Landscape and anti-predation determinants of nest-site selection, nest distribution and productivity in a Mediterranean population of Long-eared Owls Asio otus

Nest predation is an important determinant of owl breeding success. We studied Long-eared Owl Asio otus productivity and attributes of nest-sites at the microhabitat and landscape scales in a Mediterranean locality over an 8-year period. We examined the effect on nest location and productivity of protective cover in concealing the nest from aerial and terrestrial predators. A dense cover of ivy and tree-foliage at canopy level favoured nest location but not productivity. By contrast, high shrub cover beneath the nest was selected by Owls and was positively related to both the site reoccupancy rate and the overall number of young fledged. Pre-fledging Owls use the ground, where they are exposed to terrestrial predators, which are much more abundant in the study area than are aerial predators. Our results therefore support the hypothesis that Owls adapt nest-site choice to local sources of predation risk. As reported elsewhere, Long-eared Owls in our study area showed restricted territoriality and nested in clusters. As active nest-sites during the same breeding season were more than 1 km apart on average, and their productivity was never greater for clustered nests than for more isolated nests, nest aggregation could not be interpreted as a case of facultative colonial breeding, which has been reported for this species in other areas. Neither landscape variables indicative of the availability of foraging areas nor structural attributes that protect young from predators explained the remarkable scarcity of nests in half of the study area. Unmeasured factors such as human disturbance could explain the pattern of distribution of Long-eared Owl nests.     

Weight Changes of Dunlins on the Dee Estuary in May

In an attempt to increase the number of breeding pairs of Long-eared Owls at Woodwalton Fen National Nature Reserve, wicker baskets of varying design were placed throughout the reserve. Between 1981 and 1996, owls used the baskets on 77 occasions. Of 71 nests followed from incubation, 59.2% hatched young and 50.7% fledged one or more chicks. In only two nests were the complete clutches hatched and fledged. Early nesters laid larger clutches and raised more broods than those nesting later in the season. It is suggested that the level of the water table and the management regime necessary for the restoration of open fen communities affect the number of breeding owls.     

Nest Defence of Nesting Chinstrap Penguins (Pygoscelis antarctica) against Intruders

We tested some predictions of parental investment theory by studying the aggressive behaviour of colonial nesting chinstrap penguins (Pygoscelis antarctica) against human intruders into their nesting territories. We tested for differences in the aggressive behaviour of penguins according to offspring age (eggs vs. chicks), offspring number, nest location in the colonies (central vs. peripheral) and sex. Offspring age was the main factor influencing nest defence, although nest location and sex were also important. Chicks were defended more strongly than eggs, in accordance with changes in the reproductive value of offspring, and this increase in aggressiveness was not related to revisitation of the same individuals. The level of aggression of penguins breeding in central sites was higher than that of peripheral birds, a difference that could be due to the lower residual reproductive value of central-nesting, probably older, birds. The stronger aggressiveness of males could be due to a combination of factors related to sexual selection and life-history traits. Offspring number did not affect the level of nest defence.     

Habitat preferences for Long-eared Owls Asio otus and Little Owls Athene noctua in semi-arid environments at three spatial scales

Capsule There is a relationship between owl numbers and the availability of the agri-forest patchwork.

Aims To model habitat preferences at three different scales of two predators largely neglected within the framework of Environmental Impact Assessment (EIA) studies.

Methods We studied habitat preferences of Long-eared Owls and Little Owls by comparing habitat composition around 28 and 78 occupied territories respectively with 55 non-occupied territories in Alicante (eastern Spain). Generalized linear models were used to examine patterns of habitat preference at three different spatial scales: nest-site, home range and landscape.

Results At the nest-site scale, Long-eared Owls preferred wooded areas with few paved roads while Little Owls preferred arid plantations. Furthermore, the probability of finding an occupied territory increased with the proximity of another occupied territory in the surroundings. The home range scale models mirror the feeding requirements of the owls. Thus, Long-eared Owls occupied areas with high percentages of forest, arid plantations, edges between these two land uses, short distances between nests, with presence of conspecifics and little human disturbance. Little Owls occupied arid plantations with high availability of linear structures and the proximity of villages. At the landscape scale, Long-eared Owls eluded extensive forests, and Little Owls preferred arid plantations.

Conclusions We suggest a hierarchical process of habitat selection for both owls regarding fitting trophic resources at the broadest scales and adequate sites for breeding and roosting at the smallest scale. EIA studies must consider that protecting small areas around single nests may not be an efficient conservation option compared with preserving clusters of territories for both species.     

