Feeding territoriality and the reproductive consequences in brown skuas<Emphasis Type="Italic"> Catharacta antarctica lonnbergi</Emphasis>

In the maritime Antarctic, brown skuas (Catharacta antarctica lonnbergi) show two foraging strategies: some pairs occupy feeding territories in penguin colonies, while others can only feed in unoccupied areas of a penguin colony without defending a feeding territory. One-third of the studied breeding skua population in the South Shetlands occupied territories of varying size (48 to >3,000 penguin nests) and monopolised 93% of all penguin nests in sub-colonies. Skuas without feeding territories foraged in only 7% of penguin sub-colonies and in part of the main colony. Females owning feeding territories were larger in body size than females without feeding territories; no differences in size were found in males. Territory holders permanently controlled their resources but defence power diminished towards the end of the reproductive season. Territory ownership guaranteed sufficient food supply and led to a 5.5 days earlier egg-laying and chick-hatching. Short distances between nest and foraging site allowed territorial pairs a higher nest-attendance rate such that their chicks survived better (71%) than chicks from skua pairs without feeding territories (45%). Due to lower hatching success in territorial pairs, no difference in breeding success of pairs with and without feeding territories was found in 3 years. We conclude that skuas owning feeding territories in penguin colonies benefit from the predictable and stable food resource by an earlier termination of the annual breeding cycle and higher offspring survivorship.Research licence: Umweltbundesamt Bonn 13.4-94003-1/5-7.     

Occupancy of yellow‐billed and Pacific loons: evidence for interspecific competition and habitat mediated co‐occurrence

Interspecific competition is an important process structuring ecological communities, however, it is difficult to observe in nature. We used an occupancy modelling approach to evaluate evidence of competition between yellow‐billed (Gavia adamsii) and Pacific (G. pacifica) loons for nesting lakes on the Arctic Coastal Plain of Alaska. With multiple years of data and survey platforms, we estimated dynamic occupancy states (e.g. rates of colonization or extinction from individual lakes) and controlled for detection differences among aircraft platforms and ground survey crews. Results indicated that yellow‐billed loons were strong competitors and negatively influenced the occupancy of Pacific loons by excluding them from potential breeding lakes. Pacific loon occupancy was conditional on the presence of yellow‐billed loons, with Pacific loons having almost a tenfold decrease in occupancy probability when yellow‐billed loons were present and a threefold decrease in colonization probability when yellow‐billed loons were present in the current or previous year. Yellow‐billed and Pacific loons co‐occurred less than expected by chance except on very large lakes or lakes with convoluted shorelines; variables which may decrease the cost of maintaining a territory in the presence of the other species. These results imply the existence of interspecific competition between yellow‐billed and Pacific loons for nesting lakes; however, habitat characteristics which facilitate visual and spatial separation of territories can reduce competitive interactions and promote species co‐occurrence.     

Territory acquisition in loons: the importance of take-over

We examined patterns of territory acquisition and reconnaissance in common loons, Gavia immer, from northern Wisconsin. Among all territory acquisitions, 41.5% occurred through passive occupation of territories left vacant after the death or desertion of a previous resident, 17% constituted founding of new territories and the remaining 41.5% came about through take-over: either usurpation of defended territories or appropriation of territories before the seasonal return of previous owners. Take-overs occurred in both sexes, but individuals acted alone, never in pairs. Displaced breeders usually took refuge on undefended lakes near their former territories; about half of these loons later regained former territories through passive occupation or took possession of new territories elsewhere. As predicted by the reconnaissance hypothesis, usurpations occurred most often in territories that had produced chicks during the previous 12 months, suggesting that loons use the presence or absence of chicks as a cue for territorial usurpation. Large individuals of both sexes held onto territories longer than small individuals, an indication that body size might be correlated with fighting ability. In terms of life history, loons appear to locate good territories through reconnaissance, usurp them in a subsequent year and recover from displacements by reclaiming their original territories or new ones. Copyright 2000 The Association for the Study of Animal Behaviour.     

