Parental differences in brood provisioning by Hen Harriers Circus cyaneus

Capsule Females varied their provisioning patterns according to brood age and brood size, whereas males did not.

Aims To quantify how parents balance the needs of their offspring for food and protection.

Methods We studied 13 nests from hides and spent on average 101 hours per nest monitoring prey types, provisioning rate and the time spent at the nest by both sexes in relation to brood size and brood age.

Results Males always provided more food than females. Males brought similar amounts of prey items irrespective of brood size and nestling age, whereas females brought more prey and bigger items to larger and older broods. Females spent less time brooding larger broods, particularly early on.

Conclusions Hen Harrier parents share the provisioning burden, with each parent delivering prey as a function of brood care requirements, hunting capability and the behaviour of the other parent.     

Male parental care promotes early fledging in an open-nester, the Willow Warbler Phylloscopus trochilus

The importance of male parental care to female reproductive success was investigated in the monogamous Willow Warbler Phylloscopus trochilus by removing the male parent at two different stages of the breeding cycle. Females that were widowed at the start of egg-laying continued breeding and managed to raise their brood on their own with no apparent reductions in numbers fledged or fledgling body-mass. The widowed females compensated for the loss of male assistance by increasing their own food provisioning rate as compared with control females. However, widows spent less time brooding the small young, and the growth rate of nestlings was reduced. In nests where the male parent was removed 7 days after the eggs hatched, the subsequent growth rate of nestlings was still affected, which suggests that male care is influential throughout the nestling period. On average, broods reared by widows fledged 2 days later than did broods of control females. An extension of the nestling period may appreciably affect reproductive success, since 68% of nests failed due to predation, mostly during the nestling period. We suggest that the main role of male parental care in the Willow Warbler is to assure a high growth rate of nestlings, which leads to early fledging and hence a reduced risk of nest predation.     

Reversed sexual size dimorphism and parental care in the Red-footed Booby Sula sula

The 'division-of-labour' hypothesis predicts that males and females perform different roles in parental care and that natural selection acts differently on each sex so as to produce different body size optima suited to their particular roles. Reversed sexual size dimorphism in avian species (females larger than males) may therefore be an adaptive consequence of different roles of males and females in parental care. We investigated patterns of nest attendance, brooding, foraging and provisioning rate in a tropical seabird, the Red-footed Booby Sula sula , a species showing a reversed sexual size dimorphism. During incubation, females attended the nest more often than males, and spent more time brooding the small chick than did males during daytime. Males and females did not differ in the average duration of their foraging trips. During incubation, there was a positive relationship between nest attendance and the duration of foraging trips in males, but not in females. During the small-chick stage, for the same time spent at the nest, males spent significantly more time than females at sea. On average, females fed the chick more often than did males. In males, there was a significant and positive relationship between the probability of feeding the chick and the duration of the foraging trip, whereas in females, this probability was much less dependent on the duration of the foraging trip. Overall, female Red-footed Boobies achieved slightly, but significantly, more parental commitment than did males. However, these sexual differences in parental participation were small, suggesting a minimal division of labour in the Red-footed Booby. Our results suggest that the division of labour hypothesis is unlikely to explain fully the adult size dimorphism in Red-footed Boobies.     

Parental behaviour in the Lapwing Vanellus vanellus

Parental behaviour of monogamous and polygynous Lapwings was studied during incubation and brood care. Both parents attended the nest in 86% of monogamous pairs ( n = 29 pairs). In 14% of pairs, only the male parent continued incubation until the eggs hatched, whereas the female deserted the clutch before or at the end of incubation. There was a clear division of parental roles during incubation. Females spent more time incubating (64% of time) than their mates (27%), whereas males spent more time defending the nest (3%) than females (>1%). Time spent incubating did not differ between monogamous and polygynous males. However, polygynous females spent more time incubating (primary females: 95%; secondary females: 97%) than monogamous females. Biparental care was the most common pattern of post-hatching care, although in some broods either the male or the female parent deserted before the chicks fledged. Division of sex roles was less pronounced in brood care than during incubation. Females spent more time brooding (21%) than males (3%), and females attended their chicks more closely than males. Nevertheless, males and females spent similar amounts of time defending the brood from predators and conspecifics. We suggest that the apparent division of parental roles may be explained by sexual selection, i.e. the remating opportunities for male Lapwings might be reduced if they increase their share in incubation. However, the different efficiency of care provision, for example in ability to defend offspring, may also influence the roles of the sexes in parental care.     

