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.     

Aromatic plants in blue tit Cyanistes caeruleus nests: no negative effect on blood-sucking Protocalliphora blow fly larvae

Nesting birds use several behavioural or physiological defence mechanisms against parasites. On Corsica, female blue tits Cyanistes caeruleus incorporate fresh fragments of a limited number of aromatic plants in the nest cup, from the end of nest construction until fledging. Some of these plants negatively affect bacterial growth and host location by blood-sucking mosquitoes in laboratory conditions. In natural populations, Corsican blue tit chicks are exposed to the highest levels of blood-sucking ectoparasitic blow flies Protocalliphora spp. reported in Europe. These ectoparasites can have severe negative effects on chick development and survival probabilities, especially when food constraints are elevated. Here we investigated in several natural Corsican blue tit populations the hypothesis that aromatic plants brought to the nest have anti-blow fly effects during the chick-rearing stage. We predicted that: 1) the amount of aromatic plants should be negatively related to blow fly infestation intensity across nests, 2) experimental addition of aromatic plants in nests should reduce blow fly infestation intensity, and 3) nestlings should be in better physical condition in nests where aromatic plants were experimentally added. No significant relation was found between amount of aromatic plants in nests and blow fly infestation intensity. Experimental addition of aromatic plants did not reduce blow fly infestation intensity and did not affect the chick phenotypic parameters we measured. We conclude that aromatic plants in blue tit nests are not used as a defence against ectoparasitic Protocalliphora blow flies in our study population.     

Parental Investment Theory and Nest Defence by Imperial Shags: Effects of Offspring Number,Offspring Age,Laying Date and Parent Sex

Nest defence is a common form of parental care employed by birds to improve the survival of their offspring. Theory predicts that parents should adjust their nest defence according to the value of the brood at stake, defending more intensively broods with high survival and reproductive prospects. We evaluated the influence of offspring number, offspring age, laying date and parent sex on nest‐defence intensity (NDI) of the Imperial Shag Phalacrocorax atriceps, a sexually dimorphic seabird with seasonal decline in offspring survival and very limited renesting potential. We also evaluated whether NDI was correlated within pairs and whether NDI of both members of the pair was correlated with incubation and breeding success. To elicit defensive behaviour, we simulated predation attempts using a Kelp Gull Larus dominicanus model. As predicted by theory, NDI was positively correlated with the number of offspring in the nest and offspring age. NDI during chick rearing was higher than that at early and late incubation, while no differences were found between incubation stages. Contrary to our prediction, we did not find differences in NDI according to laying date. NDI for males was higher than females, while NDI was also positively correlated within pairs. NDI was not statistically related to incubation or breeding success. These results suggest that other factors, such as laying date or parental quality and age, play a much larger role in determining the outcome and productivity of a nesting attempt. Our results provide partial support for parental investment theory; while parental defence increased with brood value according to offspring number and age, parental defence was not related to laying date, a factor strongly affecting offspring survival and recruitment probabilities in this species.     

Cooperative nest defence in red-winged blackbirds: reciprocal altruism,kinship or by-product mutualism?

Male red-winged blackbirds (Agelaius phoeniceus) often cooperate with their neighbours in defending nests against predators. Some studies have suggested that this is an example of by-product mutualism, whereas others have suggested the possibility of reciprocal altruism. No study has addressed the possibility of kin-selected cooperation in nest defence in this species. Reciprocal altruism, kin selection and by-product mutualism are not mutually exclusive alternatives, but few studies of territorial neighbours have tested for multiple mechanisms simultaneously. We test for these three possibilities in a population of red-winged blackbirds. We used simulated defections to test for reciprocal altruism. We used analysis of microsatellite loci to test for kin selection between adult male neighbours. We also used microsatellite loci to test for by-product mutualism resulting from nest defence of offspring sired on neighbouring territories. We found that male red-winged blackbirds cooperate in nest defence primarily as a form of reciprocal altruism. Experimental males reduced their level of nest defence relative to controls following simulated defection by a neighbour. In contrast to some earlier studies, we found no evidence for by-product mutualism: males did not defend nests where they had sired extra-pair offspring. We also found no evidence for kin selection: males were no more cooperative with more closely related neighbours. Considered alongside the results from other studies, our study suggests that mechanisms stabilizing cooperation in red-winged blackbirds may vary among populations.     