FOOD RESOURCES, TERRITORY DENSITY AND REPRODUCTIVE SUCCESS OF AN ISLAND SILVEREYE POPULATION ZOSTEROPS LATERALIS

We examined the relationships between food resources, territory density and some breeding parameters (individual reproductive success and parental age) of a Capricorn Silvereye population during two years' detailed study on Heron Island, Great Barrier Reef. The territorial behaviour of the Silvereyes included dawn song. aggression and distinctive vocalizations. During both years, about 85% of pairs occupied territories although the population sizes differed. The dispersion pattern of territories was not regular and was correlated with the density of figs and human food scraps. Nestlings were fed more figs as they grew, and the parents foraged in fig trees outside their territories. However, only about 60%, of territorial pairs fledged young. Pairs which fledged most young were older, fed more insects to their nestlings, and nested in areas containing more fruiting fig trees per territory. We conclude that food resources were in short supply, and that access to fig trees provided breeding birds with a quick energy source while they searched for insect protein for the young. The data are consistent with the predictions of the ‘sufficient resource‘ hypothesis of the adaptive significance of territoriality but not with its assumption that the principal benefit is the food within the territory.     

Investment in territorial defence relates to recent reproductive success in common loons Gavia immer

As the value of a limited resource such as a territory increases, animals should invest more in the defence of that resource. Because reproductive success often depends on the quality of a breeding territory, reproductive success or failure may alter the perceived value of territory and affect an animal's investment in territorial defence. We used common loons Gavia immer to test the hypothesis that animals with recent breeding success would show stronger territorial defence than those with no recent breeding success. Surprisingly, successful loons responded less, not more, to a simulated intrusion. However, birds with success in the previous season also increased their territorial response as the breeding season progressed. In conjunction with past data showing that recently successful loons experience an increase in conspecific intrusions on their territories, we interpret our data to suggest that loons with recent success offset the cost of increased intrusions by adopting a more efficient strategy for territorial defence (e.g. limiting investment in resource defence until the time of the season when it is most critical).     

Territorial occupancy and breeding performance in a migratory raptor do not follow ideal despotic distribution patterns

It has commonly been argued that many territorial species select their breeding sites following an ideal despotic distribution model, in which the most productive, high-quality territories are more frequently occupied. Theoretical and empirical studies have shown that this occupancy pattern may have population regulatory consequences, leading to density dependence in heterogeneous habitats. During a 9-year research project in a forested area of south-eastern Spain, we tested some of the predictions of the ideal despotic distribution model and the site-dependent population regulation theory in a migratory raptor species, the booted eagle Hieraaetus pennatus . Contrary to the predictions of the despotic model, our results showed that the temporal pattern of territorial occupation did not differ from randomness, and that the territory occupancy rate was not significantly related to the reproductive parameters considered. At population level, the breeding variables were density independent, suggesting the absence of site-dependent regulation. In addition, we were unable to find significant differences in the habitat characteristics between high-quality and low-quality territories, classified according to the criteria of both occupancy frequency and average productivity. Overall, our results suggest that booted eagles select their territories at random, probably due to the lack of strong environmental heterogeneity, and that occupancy rate is not a good measure of territory quality for the population studied.     

Effects of spatial disturbance on common loon nest site selection and territory success

The common loon (Gavia immer) breeds during the summer on northern lakes and water bodies that are also often desirable areas for aquatic recreation and human habitation. In northern New England, we assessed how the spatial nature of disturbance affects common loon nest site selection and territory success. We found through classification and regression analysis that distance to and density of disturbance factors can be used to classify observed nest site locations versus random points, suggesting that these factors affect loon nest site selection (model 1: Correct classification = 75%, null = 50%, K = 0.507, P < 0.001; model 2: Correct classification = 78%, null = 50%, K = 0.551, P < 0.001). However, in an exploratory analysis, we were unable to show a relation between spatial disturbance variables and breeding success (P = 0.595, R² = 0.436), possibly because breeding success was so low during the breeding seasons of 2007–2008. We suggest that by selecting nest site locations that avoid disturbance factors, loons thereby limit the effect that disturbance will have on their breeding success. Still, disturbance may force loons to use sub-optimal nesting habitat, limiting the available number of territories, and overall productivity. We advise that management efforts focus on limiting disturbance factors to allow breeding pairs access to the best nesting territories, relieving disturbance pressures that may force sub-optimal nest placement. © 2011 The Wildlife Society     

Breeding performance, age effects and territory occupancy in a Bonelli's Eagle Hieraaetus fasciatus population