Brood size and body condition in the House Sparrow Passer domesticus: the influence of brooding behaviour

In many bird species, females undergo a marked decline in body condition during the first days of the nestling period. This decline may be because brooding young chicks reduces the time available for foraging. Alternatively, it might be viewed as an adaptive way to reduce flight costs when the food demand of the brood is highest. To test these hypotheses we modified the brooding commitment of House Sparrows Passer domesticus by manipulating brood size to see if changes in time spent brooding affects adult body condition. During the nestling period, females provided on average three times as much brooding as males. Reduced broods received 14% more brooding than large broods and time spent brooding declined with brood size and chick age according to an exponential decay function. Male body condition was unaffected by brood size and remained stable throughout the reproductive period. Body condition of females with enlarged broods decreased gradually during the nestling period, whereas that of females tending reduced broods dropped abruptly and significantly upon hatching. This resulted in females with reduced broods having lower body condition during the first half of the nestling period than those with enlarged broods. The sharp drop in body condition of females with reduced broods coincided with the period that brooding was most intensive. Indeed, female body condition at the end of the nestling period was negatively correlated with the proportion of time they spent brooding during the first half of the nestling period. Thus, the probable lower homeothermic capacities of reduced broods implies a higher brooding commitment for female House Sparrows that, in turn, may reduce their opportunity to forage and consequently also their body condition.     

Sex-related parental care strategies in the lesser spotted woodpecker Picoides minor: of flexible mothers and dependable fathers

We investigated sex-specific parental care behaviour of lesser spotted woodpeckers Picoides minor in the low mountain range Taunus, Germany. Observed parental care included incubation, nest sanitation as well as brooding and feeding of nestlings. Contributions of the two sexes to parental care changed in progress of the breeding period. During incubation and the first half of the nestling period, parental care was divided equally between partners. However, in the late nestling stage, we found males to feed their nestlings irrespective of brood size while females considerably decreased feeding rate with the number of nestlings. This behaviour culminated in desertion of small broods by females shortly before fledging. The fact that even deserted nests were successful indicates that males were able to compensate for the females' absence. Interestingly, the mating of one female with two males with separate nests could be found in the population, which confirms earlier findings of polyandry in the lesser spotted woodpecker. We conclude that biparental care is not essential in the later stage and one partner can reduce effort and thus costs of parental care, at least in small broods where the mate is able to compensate for that behaviour. Reduced care and desertion appears only in females, which might be caused by a combination of two traits: First, females might suffer higher costs of investment in terms of mortality and secondly, male-biased sex ratio in the population generally leads to higher mating probabilities for females in the following breeding season. The occurrence of polyandry seems to be a result of these conditions.     

Parental care and social mating system in the Lesser Spotted Woodpecker Dendrocopos minor

The sexes’ share in parental care and the social mating system in a marked population of the single‐brooded Lesser Spotted Woodpecker Dendrocopos minor were studied in 17 woodpecker territories in southern Sweden during 10 years. The birds showed a very strong mate fidelity between years; the divorce rate was 3.4%. In monogamous pairs, the male provided more parental care than the female. The male did most of the nest building and all incubation and brooding at night. Daytime incubation and brooding were shared equally by the sexes, and biparental care at these early breeding stages is probably necessary for successful breeding. In 42% of the nests, however, though still alive the female deserted the brood the last week of the nestling period, whereas the male invariably fed until fledging and fully compensated for the absent female. Post‐fledging care could not be quantified, but was likely shared by both parents. Females who ceased feeding at the late nestling stage resumed care after fledging. We argue that the high premium on breeding with the same mate for consecutive years and the overall lower survival of females have shaped this male‐biased organisation of parental care. In the six years with best data, most social matings were monogamous, but 8.5% of the females (N=59) exhibited simultaneous multi‐nest (classical) polyandry and 2.9% of the males (N=68) exhibited multi‐nest polygyny. Polyandrous females raised 39% more young than monogamous pairs. These females invested equal amounts of parental care at all their nests, but their investment at each nest was lower than that of monogamous females. The polyandrously mated males fully compensated for this lower female investment. Polygynous males invested mainly in their primary nest and appeared to be less successful than polyandrous females. Polyandry and polygyny occurred only when the population sex ratio was biased, and due to strong intra‐sexual competition this is likely a prerequisite for polygamous mating in Lesser Spotted Woodpeckers.     