Feathers,suspicions, and infidelities: an experimental study on parental care and certainty of paternity in the blue tit

Theoretical models on parental care predict that males should decrease their parental effort when paternity is in doubt. Males may use some cues to assess their certainty of paternity, and try to avoid rearing offspring sired by extra‐pair males. We have previously reported in a socially monogamous passerine, the blue tit (Cyanistes caeruleus), that males decorate their nests with feathers, and that when this ornament is manipulated, males appear to have suspicions about the presence of an intruder male. Here, we decrease the male's certainty of paternity through experimental feather supplementation to analyse whether the outcome of our experiment supports the assumptions of the parental care theory. Male C. caeruleus responded to the feather supplementation experiment by reducing their parental investment (feeding frequency and nest defence) in comparison with control males. The occurrence of extra‐pair offspring in experimental nests was double than that in controls. This suggests that the manipulation was successful not only in altering males' perceived paternity, but also, indirectly, the actual paternity. Furthermore, males that gained extra‐pair young also had a higher than average probability to lose paternity in their nest, which may imply that male C. caeruleus faced a trade‐off between obtaining extra‐pair fertilizations and maintaining paternity in their own nest. Overall, this study supports the idea that males are prone to decrease their parental effort when they perceive that the risk of losing paternity is high. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 552–561.     

Does Social Mating System Influence Nest Defence Behaviour in Great Reed Warbler (Acrocephalus arundinaceus) Males?

In birds with biparental care, males and females often conflict over how much care to provide to their offspring and it may be substantially influenced by increased level of polygamy. In accordance with sexual conflict theory, males of socially polygynous bird species provide much less care to their nestlings than do males of most socially monogamous species. Most of previous studies, however, have used feeding behaviour as an index for variations in male parental care only. However, this may be skewed if polygynous males compensate for lower feeding assistance through the provision of other parental care such as protection of nests from predators. In this paper, we examine nest defence behaviour in the facultatively polygynous great reed warbler with respect to sex and type of social mating system. We recorded latency to the first arrival, distance from the predator and defensive reaction of each parent towards a human intruder. Socially polygynous males with two simultaneously active nests defended primary females’ nests less vigorously than socially monogamous males, whereas no differences were found between monogamous and primary females. Generally, however, they took a bigger role in nest defence than males in all cases. Our results support an idea that sexual conflict is driven by polygamy and that type of social mating system can influence nest defence behaviour of facultatively polygynous birds. This finding should be taken into consideration when studying nest defence parental care in polygynous mating systems.     

Individual benefits of nestling begging: experimental evidence for an immediate effect, but no evidence for a delayed effect

The evolutionary stability of honest signalling by offspring is thought to require that begging displays be costly, so the costs and benefits of begging--and whether they are experienced individually or by the whole brood--are crucial to understanding the evolution of begging behaviour. Begging is known to have immediate individual benefits (parents distribute more food to intensely begging individuals) and delayed brood benefits (parents increase provisioning rate to the brood), but the possibility of delayed individual benefits (previous begging affects the current distribution of food) has rarely, if ever, been researched. We did this using playback of great tit Parus major chick begging and a control sound from either side of the nest. Male parents fed chicks close to the speaker more when great tit chick begging, but not other stimuli, was played back. In contrast, there was no effect of playback at the previous visit on the chicks that male parents fed. We have thus demonstrated an immediate individual benefit to begging, but found no evidence of a delayed individual benefit in this species.     