Bonelli's Eagle Hieraaetus fasciatus is one of the most endangered birds of prey in Europe. Despite mounting interest and research, several questions regarding the conservation implications of territory occupancy and site-dependent population regulation remain insufficiently explored for this species. Here, we report on a 12-year study of the territorial structure of a Bonelli's Eagle population in southeastern Spain. No signals of population decline were found in the breeding population, as mean annual productivity was stable and the presence of mixed-age pairs in the population decreased with the years. However, the average proportion of subadults occupying territories was larger than that observed in other Spanish populations. Contrary to the predictions of a despotic distribution model, we found no significant relationship between occupancy rates and breeding parameters. Our results showed significant variations in productivity attributable to differences in the quality of individuals (i.e. mixed versus adult pairs), but no variability among territories per se (i.e. caused by habitat heterogeneity). Moreover, coexistence with intraguild species did not have any significant effect on productivity, although the proximity of Eagle Owls Bubo bubo affected the occupation rate of territories. Finally, our population does not appear to experience site-dependent population regulation, as a positive relation between mean annual productivity and density was found. The threat posed by changes in land use in the study area leads us to suggest that strict protection of current territories is necessary to ensure population persistence, and we suggest that a significant population increase is only likely if new or deserted territories become available.     

A long-term large-scale study of the breeding biology of the Spanish imperial eagle (Aquila adalberti)

We present data from a 17-year study of the population biology of a growing population of Spanish imperial eagles Aquila adalberti across most of its breeding range. The objective of this study was to analyse the effects of age, supplemental feeding and rabbit haemorrhagic disease (RHD) on several breeding parameters of this population of eagles. Average clutch size was 2.2 eggs per clutch, and the average incubation time was 41.7 days per clutch. Fledging occurred an average of 76.8 days after hatching, the length of the fledgling period was not correlated to clutch size. The annual average percentage of pairs laying eggs was 88%. A significant reduction in the percentage of pairs laying eggs in the period 1992–1994 (from 91 to 81%) coincided with most of the eagles’ territories being affected by the rabbit epizootic disease, RHD, which reduced their food supply significantly. Average productivity was 1.23 chicks per monitored territory, average breeding success was 1.40 chicks in a clutch per territory and the average fledging rate was 1.69 chicks per territory with hatching success. The main known causes of breeding failure during incubation were nest collapse and human disturbance. During chick-rearing, total or partial chick losses were mainly caused by siblicide, disease, malnutrition or nest collapse. In 26.2% of the 1372 monitored breeding attempts, at least one of the breeding birds was a subadult (the male in 56.1% of the cases, the female in 15.5% and both sexes in 28.4% of cases). In cases of mixed-aged pairs (n = 205), 70.7% were the result of a substitution, and 29.3% were the result of the forming of a new pair. Egg laying took place significantly earlier and breeding success was higher in territories occupied by adults than in those occupied by subadults. Breeding parameters were higher in high-quality (rabbit-rich) territories than in low-quality (rabbit-poor) territories, but only for those territories occupied by adults. The values obtained in the territories occupied by adults were only significantly higher than in those of the subadults in high-quality territories. Age and territory quality thus simultaneously affected reproductive output.     

对白枕鹤领域的初步研究

1986-1988年的3-5月间,在黑龙江省林甸县对白枕鹤的领域进行了研究。三年中,对8个巢的领域研究结果表明,白枕鹤多年连续利用同一领域,占据的最大领域面积为10.7平方公里,平均不超过7.0平方公里,并随繁殖过程而缩小,至雏期活动和防御范围只有1.5平方公里左右。多数白枕鹤的领域内包括一片耕地为固定觅食区,但小于无耕地的领域面积。     

Pair territoriality in the longnose filefish,oxymonacanthus longirostris

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

Territory occupancy and breeding success of Peregrine Falcons Falco peregrinus at various stages of population recovery

