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1.
We studied nesting habitat selection of the endangered non‐migratory Osprey Pandion haliaetus population of the Canary Islands and evaluated the effect of human expansion in recent decades. Compared with randomly selected potential nest‐sites, Osprey nests were more frequently found on taller, southwest‐facing cliffs, characterized by lower human pressure and closer to Yellow‐legged Gull Larus michahellis colonies and Barbary Falcon Falco pelegrinoides breeding sites. Furthermore, changes in some breeding habitat features have been detected in recent decades. According to our predictive models, large areas of suitable habitat are available but unoccupied in the Canaries, and human activities are probably limiting the settlement and dispersion of new pairs.  相似文献   

2.
We studied factors affecting breeding habitat selection in a population of cliff-nesting peregrines Falco peregrinus across multiple spatial levels (cliff site, cliff context and land-use of the surrounding landscape), over a 2,100 km2 study area in the Alps (Italy and Switzerland). We detected 30 breeding pairs (density: 1.43 territorial pairs/100 km2), whose territories were uniformly distributed over the study area. We compared 15 habitat features characterising occupied cliffs and 30 randomly selected unoccupied cliffs by means of stepwise forward logistic regression and hierarchical partitioning. The logistic regression analysis showed that occupied cliffs were longer (horizontal length), steeper, and had a greater extension of urban areas in the surrounding landscape compared to non-occupied cliffs. The model had a discrimination ability of 0.95. Hierarchical partitioning indicated that the logistic regression model was appropriate. Neither the distance of cliffs to the nearest site occupied by a potential competitor and predator, the eagle owl Bubo bubo, which occurred at a low density (0.67 pairs/100 km2), nor the distance to the nearest site occupied by peregrines had any effect on the cliff suitability model. Therefore, habitat selection in cliff-nesting peregrines was mainly influenced by cliff site features and surrounding landscape characteristics, whereas the proximity to conspecifics and to an intraguild predator had no apparent effects on peregrine settlement in our study population.  相似文献   

3.
Conspecific nesting density affects many aspects of breeding biology, as well as habitat selection decisions. However, the large variations in breeding density observed in many species are yet to be fully explained. Here, we investigated the settlement patterns in a colonial species with variable breeding density and where resource distribution could be manipulated. The zebra finch, Taeniopygia guttata, is a classic avian model in evolutionary biology but we know surprisingly very little about nest site selection strategies and nesting densities in this species, and in fact, in nomadic species in general. Yet, important determinants of habitat selection strategies, including temporal predictability and breeding synchrony, are likely to be different in nomadic species than in the non‐nomadic species studied to date. Here, we manipulated the distribution of nesting sites (by providing nest boxes) and food patches (feeders) to test four non‐exclusive habitat selection hypotheses that could lead to nest aggregation: 1) attraction to resources, 2) attraction to breeding conspecifics, and 3) attraction to successful conspecifics and 4) use of private information (i.e. own reproductive success on a site). We found that wild zebra finches used conspecific presence and possibly reproductive success, to make decisions over where to locate their nests, but did not aggregate around water or food within the study areas. Moreover, there was a high degree of inter‐individual variation in nesting density preference. We discuss the significance of our results for habitat selection strategy in nomadic species and with respect to the differential selection pressures that individuals breeding at different densities may experience.  相似文献   

4.
Griffon vulture (Gyps fulvus) population surveys were conducted during 1996–2002 in the island of Crete (Greece) to document population status and structure. Fieldwork was carried out during the breeding period when birds could be monitored in their colonies. Total population size was estimated at 379 individuals (range = 341–417) with adult birds comprising 63%. The breeding population was estimated at 141 pairs, which were distributed on an average in 23 colonies per year (range = 16–30) while the mean number of breeding pairs that laid eggs was 98 (range= 64–126). Crete thus supports the largest insular population of the species in the world and hosts 70–80% of the breeding population of the species in Greece. Population density was estimated at 6.9 individuals/100 km2, 2.6 breeding pairs/100 km2 and 1.8 nesting pairs/100 km2. The average home range of an occupied colony (i.e., breeding group) was estimated at ca. 204 km2 producing a theoretical foraging range of 8 km radius around the breeding cliff. No trends in the total number of individuals and breeding pairs appeared to exist, although significant differences in population size of individual colonies occurred between the years. The majority of the population was concentrated in small-sized colonies, which showed a low occupancy rate. The number of abandoned sites and the colonization of new ones could represent a shift of breeding pairs to alternative colonies provoked by local food abundance and conspesific attraction.  相似文献   