Does Nest Defence Deter or Facilitate Cowbird Parasitism? A Test of the Nesting-cue Hypothesis

The nesting-cue hypothesis poses that avian brood parasites use nest-defence responses directed toward them by hosts as cues to locate nests to parasitize. Hosts that respond more intensely to brood parasites should provide more cues about nest location than those hosts giving lower intensity responses. Thus, the nesting-cue hypothesis predicts that within a species parasitized nest owners should respond more intensely than unparasitized nest owners to cowbirds perched near and away from nests. This assumes that hosts respond to cowbirds when they are encountered away from the nest and that host responses gradually increase in intensity as the cowbird nears the nest. The nesting-cue hypothesis, its assumptions and prediction were tested using six host species of the brown-headed cowbird (Molothrus ater, Icterinae). We presented a female cowbird model at three distances from host nests and compared the responses elicited. All species responded to the cowbird at all distances, which supports the first assumption. Some of the rarely elicited responses (e.g. contacts) and the proximity of the host to the model varied significantly with distance, which suggests that cowbirds could use nest defence by the host as cues to the location of an active nest. However, parasitized nest owners did not respond more intensely than unparasitized nest owners to the cowbird positioned at any of the distances from the nest, which does not support the nesting-cue hypothesis itself. Further considerations are discussed that suggest that nest defence is not likely to be used as a nest-location cue.     

Sex‐specific food provisioning in a monomorphic seabird,the common guillemot Uria aalge: nest defence,foraging efficiency or parental effort?

Sexual differences in food provisioning rates of monomorphic seabirds are well known but poorly understood. Here, we address three hypotheses that attempt to explain female-biased food provisioning in common guillemots Uria aalge : (1) males spend more time in nest defence, (2) females have greater foraging efficiency, and (3) males allocate a greater proportion of foraging effort to self-maintenance. We found that males spent no more time with chicks than females but made longer trips and travelled further from the colony. There was extensive overlap between sexes in core foraging areas, indicating that females were not excluding males from feeding opportunities close to the colony. However, as a result of their longer trips, the total foraging areas of males were much greater than those of females. There was no difference between sexes in overall dive rate per hour at sea, in behaviour during individual dives or in a number of other measures of foraging efficiency including the frequency, depth and duration of dives and the dive: pause ratio during the final dive bout of each trip, which was presumably used by both sexes to obtain prey for the chick. These data strongly suggest that sexes did not differ in their ability to locate and capture prey. Yet males made almost twice as many dives per trip as females, suggesting that males made more dives than females for their own benefit. These results support the hypothesis that female-biased food provisioning arose from a difference between sexes in the allocation of foraging effort between parents and offspring, in anticipation of a prolonged period of male-only post-fledging care of the chick, and not from differences in foraging efficiency or time spent in nest defence.     

Nest Defence Behaviour of the Eurasian Kestrel (Falco tinnunculus) Against Human Predators

Man has become the main predator of many animal species. Because the characteristics of humans are quite distinct with respect to other terrestrial predators, the cost and benefits of defence behaviour may also differ. In this paper, we study the factors affecting nest defence behaviour of the Eurasian Kestrel ( Falco tinnunculus canariensis ) against a potential human predator throughout the reproductive cycle, as well as the balance of cost and benefits of this behaviour. The study population inhabits the island of Tenerife, and the nests are located on cliffs. The intensity of the defensive behaviour was unrelated to the frequency of human visits, prey abundance (Orthoptera, Coleoptera, lizards, birds and rodents), laying date, or number of offspring in the nest. Both males and females increased their defensive behaviour as the nesting period advanced, particularly when the chicks were older than 15 days. Moreover, the intensity of the defensive behaviour, especially of males, decreased when nests were more inaccessible. Although nest defence behaviour against humans appeared to be similar to those against other predators, the benefits are not clear because the probability of nest robbing was greater for these more aggressive pairs.     