Organochlorine pesticides disrupted reproduction and killed many raptorial birds, and contributed to population declines during the 1940s to 1970s. We sought to discern whether and to what extent territory occupancy and breeding success changed from the pesticide era to recent years in a resident population of Peregrine Falcons Falco peregrinus in southern Scotland using long‐term (1964–2015) field data and multi‐state, multi‐season occupancy models. Peregrine territories that were occupied with successful reproduction in one year were much more likely to be occupied and experience reproductive success in the following year, compared with those that were unoccupied or occupied by unsuccessful breeders in the previous year. Probability of territory occupancy differed between territories in the eastern and western parts of the study area, and varied over time. The probability of occupancy of territories that were unoccupied and those that were occupied with successful reproduction during the previous breeding season generally increased over time, whereas the probability of occupancy of territories that were occupied after failed reproduction decreased. The probability of reproductive success (conditional on occupancy) in territories that were occupied during the previous breeding season increased over time. Specifically, for territories that had been successful in the previous year, the probability of occupancy as well as reproductive success increased steadily over time; these probabilities were substantially higher in recent years than earlier, when the population was still exposed to direct or residual effects of organochlorine pesticides. These results are consistent with the hypothesis that progressive reduction, followed by a complete ban, in the use of organochlorine pesticides improved reproductive success of Peregrines in southern Scotland. Differences in the temporal pattern of probability of reproductive success between south‐eastern and south‐western Scotland suggest that the effect of organochlorine pesticides on Peregrine reproductive success and/or the recovery from pesticide effects varied geographically and was possibly affected by other factors such as persecution.     

Responses of breeding common loons to human activity in upper Michigan

Breeding populations of the common loon (Gavia immer) in Michigan have declined in the past several decades, resulting in classification of this species as threatened under state law. Factors responsible for the decline are unknown, but may include toxic contaminants, mortality in commercial fish nets, and human disturbance of breeding sites. To assess the latter possibility, 960 hours of observation were devoted to observing human-loon interactions on two sets of lakes, one with restricted human use (minimal or no shoreline development and open to canoe use only), and another with unrestricted use (varying amounts of shoreline development and motorboat traffic). Six mated pairs of loons on six restricted use (r-u) lakes were compared to eight loon pairs on seven open-use (o-u) lakes. The number of nests that hatched young per nest started was not significantly different between the two sets of lakes (7 of 13 (0.62) on r-u lakes vs 8 of 17 (0.47) on o-u lakes), despite significantly more human activity on o-u lakes. Chicks hatched per pair of loons were likewise not significantly different (1.1 vs 1.2 on r-u and o-u lakes, respectively). Fledging success was significantly lower on r-u lakes (7 chicks fledged of 11 hatched) than on o-u lakes (13 fledged of 13 hatched). Human activity on o-u lakes was 2–3 times that on r-u lakes during chick rearing, but time spent by adult loons tending and feeding chicks was not significantly different between the two types of lakes. The larger size of most o-u lakes may have allowed loons a greater opportunity to avoid human disturbance. Higher levels of human activity did not affect production of chicks by loons under the conditions observed, but these results should not be extrapolated to lakes experiencing much higher human use.     

Territory occupancy rate of goshawk and gyrfalcon: no evidence of delayed numerical response to grouse numbers

Two recent studies on territory occupancy rates of goshawk Accipiter gentilis and gyrfalcon Falco rusticolus report a 2–3-year-delayed numerical response to grouse numbers, which is a requirement for a hypothesis of predator-generated grouse cycles. The time lags were assumed to reflect the average age of sexual maturity in the raptor species. In southern Norway, however, subadult (two-year-old) goshawk hens occupied only 18–25% of territories where occupancy was not recorded in the preceding year, and there was no significant relationship between the proportion of subadults among recruits and grouse indices two years earlier. We argue that territory occupancy rates are not appropriate indices of total raptor population levels, but rather reflect the proportion of territorial pairs that attempt to nest. Because this depends on the body condition of the hens, fluctuations in other important winter resident prey species (most important for the goshawk) and winter weather (most important for the gyrfalcon) should also be addressed. During 1988–2006, the annual proportion of goshawk territories with recorded nesting attempts in southern Norway was most closely related to the preceding autumn’s population indices of black grouse Tetrao tetrix and mountain hare Lepus timidus, whereas the annual proportion of gyrfalcon territories with observations of falcons or with confirmed breeding attempts in central Norway were best explained by population indices of willow grouse Lagopus lagopus and ptarmigan L. mutus from the previous autumn, and by December temperatures. Hence, our studies do not support the predation hypothesis for grouse cycles.     