5.
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.  相似文献   

6.
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.  相似文献   

7.
Species ranges often change in relation to multiple environmental and demographic factors. Innovative behaviors may affect these changes by facilitating the use of novel habitats, although this idea has been little explored. Here, we investigate the importance of behavior during range change, using a 25‐year population expansion of Bonelli's eagle in southern Portugal. This unique population is almost exclusively tree nesting, while all other populations in western Europe are predominantly cliff nesting. During 1991–2014, we surveyed nest sites and estimated the year when each breeding territory was established. We approximated the boundaries of 84 territories using Dirichlet tessellation and mapped topography, land cover, and the density of human infrastructures in buffers (250, 500, and 1,000 m) around nest and random sites. We then compared environmental conditions at matching nest and random sites within territories using conditional logistic regression, and used quantile regression to estimate trends in nesting habitats in relation to the year of territory establishment. Most nests (>85%, n = 197) were in eucalypts, maritime pines, and cork oaks. Nest sites were farther from the nests of neighboring territories than random points, and they were in areas with higher terrain roughness, lower cover by agricultural and built‐up areas, and lower road and powerline densities. Nesting habitat selection varied little with year of territory establishment, although nesting in eucalypts increased, while cliff nesting and cork oak nesting, and terrain roughness declined. Our results suggest that the observed expansion of Bonelli's eagles was facilitated by the tree nesting behavior, which allowed the colonization of areas without cliffs. However, all but a very few breeding pairs settled in habitats comparable to those of the initial population nucleus, suggesting that after an initial trigger possibly facilitated by tree nesting, the habitat selection remained largely conservative. Overall, our study supports recent calls to incorporate information on behavior for understanding and predicting species range shifts.  相似文献   

8.
Determining habitat quality for wildlife populations requires relating a species' habitat to its survival and reproduction. Within a season, species occurrence and density can be disconnected from measures of habitat quality when resources are highly seasonal, unpredictable over time, and patchy. Here we establish an explicit link among dynamic selection of changing resources, spatio‐temporal species distributions, and fitness for predictive abundance and occurrence models that are used for short‐term water management and long‐term restoration planning. We used the wading bird distribution and evaluation models (WADEM) that estimate (1) daily changes in selection across resource gradients, (2) landscape abundance of flocks and individuals, (3) conspecific foraging aggregation, and (4) resource unit occurrence (at fixed 400 m cells) to quantify habitat quality and its consequences on reproduction for wetland indicator species. We linked maximum annual numbers of nests detected across the study area and nesting success of Great Egrets (Ardea alba), White Ibises (Eudocimus albus), and Wood Storks (Mycteria americana) over a 20‐year period to estimated daily dynamics of food resources produced by WADEM over a 7490 km2 area. For all species, increases in predicted species abundance in March and high abundance in April were strongly linked to breeding responses. Great Egret nesting effort and success were higher when birds also showed greater conspecific foraging aggregation. Synthesis and applications: This study provides the first empirical evidence that dynamic habitat selection processes and distributions of wading birds over environmental gradients are linked with reproductive measures over periods of decades. Further, predictor variables at a variety of temporal (daily‐multiannual) resolutions and spatial (400 m to regional) scales effectively explained variation in ecological processes that change habitat quality. The process used here allows managers to develop short‐ and long‐term conservation strategies that (1) consider flexible behavioral patterns and (2) are robust to environmental variation over time.  相似文献   

9.
Nesting is a critical yet hazardous life stage for many birds. For colonial‐breeding birds, the conspicuousness of the colony to predators suggests immense pressure to select optimal colonial nesting sites. But what drives selection of those sites? As with solitary nesting birds, reducing access by predators may be the single most important factor. If so, knowledge of the predators involved and the attributes of different potential colony sites can allow us to predict the features that make a site especially safe. We examined the attributes of trees used by breeding colonies of metallic starlings Aplonis metallica in tropical Australia, and experimentally tested if those attributes prevented nest access by predatory snakes. Our surveys confirmed that tree choice by starling nesting colonies is highly non‐random, with all colonies located in tall trees in rainforest clearings, with no low branches and smooth bark. Experimental tests demonstrated that the climbing ability of predatory snakes depends upon bark rugosity, and that colony access by snakes depends on tree attributes such as bark rugosity and canopy connectivity. Our study confirms that colonial‐nesting starlings select colony sites that provide a safe refuge from predation. Intense predation pressure may have driven the evolution of stringent breeding habitat criteria in many other species of colonial‐breeding birds.  相似文献   