Correlation between two components of parental investment: nest defence intensity and nestling provisioning effort of willow tits

Nest defence intensity and nestling provisioning effort of female willow tits (Parus montanus) were significantly correlated at the end of nestling period: well-fed young were defended most intensely. Increased effort was rewarded, since broods with the highest female per-offspring provisioning rates were the most likely to produce local recruits. This suggests that the feeding ability is an important cue for parental investment decisions, at least in a species like the willow tit which has adopted the clutch adjustment strategy. Thus, the most valuable broods would not necessarily be the largest ones, but the ones in which the original number of young could be fed most adequately. However, no associations were found between the level of parental effort and offspring weight, size or condition, nor did the broods producing recruits differ from other broods in timing of breeding or number and size of offspring. The female behaviour may suggest that they invest the most time, energy and risk in the young whose chances of joining the winter flock are the best. The first well-fed young also gain an advantage of prior residency in joining the flock. The first to join normally obtain higher social status, and hence better winter survival, than latecomers. The corresponding patterns in male parental investment behaviour were weak or absent, which suggested that the male effort was affected by the female behaviour. Males seemed to invest in nestling provisioning in such a way as to supplement the female effort. During nest defence action males also seemed to invest in protection of females against predation.     

REPRODUCTIVE COST,AGE-SPECIFIC SURVIVAL AND A COMPARISON OF THE REPRODUCTIVE STRATEGY IN TWO EUROPEAN TITS (GENUS PARUS)

Theoretical analyses of optimal reproductive rates usually assume a trade-off between offspring production and parental survival. This study verified a survival cost for willow tit males; nonbreeding males survived better than males attending a brood. Theory also predicts a smaller clutch size in birds that are less successful in transforming reproductive investments into mature offspring. As predicted, we found that crested tits, suffering a higher nest predation rate, laid smaller clutches than willow tits. The generally lower survival rate of willow tit adults may largely be attributed to their higher reproductive commitment (larger willow tit clutch size), because no significant interspecific survival difference remained between nonbreeding males. Finally, in willow tits we found a positive correlation between average clutch size and juvenile survival rate (density-dependent) the ensuing year, suggesting that willow tits may adjust clutch size in response to changing survival prospects for their young by using the breeding density as a cue.     

Egg speckling patterns do not advertise offspring quality or influence male provisioning in great tits

Many passerine birds lay white eggs with reddish brown speckles produced by protoporphyrin pigment. However, the function of these spots is contested. Recently, the sexually selected eggshell coloration (SSEC) hypothesis proposed that eggshell color is a sexually selected signal through which a female advertises her quality (and hence the potential quality of her future young) to her male partner, thereby encouraging him to contribute more to breeding attempts. We performed a test of the SSEC hypothesis in a common passerine, the great tit Parus major. We used a double cross-fostering design to determine whether males change their provisioning behavior based on eggshell patterns they observe at the nest. We also tested the assumption that egg patterning reflects female and/or offspring quality. Because birds differ from humans in their color and pattern perception, we used digital photography and models of bird vision to quantify egg patterns objectively. Neither male provisioning nor chick growth was related to the pattern of eggs males observed during incubation. Although heavy females laid paler, less speckled eggs, these eggs did not produce chicks that grew faster. Therefore, we conclude that the SSEC hypothesis is an unlikely explanation for the evolution of egg speckling in great tits.     

Sex differences in nest defence by the red-backed shrike<Emphasis Type="Italic"> Lanius collurio</Emphasis>: effects of offspring age,brood size,and stage of breeding season

The red-backed shrike Lanius collurio is an open-cup nesting passerine bird, which protects its nest aggressively attacking predators near the nest. The response of parents to a human observer was investigated during the nesting period 1999–2001 in Poland. The intensity of nest defence increased as the breeding cycle progressed in accordance to the offspring value hypothesis. However, we did not find an increase in the level of aggression according to the number of offspring and the time of season. Contrary to previous predictions, we did not find gender differences in nest defence. More aggressive parents, both females and males, have significantly better breeding success than quieter individuals.     