Prospecting in a solitary breeder: chick production elicits territorial intrusions in common loons

In many species, young animals learn about various breedingpatches in one year and use what they have learned to settlein a promising patch the next. Common loons (Gavia immer) seemgood candidates for such prospecting as prebreeders and displacedbreeders intrude frequently into breeding territories defendedby monogamous pairs yet engage in no extrapair copulations.We tested 3 hypotheses for prospecting in loons. The permanentattributes hypothesis gained little support as we found no consistentdifferences in quality between territories and no physical orbiotic trait that predicted reproductive success in a territory.We found some support for the conspecific attraction hypothesisas intruders were attracted to conspecifics in a lake in theshort term; however, intrusions were not more frequent in territoriesthat had experienced regular use by a pair the previous yearthan in territories that had previously been vacant. Instead,the increase in intrusion rate after a year of chick productionsupported the habitat-copying hypothesis, which states thatfloaters use the presence of chicks as a cue to target territoriesfor future attempts at territorial takeover. Despite this systemof prospecting, founding of new territories was common. Onestriking finding was the tendency of territorial breeders toconceal chicks from flying intruders, perhaps to avoid futureterritorial takeover.     

Couch potatoes do better: Delayed dispersal and territory size affect the duration of territory occupancy in a monogamous mammal

In territorial, socially monogamous species, the establishment and defense of a territory are an important strategy to maximize individual fitness, but the factors responsible for the duration of territory occupancy are rarely studied, especially in long‐lived mammals. A long‐term monitoring program in southeast Norway spanning over 18 years allowed us to follow the individual life histories of Eurasian beavers (Castor fiber) from adolescence in their natal family group to dispersal and territory establishment until the end of territory occupancy. We investigated whether territory size, resource availability, population density, and dispersal age could explain the duration of territory occupancy, which ranged from 1 to 11 years. The duration of territory occupancy was positively related to dispersal age, suggesting that individuals that delayed dispersal had a competitive advantage due to a larger body mass. This is in support with the maturation hypothesis, which states that an animal should await its physical and behavioral maturation before the acquisition of a territory. Further, we found that individuals that established in medium‐sized territories occupied them longer as compared to individuals in small or large territories. This suggests that large territories are more costly to defend due to an increased patrolling effort, and small territories might not have sufficient resources. The lifetime reproductive success ranged from zero to six kits and generally increased with an increasing duration of territory occupancy. Our findings show the importance of holding a territory and demonstrate that dispersal decisions and territory selection have important consequences for the fitness of an individual.     

Territory Occupancy and Parental Quality as Proxies for Spatial Prioritization of Conservation Areas

In order to maximize their fitness, individuals aim at choosing territories offering the most appropriate combination of resources. As population size fluctuates in time, the frequency of breeding territory occupancy reflects territory quality. We investigated the relationships between the frequency of territory occupancy (2002–2009) vs. habitat characteristics, prey abundance, reproductive success and parental traits in hoopoes Upupa epops L., with the objective to define proxies for the delineation of conservation priority areas. We predicted that the distribution of phenotypes is despotic and sought for phenotypic characteristics expressing dominance. Our findings support the hypothesis of a despotic distribution. Territory selection was non-random: frequently occupied territories were settled earlier in the season and yielded higher annual reproductive success, but the frequency of territory occupancy could not be related to any habitat characteristics. Males found in frequently occupied territories showed traits expressing dominance (i.e. larger body size and mass, and older age). In contrast, morphological traits of females were not related to the frequency of territory occupancy, suggesting that territory selection and maintenance were essentially a male''s task. Settlement time in spring, reproductive success achieved in a given territory, as well as phenotypic traits and age of male territory holders reflected territory quality, providing good proxies for assessing priority areas for conservation management.     