10.
An influential period in avian life‐cycles is the annual breeding season, when competition over suitable nesting sites and territories is a key factor that can determine fitness and distribution, especially for species that are highly selective in their nesting habitats. We analysed nest‐site characteristics, breeding success and competitive interactions between two apex predator populations. Whereas the Short‐toed Eagle Circaetus gallicus has nested in the Judean Foothills (Israel) for a long time, the Long‐legged Buzzard Buteo rufinus has only invaded the nesting habitat of the Short‐toed Eagle during their breeding season in the last two decades. These two recently sympatric species have similar nesting ecology and frequently use the same nests. They are therefore expected to compete over nesting sites and territories. We analysed interspecific interactions between these two species by combining information from comprehensive observational, experimental, GIS analysis and remote sensing data, deriving 65 variables to characterize the nest‐sites used and the breeding success in 381 breeding attempts over four consecutive breeding seasons. To assess interspecific and intraspecific territorial behaviour and aggressiveness, stuffed Long‐legged Buzzards and Short‐toed Eagles were presented close to nests. Nest‐site characteristics overlapped substantially between species, and Long‐legged Buzzards occupied 21% of all Short‐toed Eagle nests. Intraspecific aggression rates among Long‐legged Buzzards were higher than their interspecific aggression rates with Short‐toed Eagles and also higher than intraspecific aggression among Short‐toed Eagles. Long‐legged Buzzard and Short‐toed Eagle breeding densities (1.59 ± 0.11 and 2.96 ± 0.11 pairs per 10 km2, respectively) are likely to be the highest across their respective breeding distributions, with a maximum productivity of 0.96 ± 0.01 and 0.56 ± 0.05 (young fledged/breeding pair) for Long‐legged Buzzard and Short‐toed Eagle, respectively. Intraspecific interactions among both species play an important role in determining their breeding success and the spatial distribution of nesting sites. Our results suggest that interspecific competition over nesting sites and territories between both species, and the potential dominance of Long‐legged Buzzard, has both direct and indirect impacts on the spatial and demographic distribution of Short‐toed Eagles due to the recent establishment of Long‐legged Buzzard territories in the Judean breeding area.  相似文献   

11.
The Demoiselle crane population was studied in the southern Chelyabinsk region (52°12′N; 60°21′E) during 1988–2008. Counts of nesting pairs were annually conducted over an area of 100–130 km2 in May. The number of breeding pairs varied from 6.5 to 9.5 per 100 km2; on average, there were 8 pairs per 100 km2 during 1989–1991. In 2000–2008, the population density increased from 7.5 to 12 pairs per 100 km2, on average, 9.3 pairs per 100 km2. Demoiselle crane nesting in steppe pastures and perennial grass fields has been rarely observed for the last decade. Now the main nesting biotopes of the species are stubble fields accounting for up to 70% of all nidification cases documented. The decline in agricultural production in recent decades has resulted in the appearance of long-fallow lands, in which 17 to 35% crane pairs nest.  相似文献   

12.
Jouke Prop  John L. Quinn 《Oikos》2003,102(3):571-580
In this paper we aim to explain the distribution of red-breasted geese Branta ruficollis over different nesting habitats. To be safe from land predators red-breasted goose colonies were restricted to i) islands on rivers, ii) cliffs with peregrine falcons Falco peregrinus , and iii) the close proximity of snowy owl Nyctea scandiaca and rough-legged buzzard Buteo lagopus nests. Among years nest site availability varied by fluctuations in numbers of owls and buzzards in association with cycles in lemming abundance, but the total number of goose nests found in the study area did not vary. The distribution of geese, in combination with data on reproductive success, suggested a despotic mechanism: at cliffs, goose numbers were constant among years with an invariably high reproductive success, whereas large fluctuations in numbers on islands coincided with opposite trends in success. Apparently, geese nesting with owls or buzzards moved to the few islands present in the study area during years when these birds of prey were absent. Consequently, in such years the average density of geese on islands was more than twice as high as at cliff colonies (5.4 and 2.3 pairs per ha of foraging habitat, respectively). Colony size at cliffs may have been restricted by territorial behaviour of the geese, though there is evidence that, additionally, the host falcons also limited the number of nesting geese. Apparently rare in closely related species, we observed a negative density-dependent effect on reproductive success during the nest phase, and attribute this to limited food resources, reinforced by the high frequency of territorial interactions. This leads to the conclusion that, in addition to predation pressure, nesting density is an important agent in the link between lemming cycles and goose breeding success.  相似文献   