Correlations between body size, defensive behaviour and reproductive success in male Little Blue Penguins Eudyptula minor: implications for female choice

Field studies exploring relationships among body size, defensive behaviour (including vocalizations) and reproductive success in male Little Blue Penguins Eudyptula minor were conducted to identify behavioural and physical attributes that might provide females or opponents with information on male quality. Head length provided a simple estimator of body size in both sexes. Nests defended by large males produced eggs and chicks earlier in the breeding season, and contained chicks that grew more quickly than chicks in nests defended by small males. In contrast, female body size did not affect any of the reproductive parameters we measured. The pitch of male defensive calls varied significantly with male size, so females (or opponents) could potentially gauge male size by assessing the calls. Large males were also more likely to respond vocally to simulated nest intrusions. As a result, by 'keeping company' with prospective partners early in the breeding season, females can obtain information about males that would allow them to select better partners and improve their own reproductive potential.     

A molecular genetic analysis of the communal nesting of the ostrich (Struthio camelus)

The ostrich breeding system is complex and unique; communal clutches are laid by several females, although only one female, the major female, and the resident territorial male provide parental care. More eggs are laid in the nest than can be incubated and the major female ejects surplus eggs from the incubated central clutch. Microsatellite markers were used to analyse the parentage of communal nests in Nairobi National Park. This revealed that major females contributed a disproportionate number of fertile eggs to the central, incubated clutch and that multiple paternity and maternity within a nest were common; 68.9% of all incubated eggs on a nest were not parented by both the resident territorial male and the major female of that nest. All the males fertilized eggs on the clutches of neighbouring males. Unexpectedly, every major female with her own nest was also simultaneously a minor female with incubated eggs on neighbouring clutches. The relatedness between females laying in the same nest was not significantly different from the population average and significantly less than that between chicks hatched from the same nest.     

The nest defence by the red‐backed shrike Lanius collurio – support for the vulnerability hypothesis

The majority of altricial bird species defend their brood against predators more intensively in nestlings rather than eggs stage. Several hypotheses have been proposed to explain this difference. The majority of existing experimental studies have recorded a gradually increasing intensity of nest defence supporting the reproductive value hypothesis. We have compared nest defence in two nesting stages of the red‐backed shrike against two predators of adult birds and against two predators of nests. While the nests with nestlings were defended by parents against three out of four predators, nests with eggs were almost not defended at all. This rapid change in parent nest defence supports rather the vulnerability hypothesis, predicting that the threat to nests with nestlings increases rapidly after hatching, as they became more conspicuous due to their begging and parental provisioning. Unlike most of the species tested previously, the red‐backed shrike uses very vigorous mobbing towards predators. We suggest that the occurrence of this active mobbing (strikes, including physical contact) is a good proxy of the current threat to the nest.     

Aspects of the Social Organisation of the European Wild Rabbit (Oryctolagus cuniculus)

The social organisation of a free-living population of wild rabbits was examined. Discrete breeding groups were recognised which consisted of individuals who shared access to underground refuges. Males attempted to maximise their access to females by a) occupying large home ranges; b) defending female access indirectly through territorial defence; c) defending female access directly within groups. Females defended nest site access within groups. Intergroup interactions between females were rare because of limited home range overlap. Interruption of female-female aggression by males was observed suggesting that conflicts of interest arose between the sexes. Adult-juvenile aggression was primarily intra-sexual; females initiated more interactions than males who adopted a protective role in some cases.     

When do altricial birds reach maximum of their brood defence intensity?

It has been suggested that the brood defence by parents of altricial birds should increase during the breeding attempt until the young depart from the nest. The two proximate hypotheses provide alternative predictions about the peak of brood defence intensity: (1) the vulnerability hypothesis predicts a rapid rise in brood defence after hatching of the chicks, with maximum defence intensity just before fledging and strong decline afterwards; (2) the feedback hypothesis predicts that brood defence intensity will, after a rapid rise, reach a plateau at the end of the nestling period and early after fledging and then slowly decline. I compared brood defence behaviour of altricial meadow pipit (Anthus pratensis) breeding in the Czech Republic during the late nestling stage and during the fledging time. A stuffed stoat (Mustela erminea) was placed 5 m from a meadow pipit nest and the defence behaviour of parents was recorded for 10 min from a hide. Brood defence intensity was higher during the fledgling time than during the late nestling stage, and this trend was more evident in males than in females. Regardless of the proportion of already fledged chicks and those still present in the nest, brood defence did not significantly decrease during the fledgling time in males or females. The results do not agree with the predictions of the vulnerability hypothesis and support the predictions of the feedback hypothesis.     

