraspecific variation in size and density of Avocet colonies: effects of nest-distances on hatching and breeding success

In many colonial bird species there is considerable intraspecific variation in colony size and inter‐nest distance (colony density). Possible causes of this variation and its effects on hatching success (survival of eggs) and breeding success (probability of a pair raising chicks) were studied in 48 Avocet Recurvirostra avosetta colonies in Schleswig‐Holstein (Germany) between 1991 and 1996. Colony density was influenced by time of year and habitat (categories: island or mainland, close to or far from feeding grounds). Colonies on islands had the highest densities. When all available space at a colony site was used, colonies became very dense (mean nearest‐neighbour nest distance less than 1 m). Colony size (number of clutches) was influenced by time of year, but not by habitat. Hatching success was low in high density colonies and in very low density ‘colonies’ (single nests) and high over a broad range of intermediate nest densities. The low success rate of single nests was caused by a very high predation rate, whereas the low success rate in very dense colonies was caused by a high rate of nest abandonment. Nest abandonment in very dense colonies was associated with a high level of aggressiveness among Avocets during the egg‐laying period. Due to territorial behaviour, Avocets seemed to be expelled from the densest breeding sites. In very dense colonies, high frequencies of clutches of unusual size occurred due to conspecific nest parasitism. The number of Avocets taking part in attacks on potential egg predators was small and (in colonies of more than one clutch) depended neither on colony size nor on colony density. Despite a low hatching success in very dense colonies, individuals breeding in the densest colonies had significantly better chances of raising chicks than Avocets breeding in less dense colonies. Coloniality seemed to be obligatory for Avocets in order to ensure hatching success. The size and density of colonies seemed to be associated with the availability of suitable nesting habitats (islands).     

Population biology of Senecio integrifolius (Compositae), a rare plant in Sweden

In a study of natural populations of Senecio integrifolius in southern Sweden (1979–1983) seeds were found to disperse from mid-June to late July and most of them germinated in autumn. No soil seed bank was observed. On a heavily grazed site few seeds were produced but the percentage of estimated germination was high (c. 75%). In two moderately grazed habitats 8–10% of the estimated number of seeds produced in permanent plots germinated. In a lightly grazed habitat many seeds were produced but only a small percentage germinated (1.4%). In field experiments the average germination was 50–53% when newly harvested seeds were sown where the vegetation had been removed, and 20–33% when sown in undisturbed vegetation at a moderately grazed site. Germination was much lower in a lightly grazed habitat (3–12%). Survival of seedlings was much higher in heavily and moderately grazed habitats than in lightly grazed habitats. Mortality tended to be higher during the growing season (mid-April to early November) than during the winter, and increased markedly during a drought period in the summer of 1982. The half-life of plants established in 1980 varied from 39.3 years at the most intensively grazed site to 7.2 at the lightly grazed site. The number of flowering stems varied between years mainly according to weather. Few plants in the permanent plots flowered every year, the flowering being most frequent at heavily grazed sites. It is concluded that heavy grazing by cattle after seed dispersal is the appropriate management for maintaining S. integrifolius in Sweden.     

Breeding success and chronology of Wood Storks Mycteria americana in northern and central Florida, U.S.A.

The breeding success and chronology of Wood Storks Mycteria americana were studied at eight colonies in northern and central Florida during 1981–1985. Mean ± s.d. clutch size for all colony-years was 3.07 ± 0.56 (n = 2694 nests), with three-egg clutches (72%) most frequent. Mean clutch size among all colonies and years ranged from 2.73 ± 0.55 to 3.41 ± 0.61. Many colonies exhibited significant negative trends in clutch size with, hatching date because of a proportional decrease in four-egg clutches later in the season. Mean colony clutch size was not correlated with nest numbers, nesting density or mean hatching date within most years. Mean ± s.d. number of fledglings for all colonies and years was 1.29 ± 1.16 fledglings per nest (n = 2812 nests). Mean annual fledging rates in colonies ranged from 0 (colony failed) to 2.66 fledglings per nest. Most breeding failure occurred prior to egg hatching, and the second highest mortality occurred between hatching and 2 weeks of age. Four-egg clutches fledged more storks than three-egg clutches, which in turn were more successful than two-egg clutches. However, all clutch sizes showed similar fledgling per egg rates. The seasonal decline in productivity was associated proportionally with smaller clutch sizes later in the breeding season. An increase in mean hatching date was correlated with an increase in latitude. There was greater within-year breeding synchrony among colonies than interyear breeding synchrony within each colony. Breeding synchrony was not correlated with mean hatching date, latitude, longitude, nest numbers or nesting density.     