13.
Some birds use social cues, such as the presence of conspecifics, when selecting breeding habitat. This phenomenon, known as conspecific attraction, has been well‐documented in migratory species, but has not been assessed for resident species of birds. We used Dupont's Larks (Chersophilus duponti) as a model species to determine if conspecific attraction plays a role in habitat selection by resident species of birds. At our study site in Soria province in central Spain, we monitored two potential habitat patches and one managed site where management actions had provided apparently suitable habitat. At each site, we broadcast recordings of the songs and calls of male Dupont's Larks, and monitored their presence during the breeding season and dispersal period in 2018 using automated recorders and field surveys. No birds were attracted to our study sites. Our results suggest that management of patches of suitable habitat should occur close to areas (within 1 km) already occupied by Dupont's Larks to encourage natural colonization because, based on our results, playback of conspecific vocalizations may not attract the species to new breeding areas. However, additional studies are needed before drawing conclusions about the effectiveness of conspecific attraction for this and other resident species of birds.  相似文献   

14.
Habitat selection is a complex process, that is affected by several factors, including habitat characteristics, environmental conditions, and both intra‐ and interspecific interactions. We analysed habitat preferences of two top avian predators, Peregrine Falcon Falco peregrinus, a medium‐sized diurnal raptor, and Eagle Owl Bubo bubo, a large nocturnal raptor. These two species are known to compete for preferred nest‐sites, and proximity to cliffs with Eagle Owls may reduce Peregrine breeding output through predation of young Falcons. We investigated the environmental factors affecting occurrence and coexistence of the two species and the potential role of habitat suitability in favouring co‐occurrence in 3519 km2 of the central pre‐Alps of Italy, where the two species breed on cliffs and sometimes co‐occur on the same cliff. Peregrines settled on long, steep and favourably orientated cliffs in woodland landscapes close to urban areas. Eagle Owls settled on topographically similar cliffs, but in lower rainfall areas compared with cliffs occupied by Peregrines and cliffs unoccupied by either species. Sites where the two species co‐occurred were characterized by more horizontally extended cliffs compared with sites of exclusive occurrence of each species. An analysis of relative habitat suitability revealed that sites where the two species co‐occurred had the highest predicted probability of occupancy for both species, suggesting that those sites should be regarded as high‐quality sites. Breeding productivity of Eagle Owls was negatively affected by the co‐occurrence of Peregrines, whereas the effect of Eagle Owl proximity on Peregrine productivity varied according to cliff suitability for the Peregrines. Habitat selection had fitness consequences for Eagle Owls because breeding productivity increased with cliff length. Environmental conditions, particularly climatic factors, could allow the widespread coexistence of these competing raptors at the landscape scale, whereas at the local scale co‐occurrence could take place only on larger cliffs. These were preferred sites for both species, presumably because breeding at such sites offsets the costs of settling close to the competitor species.  相似文献   

15.
Animals are expected to select a breeding habitat using cues that should reflect, directly or not, the fitness outcome of the different habitat options. However, human‐induced environmental changes can alter the relationships between habitat characteristics and their fitness consequences, leading to maladaptive habitat choices. The most severe case of such nonideal habitat selection is the ecological trap, which occurs when individuals prefer to settle in poor‐quality habitats while better ones are available. Here, we studied the adaptiveness of nest box selection in a tree swallow (Tachycineta bicolor) population breeding over a 10‐year period in a network of 400 nest boxes distributed along a gradient of agricultural intensification in southern Québec, Canada. We first examined the effects of multiple environmental and social habitat characteristics on nest box preference to identify potential settlement cues. We then assessed the links between those cues and habitat quality as defined by the reproductive performance of individuals that settled early or late in nest boxes. We found that tree swallows preferred nesting in open habitats with high cover of perennial forage crops, high spring insect biomass, and high density of house sparrows (Passer domesticus), their main competitors for nest sites. They also preferred nesting where the density of breeders and their mean number of fledglings during the previous year were high. However, we detected mismatches between preference and habitat quality for several environmental variables. The density of competitors and conspecific social information showed severe mismatches, as their relationships to preference and breeding success went in opposite direction under certain circumstances. Spring food availability and agricultural landscape context, while related to preferences, were not related to breeding success. Overall, our study emphasizes the complexity of habitat selection behavior and provides evidence that multiple mechanisms may potentially lead to an ecological trap in farmlands.  相似文献   