Breeding behaviour of the Bearded Vulture Gypaetus barbatus: minimal sexual differences in parental activities

Monogamous biparental care is expected to occur when opportunities for extra-pair copulations are rare, and both parents are required to raise the chick. Bearded Vultures Gypaetus barbatus fulfill these conditions. Contributions by male and female Bearded Vulture to nest building, nest defence, incubation, nest attendance and chick feeding were studied over five years in eight pairs from the Pyrenees [Catalonia, northeast Spain). Overall, the sexes show equal investment, although the degree of parental effort developed differs depending on the specific activities. During pre-laying, males were significantly more active than females in supplying material to the nest and in territorial defence behaviour, which increased (in both sexes) as the season advanced. Incubation was shared equally both by day and by night. During chick-rearing, the nest was attended by both sexes and the presence of both parents at the nest decreased in parallel with the growth of the chick. Activities related to chick feeding were also equally divided. These results are discussed in the context of female selection of mates and the particular ecology of this species.     

Age and brood defence in male CRL:NMRI BR laboratory mice, Mus musculus domesticus

According to Lack's principle, older mice should invest more in their offspring than younger ones. I tested this principle with pairs of CRL:NMRI BR mice protecting their young against an intruder, an unfamiliar same-stock male or female. The intruder was presented within a small wire-mesh cage in an extension, a cage the same size as the pair's home cage. The dam, her mate and the young in the nest were present in all tests. Although both parents had full access to the intruder, only the male interacted with it. Young breeding pairs and young intruders were 6 months old, the old pairs and old intruders 12 months. Each breeding pair was tested with both young and old male intruders. Older males defended their pups more vigorously than younger ones. The old males showed a higher preference to stay near (to interact with) the opponent; they also showed more tail rattling and biting attempts than the young ones. This indicates that older males take greater risks in defending their offspring than younger males. Older intruders also elicited a stronger brood defence in older males and a weaker response in younger ones.     

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.     

Cooperative nest‐defence behaviour and territory quality in a resident and socially monogamous passerine

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.     

Parental sex roles of Malaysian plovers during territory acquisition,incubation and chick-rearing

Shorebirds show high variability in parental care strategies among species, populations, and environments. Research on shorebird parental sex roles can help to understand the selective pressures that shape avian breeding strategies. Although several studies have examined parental care strategies in holarctic shorebirds, very little research has been conducted in the tropics. Here we examined parental sex roles during territorial defence, incubation, and chick-rearing in Malaysian plovers Charadrius peronii in the Gulf of Thailand. The costs and gains of particular parental behaviour may vary between the sexes and can be affected differently by environmental factors and chick age. Thus we also examined how temperature, prey availability, chick or embryo age, and time of day affected parental sex roles. Males spent more time defending territories and were further away from chicks whereas females spent more time incubating eggs. Both adults contributed to chick defence during disturbances throughout the entire chick-rearing period. Total nest attendance (sum of both sexes) was affected by the modelled temperature of an unincubated egg. Prey availability, embryo age, and time of day had no effect on total nest attendance. Males adjusted incubation effort in response to temperature only at high temperatures (>36°C) whereas females adjusted nest attendance at high and low temperatures. Chick age had no effect on the proportion of time adults spent defending territories or responding to disturbance. Pairs were more likely to fledge chicks if both the male and female spent more time defending territories. For Malaysian plovers, high cooperation between the sexes during parental care may help to achieve high quality breeding territories, maintain body conditions during hot days, protect offspring from predators and attacking conspecifics, and contribute to high lifetime reproductive success.     

Parental defence in Long-eared Owls Asio otus: effects of breeding stage, parent sex and human persecution

The responses of male and female Long-eared Owls to a human approaching the nest were examined. Each nest was only tested once. Nest defence increased significantly throughout the breeding season because older chicks were defended more strongly than younger chicks and eggs. No correlations were however found between defence intensity and laying date, clutch- or brood-size. These results generally do not support either the renesting-potential hypothesis or the positive reinforcement hypothesis but are in accordance with both the age-investment hypothesis and the vulnerability hypothesis. Females defended nestlings more often and more vigorously than did males. Given division of labour between sexes and the related reversed sexual dimorphism in size, female Long-eared Owls may be more willing to engage in nest defence simply because they are always nearer eggs and chicks during all breeding stages and are larger/heavier than males; consequently they may allocate more time and energy to this activity. Finally, owls experiencing higher levels of human persecution took smaller risks when defending nests than owls breeding in an undisturbed area.