Male Rock Sparrow (Petronia petronia) Nest Defence Correlates with Female Ornament Size

We investigated the relationship between male nest defence and female breast patch size in an alpine population of rock sparrow (Petronia petronia) in northern Italy. We presented a mounted weasel (Mustela nivalis), a common nest predator, to 28 pairs breeding in nest boxes, with 12–13‐d‐old nestlings, and measured the intensity of male and female defence reaction. We measured the frequency of attack flights, intensity of alarm calling and total time spent in view, and then combined these for each individual, in a single defence factor by principal component analysis. All the females arrived to defend the nest while only 21 males arrived, and females defended the nest more intensely than males. We analysed, by stepwise regression, the relationship of male defence factor to female behaviour and phenotype (breast patch size, a measure of quality) and brood properties (size, mass, phenology). Male defence factor was significantly related only to female breast patch size. We argue that male rock sparrows apparently make parental investment decisions according to their mate's quality, and examine possible alternative hypotheses.     

Sex-differences in nest defence in Fieldfares Turdus pilaris in relation to their size and physical condition

The nest defence of solitary-breeding Fieldfares Turdus pilaris against a human intruder, a corvid and a hawk was studied in a mountain birch forest in central Norway in 1990–2002. Females defended their nests against a human intruder more vigorously than did males before hatching, and vice versa after hatching. Although both sexes increased their nest defence, only males showed a significant increase and showed a greater defence than females at the chick stage. Females with 1-week-old nestlings had longer wings, were heavier and in a better physical condition than those that failed to get nestlings to that stage. Females, but not males, lost condition during nesting. Birds in better condition were more aggressive. The pattern of nest defence by males and females against the corvid was similar to that shown against human intruders, whereas males took greater risks than females towards the raptor. There was weak evidence that birds that were more successful showed greater levels of nest defence. The sex-role reversal in nest defence may be connected to differences in costs in relation to the renesting potential and to the physical condition of the sexes.     

Roadway mortality of barn owls in Idaho,USA

We examined the temporal, spatial, and demographic factors that influenced roadway mortality of barn owls (Tyto alba) along a 248-km stretch of Interstate 84 in southern Idaho using systematic road surveys. Counts of dead animals from surveys can be underestimated because of sampling biases; therefore, we also conducted experiments to assess the effects of search and removal bias on the estimates of roadway mortality of owls. We conducted surveys every 2 weeks over a 2-year period and detected 812 dead barn owls (unadjusted mortality rate of 1.64 owls/km/yr). After adjusting this estimate for search and removal bias, we documented mortality rates of up to 5.99 owls/km/year. Owl mortality was not random in relation to sex, age class, or location along the highway. Females and juveniles, which represent individuals more likely to disperse long distances, were killed more frequently than males and adults. During the nonbreeding season, owls were killed more often near agricultural lands than in shrub-steppe, but this pattern was not apparent during the breeding season. Owls were also killed more often on portions of the roadway closer to the Snake River canyon, perhaps because of the availability of nest and roost sites. Mortality rates differed markedly between the 2 years of study, which could have been related to variability in weather and its subsequent effect on owl productivity. Our data suggest that barn owls in this region may not persist under this level of mortality without significant immigration or management. Thus, roadway management to reduce or prevent owl use of roadways, reduce rodent populations near major roads, alert motorists to the presence of owls, or otherwise reduce the chances that vehicles and owls collide would improve barn owl survival and population persistence. © 2012 The Wildlife Society.     

Are House Wren Troglodytes aedon eggs unusually strong? Test of the predicted effect of intraspecific egg destruction

As a result of opposing selective forces, the external strength of avian eggs should be near some size-specific optimum. However, in certain situations there should be selection on females to lay unusually strong eggs. According to one hypothesis, intraspecific egg destruction should favour increased egg strength as a means of defence against conspecific intruders. This hypothesis predicts that House Wrens Troglodytes aedon , a species well known for its tendency to destroy conspecific clutches, should be under selection for unusually strong eggs. However, the intensity of selection for strong eggs should also be modified by efficacy of nest defence against conspecific intruders in a given species (i.e. efficient nest defence by the breeding pair should weaken selection for unusually strong eggs). The goals of our study were: (1) to establish whether House Wren eggs are stronger than expected for their size; (2) to determine which structural mechanisms are responsible for their unusual strength; and (3) to test a hypothesis that, between wren species, the efficacy of nest defence and the intensity of egg-destroying behaviour affect the intensity of selection for unusually strong eggs. Our results demonstrated that: (1) House Wren eggs are 1.9 times stronger than expected for their size; (2) their unusual strength is achieved mostly by their unusually thick shells; and (3) eggs of the House Wren (extensive paternal nest defence; male egg-destroying behaviour suppressed during incubation) are significantly weaker structurally than eggs of the Marsh Wren Cistothorus palustris (reduced paternal nest defence; male egg-destroying behaviour present throughout incubation). These results are consistent with the hypothesis that the intraspecific egg-destroying behaviour and the efficacy of nest defence by the breeding adults have played a role in the evolution of strength of House Wren eggs.     