Complex effects of habitat loss on Golden Eagles Aquila chrysaetos

The development of commercial forests presents potential threats to large raptors that rely on prey caught in open country. We examined the effect of afforestation of breeding habitat used by a population of Golden Eagles Aquila chrysaetos in Scotland where, over the last 50 years, extensive stands of exotic conifers have been planted. Using data for 31 years on territory occupancy and breeding success, together with spatiotemporally dynamic mapping of forest cover and predicted areas of territory‐use in a Geographical Information System, we examined relationships between forest cover and Eagle ecology at landscape and individual territory scales. Several territories were abandoned during the earliest phases of forest planting, but relatively few were apparently lost to later plantings. Territories with poorer breeding productivity appeared to be more vulnerable to abandonment than territories with better breeding productivity. At the landscape scale, temporal differences in breeding productivity were negatively related to the extent of forest cover, although productivity of individual territories showed no clear relationship with forest cover. Several territories with less than a 5% increase in forest cover experienced reduced productivity; however, territories least constrained by neighbouring pairs of Eagles showed an increase in productivity. Territories experiencing the greatest increases in forest cover showed a greater use of spatially separated nest‐sites by occupying pairs. Hence, pairs that were less constrained by neighbours appeared to compensate for loss of open habitat by shifting their territory‐use, whereas pairs that were more constrained could not compensate for open habitat loss and suffered reduced productivity (and, probably in some cases, abandoned the territory). We suggest that simple guidelines based on the extent and locations of habitat loss are inadequate when predicting effects on large territorial raptors such as Golden Eagles. Consideration should also be given to the ‘quality’ of a territory or occupying pair, as well as the extent to which territory‐use is constrained by neighbouring pairs or other ‘unsuitable habitat’ which may have been affected by previous episodes of open habitat loss.     

Winter territoriality and its implications for the breeding ecology of White-throated Dippers Cinclus cinclus

Capsule: Pairs of White-throated Dippers Cinclus cinclus which defended winter territories bred earlier than non-territorial individuals, but there was no difference in reproductive success.

Aims: The effect of winter territoriality on breeding ecology has rarely been studied in resident birds. We carried out a preliminary investigation of whether winter territorial behaviour and territory size affect the timing of reproduction, breeding territory size and reproductive success in a riverine bird, the White-throated Dipper.

Methods: We monitored an individually marked population of White-throated Dippers in the UK. Wintering individuals were classified as either territorial or ‘floaters’ according to their patterns of occurrence and behaviour, and their nesting attempts were closely monitored in the subsequent months. Winter and breeding territory sizes were measured by gently ‘pushing’ birds along the river and recording the point at which they turned back.

Results: All birds defending winter territories did so in pairs, but some individuals changed partners before breeding. Territorial pairs that were together throughout the study laid eggs significantly earlier than pairs containing floaters and those comprising territorial birds that changed partners. However, there were no significant differences in clutch size, nestling mass or the number of chicks fledged. There was no relationship between winter territory length and lay date or any measure of reproductive success, although sample sizes were small. Winter territories were found to be significantly shorter than breeding territories.

Conclusion: Winter territoriality may be advantageous because breeding earlier increases the likelihood that pairs will raise a second brood, but further study is needed. Territories are shorter in winter as altitudinal migrants from upland streams increase population density on rivers, but this may also reflect seasonal changes in nutritional and energetic demands.     

Application of a predictive model to detect long-term changes in nest-site selection in the Bearded Vulture Gypaetus barbatus: conservation in relation to territory shrinkage

In this study we examined long-term variation in the selection of nesting cliffs for the recovering population of the Bearded Vulture Gypaetus barbatus inhabiting the Spanish Pyrenees. We focussed on variables indicating a high probability of cliff occupancy as determined by a previously published model. Although the breeding population increased from 53 to 93 territories between 1991 and 2002, the breeding range expanded only slightly. New and old nesting cliffs had similar habitat features in relation to topography, altitude and degree of human influence, but the distance between occupied cliffs was reduced (from 11.1 to 8.9 km). Thus the probabilities of occupation predicted by the model were lower for newly colonized locales. Our study shows that territory compression may occur without serious modification of nesting habitat quality. These results may arise from the lack of strong territorial behaviour by Bearded Vultures and the availability of high-quality cliffs. The relatively low quality of sites in adjacent mountains may prevent the expansion of the breeding range, but conspecific attraction may also play a role. Our study confirms that monitoring changes in key variables important to habitat selection is useful in determining long-term trends in settlement patterns in heterogeneous environments. The results also suggest that the available nest-site selection model may accurately predict cliff occupancy by Bearded Vultures in those areas where the distance to the nearest neighbour is not a limiting factor. In particular, the model may be useful in establishing priority areas for reintroduction.