Annual productivity and individual female reproductive success in a Great Bustard Otis tarda population

The reproductive success of Great Bustards Otis tarda in north-western Spain was studied between 1987 and 1998, both at the population ( c . 700 adult females breeding in our study area) and the individual level (sample of 32 marked females). Overall productivity was low, with a population mean of only 0.14 chicks reared per adult female, and an average breeding success of 0.15 chicks per year in the sample of marked females, but interannual variability was high (0.04–0.29). Population productivity was positively correlated with winter (October–March) precipitation prior to each breeding season, and negatively correlated with the number of days of rain during the hatching period. High annual productivity resulted from a high proportion of females rearing two chicks. Reproductive success was higher in females older than 6 years than in younger birds. The proportion of females in the marked sample that failed in breeding after having bred successfully the previous season was significantly higher than the proportion of those that did not. Finally, females with a higher than average breeding success tended to breed successfully in years of both low and high population productivity, whereas those with lower than average breeding success did so only in years of high productivity.     

Factors affecting nesting success of a bird assembly in the central Monte Desert, Argentina

We estimated nesting success of a breeding bird assemblage in the central Monte Desert in four breeding seasons (1995–1999). We analyzed the effect of year, period within the breeding season, nest type, and nest location on daily survival rates (DSR). Averaged nesting success of passeriforms in natural habitats was very low (<20% on average, n=12 species). DSR tended to be lower during the egg-laying stage in Furnariidae, Rhinocryptidae, Mimidae, and Emberizidae. Tyrannidae had the highest DSR for all nesting stages and Emberizidae the lowest. Predation was the main cause of nest failure (>90% on average, n=12 species). DSR differed among years, being higher in the driest year (1995) and lower in the most humid year (1997). Nests initiated in the first half of the breeding season had higher DSR than those initiated in the second half. Closed nests had similar survival rates to open nests. DSR differed according to species of plant that supported the nest. Nest predation in the central Monte Desert may be an important selective pressure in birds' life history, which is in accordance with other results for South American temperate birds.     

Breeding density,habitat selection and reproductive rates of the Peregrine Falcon Falco peregrinus in Álava (northern Spain)

A population of 33–35 pairs of Peregrine Falcon in álava (north Spain) was studied with the aim of assessing the habitat attributes that influence breeding density, habitat selection and breeding success. A strong relationship was found between density of the species in each UTM (Universal Transverse Mercator) square of 10 × 10 km and cliff availability. Habitat selection was analysed by comparing 15 variables in 33 occupied and 25 unoccupied cliffs located at least 2 km from the nearest Peregrine pair. Significant differences were found in five variables: cliff dominance, distance to the nearest Golden Eagle Aquila chrysaetos nesting cliff, steepness and altitude (all showing larger values in occupied cliffs), and cliff orientation, with occupied cliffs facing preferably south and east. Orientation, dominance and distance to the nearest Golden Eagle pair, and the distance to the nearest Eagle Owl Bubo bubo nesting cliff, were included in a discriminant analysis which classified 82.76% of the cliffs correctly. The productivity of the studied population was 1.44 young/territorial pair (n = 45), and no consistent relationship was found between breeding success and habitat variables.     

Factors affecting population regulation of a colonial vulture

Two hypotheses have been proposed to link population regulation to density‐dependent changes in demographical parameters: the habitat heterogeneity hypothesis (HHH) states that, as population density rises, an increasing proportion of individuals are forced to occupy low‐quality territories, which provokes a decline in average per‐capita survival and/or productivity although some individuals show no decline in fecundity; and the individual adjustment hypothesis (IAH), which suggests that increased densities lead to reductions in survival and/or fecundity by enhancing agonistic interactions, which affect all individuals to a similar extent. However, density‐dependent effects can be affected by density‐independent factors (DIF), such as weather. We test the effects of density dependence on annual reproductive success in Griffon Vultures Gyps fulvus at four spatial scales, nest‐site, cliff, colony and metacolony, in northern Spain from 2008 to 2015. Our results showed most support for the HHH at all scales. At the colony and cliff scale, IAH and DIF had similar importance, whereas there was little evidence of IAH at the metacolony and the nest scale. The best protected eyries (caves, potholes and sheltered ledges) produced the most fledglings and were used preferentially, whereas low‐quality eyries (exposed ledges or open crevices) were used only when the number of breeders increased. The significant interaction between breeding failure and density found for the more exposed eyries suggests that at higher densities, breeding pairs are forced to use poorer nesting areas, and the negative effect of density at the cliff scale could be due to the combined effect of a higher proportion of pairs using low‐quality eyries and the negative effect of rainfall.