Cooperation and conflict among dunnocks, Prunella modularis, in a variable mating system

The mating system of a population of 90 breeding dunnocks (or hedge sparrow, Prunella modularis) included monogamy, polygyny, polyandry and polygynandry. Monogamous males guarded females during their fertile period to prevent neighbouring males from copulating. The most intense guarding occurred where two (unrelated) males shared a territory. Here, the alpha male tried to prevent the beta male from copulating with the female. Beta males were seen to copulate in only half the cases. They were more likely to succeed when the alpha male found it difficult to guard the female closely because her range was large, the vegetation was dense or there were other females breeding synchronously on the same territory. Close guarding and chasing by males reduced the female's feeding rate and was correlated with unhatched eggs in the nest. Females attempted to escape the alpha male's attentions and actively encouraged the beta male to mate. Beta males only helped to feed the young if they copulated with the female. Nestlings fed by two males and a female got more food and weighed more than those fed by just one male and a female. Indirect evidence suggested that when beta males failed to copulate, they destroyed eggs or young chicks. Females laid larger clutches when two males mated with them as opposed to one, thus adapting their clutch size to the amount of parental care they expected. The results of natural removal experiments and matched comparisons of the same female in different mating systems support these conclusions. For females, selection favours cooperative polyandry, whereas for males if favours polygyny; the variable mating system may reflect the different outcomes of this sexual conflict.     

Is female investment in defence correlated with breeding success in Hen Harriers Circus cyaneus

CapsuleThe intensity of parental defence, irrespective of the value of offspring, may be one of the proximate causes of breeding success, reflecting the quality of breeders.

Aims To test whether female Hen Harrier investment in defence towards a human predator influences breeding success.

Methods Generalized linear mixed models were used, with nest content, date of visit, the interaction (nest content*date), breeding success (number of fledglings/clutch size) and presence or absence of male as explanatory variables. ‘Female’ was included as a random variable in the model. Alarm call rate by a female Hen Harrier during the first three minutes after my arrival at a nest was used as a dependent variable (as an indicator of parental investment in defence towards a human predator).

Results Females were present and alarmed in 100% of visits whereas males were present in 37% of visits. The individual variations in female investment in defence, after controlling for the principal determinants of nest defence, were significantly correlated with breeding success of each pair.

Conclusion The relationship between female investment in nest defence and reproductive success can be explained through differences in female quality. Good quality females, as measured by the level of investment in nest defence, probably also invested more in other breeding aspects not controlled in this study obtaining higher reproductive success.     

Mating system and mate choice of Tengmalm's Owls Aegolius funereus

The mating system and mate choice of Tengmalm's Owls were studied in relation to population fluctuations of the staple food (voles) for seven years in western Finland. Three age classes of owls were differentiated: first-year, second-year and older owls. Despite a surplus of males, 11 % of them showed simultaneous bigyny in peak vole years. The mean distance between primary and secondary nests was 1158 m, and usually there was a free nest-box between the two nests. Because primary nests produced more fledglings than secondary ones and secondary nests produced fewer fledglings than simultaneous or even eight-days-later monogamous nests, this bigyny could be best explained by the 'deception' hypothesis.
In successive biandry, females deserted their first brood when young were still in the nest, thereby transferring the burden of raising the young to their males. The tendency of assortative mating by age (more yearling x yearling and + 2-year-old ×+ 2-year-old pairs than expected by chance) showed that females chose older males when these were available.     

Tawny Owls Strix aluco with reliable food supply produce male-biased broods

Tawny Owls Strix aluco have been reported to skew the sex ratio of their offspring towards males when facing food shortage during the nestling period (and vice versa), because female fitness is more compromised by food shortage during development than male fitness. To test the generality of these results we used a DNA marker technique to determine the sex ratio in broods of Tawny Owls in Danish deciduous woodland during two years of ample food supply (rodent population outbreak) and two years of poor food supply. Of 268 nestlings, 59% were males (95% CI: 53–65%). This proportion was higher than previously reported for the species (49% in Northumberland, UK, and 52% in Hungary), but consistent with Fisherian sex allocation, which predicts a male bias of c . 57% based on inferred differences in energy requirements of male and female chicks. Contrary to previous results, brood sex ratios were not correlated with the resource abundance during the breeding seasons, despite considerable variation in breeding frequency, brood size or hatching date across years. Brood sex ratios were unaffected by brood reduction prior to DNA sampling, and nestling mortality rates after DNA sampling were not related to gender. The inconsistency between the sex ratio allocation patterns in our study and previous investigations suggests that adaptive sex allocation strategies differ across populations. These differences may relate to reproductive constraints in our population, where reproductive decisions seem primarily to concern whether to lay eggs at all, rather than adjust the sex ratio to differences in starvation risk of nestlings.     