16.
Pink-footed geese Anser brachyrhynchus nest in two contrasting but commonly found habitats: steep cliffs and open tundra slopes. In Svalbard, we compared nest densities and nesting success in these two environments over ten breeding seasons to assess the impact of spring snow cover, food availability to nesting adults and arctic fox Vulpes lagopus (main terrestrial predator) abundance. In years with extensive spring snow cover, fewer geese at both colonies attempted to breed, possibly because snow cover limited pre-nesting feeding opportunities, leaving adults in poor breeding condition. Nesting success at the steep cliff colony was lower with extensive spring snow cover; such conditions force birds to commit to repeated and prolonged recess periods at far distant feeding areas, leaving nests open to predation. By contrast, nesting success at the open tundra slope was not affected by spring snow cover; even if birds were apparently in poor condition they could feed immediately adjacent to their nests and defend them from predators. Foxes were the main nest predator in the open tundra slopes but avian predators likely had a larger impact at the steep cliffs colony. Thus, the relative inaccessibility of the cliffs habitat may bring protection from foxes but also deprives geese from readily accessing feeding areas, with the best prospects for successful nesting in low spring snow cover years. Our findings indicate that spring snow cover, predator abundance and food proximity did not uniformly influence nesting success of this herbivore, and their effects were dependent on nesting habitat choice.  相似文献   

17.
Much of the breeding range for the mountain plover (Charadrius montanus) occurs in shortgrass steppe and mixed-grass prairie in the western Great Plains of North America. Studies of mountain plovers in shortgrass steppe during the 1970s and 1990s were focused in Weld County, Colorado, which was considered a key breeding area for the species. These studies, however, did not include habitats influenced by black-tailed prairie dogs (Cynomys ludovicianus) or prescribed fire. The role of these 2 rangeland disturbance processes has increased substantially over the past 15 years. During 2008–2009, I used radial distance point count surveys to estimate mountain plover densities early in the nesting season in 4 habitats on public lands in Weld County, Colorado. All 4 habitats were grazed by cattle during the growing season at moderate stocking rates but had different additional disturbances consisting of 1) dormant-season prescribed burns, 2) active black-tailed prairie dog colonies, 3) black-tailed prairie dog colonies affected by epizootic plague in the past 1–2 years, and 4) rangeland with no recent history of fire or prairie dogs. Mountain plover densities were similar on active black-tailed prairie dog colonies ( = 6.8 birds/km2, 95% CI = 4.3–10.6) and prescribed burns ( = 5.6 birds/km2, 95% CI = 3.5–9.1). In contrast, no plovers were detected at randomly selected rangeland sites grazed by cattle but lacking recent disturbance by prairie dogs or fire, even though survey effort was highest for this rangeland habitat. Mountain plover densities were intermediate (2.0 birds/km2, 95% CI = 0.8–5.0) on sites where black-tailed prairie dogs had recently been extirpated by plague. These findings suggest that prescribed burns and active black-tailed prairie dog colonies may enhance breeding habitat for mountain plovers in shortgrass steppe and illustrate the potential for suppressed or altered disturbance processes to influence habitat availability for declining wildlife species. © 2011 The Wildlife Society.  相似文献   

18.
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.  相似文献   

19.
20.
Nest defence is a fundamental aspect of parental care in secondary cavity‐nesting birds, and predation or competition for nesting sites can involve different defensive behaviours. Because habitat quality determines breeding success, we were interested in whether breeding pairs of the Eurasian nuthatch, Sitta europaea, established in more favourable environment also manifest higher probability of cooperative behaviour during their nest‐site defence. To explore this relationship, we quantified behavioural displays of both parents and analysed activity budget ethogram data from simulated territorial intrusions performed in the chick‐feeding phase with one conspecific and two different heterospecific stimuli (dummies of nuthatch, starling and woodpecker). We found that paired individuals shared their roles during nest‐site defence to a considerable extent. Males had a significantly higher number of attacks on intruders than females, and females performed more threat displays and controls of the brood than males. Multinomial analysis of the cooperative behaviour suggested that pairs in a high‐quality territory had higher probability of reciprocal substitution of different roles towards a balance between attacks, threat displays and nest controls. Contrary to this, pairs in a low‐quality territory had less likely pairwise combinations of simultaneous behavioural states that are associated with effective nest‐site defence. The difference in response probability according to territory quality was, however, highly variable in view of the stimulus that was used in simulated territorial intrusion. Because individual roles and the complex behavioural repertoire of pairs altered in response to territory quality and potential nest‐site competitor or brood predator, our results suggest that the cooperative nest‐defence behaviour could be linked to the breeding success of this year‐round territorial species living in a heterogeneous forest habitat.  相似文献   

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