Reproductive biology and ecology of white‐winged trumpeters (Psophia leucoptera) and recommendations for the breeding of captive trumpeters

The reproductive biology and ecology of a wild population of white‐winged trumpeters (Psophia leucoptera) were studied in southeastern Peru from 1983 to 1987. Because little information is available about any of the trumpeter species and because trumpeters have proven difficult to breed in captivity, information relevant to breeding and management of captive trumpeters is reported in this paper. White‐winged trumpeters lived in territorial social groups that ranged in size from four to 13 individuals. A typical territorial group contained three adult males, two adult females, and several sexually immature offspring, but smaller temporary groups sometimes formed for the duration of the breeding season. Only the dominant female contributed eggs to the clutch, and all adult males in the group competed to obtain copulations with her. Eggs were laid in elevated nesting cavities and no nest was constructed. The average clutch size was three eggs and incubation was not begun until the final egg was laid. The dominant male and female shared most of the incubation duties, but subordinate males covered approximately 15% of the incubation shifts. Eggs hatched approximately 27 days after incubation was begun and chicks left the nesting cavity the day after they hatched. Chicks were completely dependent on older birds to feed them for their first 3 weeks and then gradually began to feed themselves more and more food. The subordinate adult males fed chicks the most food, the dominant male and female and older offspring fed chicks an intermediate amount, and the subordinate adult female fed chicks the least. Young chicks behaved aggressively toward each other but were separated by adults before they injured each other. If at least one chick from the clutch survived, trumpeters did not breed again until the beginning of the next breeding season the following year. Zoo Biol 19:65–84, 2000. © 2000 Wiley‐Liss, Inc.     

Maternal yolk androgens stimulate territorial behaviour in black-headed gull chicks

Avian eggs contain substantial amounts of maternal androgens. The concentrations of these yolk androgens are affected by the maternal environment, such as the level of social competition, parasite exposure or food conditions. Since yolk androgens have been shown to affect a wide array of offspring traits, they may adjust the chicks to the expected post-hatching environment, but experimental evidence is still scarce. We investigate in colonial breeding black-headed gulls whether high concentrations of yolk androgens, such as those found in environments with high numbers of social interactions, facilitate aggressiveness and territorial behaviour of the chicks. Black-headed gulls are highly suitable for this, as the semi-precocial chicks defend the natal territory and food against intruders. We manipulated yolk androgen concentrations and investigated their role in both within-nest and between-nest aggression. We found that chicks hatching from androgen-treated eggs defended the natal territory more often than their nest mates from control eggs, without increasing sibling aggression. This suggests that variation in yolk androgen concentrations in relation to the social environment of the mother may indeed allow adjustment of the offspring''s behaviour to the expected frequency of territorial interactions with conspecifics post-hatching.     

The common cuckoo Cuculus canorus and its cavity nesting host, the redstart Phoenicurus phoenicurus: a peculiar cuckoo-host system?

We examined redstart Phoenicurus phoenicurus populations over a period of fifteen years to study interactions between the cuckoo Cuculus canorus and its cavity-nesting host. Over 380 redstart nests were checked and more than 100 cuckoo eggs were found during the study period. The average parasitism rate was 20%. The cuckoos' breeding success was extremely low, only 18 chicks surviving to the fledgling stage. When redstarts were parasitized experimentally with artificial cuckoo eggs, they rejected eight percent of mimetic eggs and 44% of non-mimetic eggs. We were not able to record any rejection of the real cuckoo eggs. However, about 30% of the real cuckoo eggs were found outside the redstart's nest cup. This could be the result of laying failures by the cuckoo, rather than of a strong rejection behaviour by the redstart. We suggest that redstarts' cavity nesting itself was a factor that reduced the cost of the parasitism dramatically. Firstly, it makes it difficult for the female cuckoo to lay her egg correctly in the nest and secondly, it is more difficult for the cuckoo chick to evict the host's eggs or nestlings effectively from the nest. Only 54% of the cuckoo chicks were able to evict all the host eggs or chicks from the nest. When reared in mixed broods, cuckoo chicks survived only in every second case to fledgling age, while at least one redstart chick from every brood managed to leave the nest.