Behavioral Plasticity in Response to Perceived Predation Risk in Breeding House Wrens

Predation is a significant cause of nest failure in passerine birds, and, thus, natural selection is expected to favor behavioral plasticity to allow birds to respond to perceived changes in predation risk. However, behavioral plasticity in response to perceived predation risk, and its potential fitness-related costs, are understudied. In a wild population of breeding house wrens (Troglodytes aedon), we tested the hypotheses that (1) birds show behavioral plasticity in response to perceived nest-predation risk to reduce self-risk or risk to offspring, but (2) this plasticity incurs fitness-related costs. We experimentally increased the perceived risk of nest predation by enlarging the diameter of the nestbox entrance from the standard 3.2 to 5.0 cm once incubation began. Unexpectedly, large-hole females spent significantly less time being vigilant than small-hole (control) females during late incubation. Both males and females also exhibited plasticity in their provisioning behavior. Large-hole males increased and large-hole females decreased provisioning visits with increasing brood size, whereas small-hole males and females behaved similarly and were unaffected by brood size. Females did not show plasticity in their incubation or brooding behavior. Notwithstanding this behavioral plasticity in response to increased perceived predation risk, treatment had no effect on hatching success or early hatchling survival, nor did it affect nestling body condition or fledging success. We conclude, therefore, that house wrens show behavioral plasticity in response to perceived nest-predation risk, but that any short-term fitness-related costs associated with this flexibility appear negligible.     

Shared care provides time-budgeting advantages for female eiders

Shared parental care resulting from brood amalgamation is often considered beneficial to parents. However, the benefit of grouping (a decline in individual vigilance despite an increase in collective vigilance, with a concomitant increase in individual feeding time) has not been evoked as a factor promoting shared care. Eider females, Somateria mollissima, are subject to substantial energetic costs during breeding, and sometimes share brood-rearing duties. We compared the activity budgets and feeding behaviour of lone tenders and multifemale tenders. We also measured the collective vigilance of multifemale tenders and examined whether the coordination of feeding activity among females changes with time. As expected, the proportion of time spent feeding increased, and the proportion of time spent vigilant decreased, as the number of females attending the brood increased. None the less, the collective vigilance of multifemale tenders was at least 20% higher than the vigilance of lone tenders. Furthermore, multifemale tending allowed females to feed more optimally, as dive duration increased with the number of females tending the brood. The level of parental investment by individual females after hatching of the eggs is therefore related to the number of tending females and, based on previous work, a female's body condition when the eggs hatch is also logically connected with the number of tending females. Whether females sharing brood care on a permanent basis coordinate their feeding activity, so as to increase the potential protection of ducklings against predation, remains equivocal. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Sex‐Specific Parental Care in Response to Predation Risk in the European Roller,Coracias garrulus

Sublethal effects of predation constitute an important part of predation effects, which may modulate prey population and community dynamics. In birds, the risk of nest predation may cause a reduction in parental activity in the care of offspring to reduce the chance of being detected by predators. In addition, parents may modify their parental food allocation preferences within the brood in response to predation risk. Our aim in this study was to evaluate the effects of risk of nest predation on parental care and within‐nest food allocation in the European Roller (Coracias garrulus), an asynchronously hatching bird. We manipulated brood predation risk by placing a snake model near the nests that simulates the most common nest predator in the Mediterranean region. Our results show that males but not females increased their provisioning rate when they were exposed to the model and that despite this, nestlings’ body mass decreased in response to this temporary increase in predation risk. We did not find evidence that parents changed their food allocation strategy towards senior or junior nestlings in their nests in response to predation risk. These results show that the European roller modifies parental care in response to their perception of predation risk in the nest and a sex‐specific sensitivity to the threat, which suggests a different perception of offspring reproductive value by parents. Finally, our results show that changes in parental behaviour in response to nest predation risk might have consequences for nestling fitness prospects.     

Who Cares? Males Provide Most Parental Care in a Monogamous Nesting Cuckoo

Cuckoos hold a prominent position in the study of parental care, because they show the greatest variation of any bird family in the way they care for their offspring. Despite this, few data are available on cuckoos with biparental care even though it is the most common form of parental care in cuckoos and birds in general. Here I describe the breeding behaviour of the pheasant coucal, Centropus phasianinus , an old-world nesting cuckoo. I show that males almost exclusively build the nest and incubate and brood the young alone both at night and during the day. The incubation pattern of short, c . 40 min bouts on the nest interspersed with 20 min frequent recesses is well suited to incubation by a single parent and is widespread amongst birds. Males also defend the nestlings and deliver 80% of the feeds to the young consisting of insects (65%), frogs and reptiles. Females did not compensate for their fewer feeding visits by delivering more vertebrates than males. Although coucal males get little help feeding, the hourly feeding rate of 3.8 feeds per brood (1.4 feeds/nestling) exceed that of the few nesting cuckoos studied to date and matches that of other tropical non-passerines. Predominantly male care is found in less than 5% of birds, mainly species with reversed sexual size dimorphism or reversed sex-roles. Its evolution, especially in monogamous species, remains puzzling. The breeding behaviour of pheasant coucals illustrates a possible transitional stage in the evolution of polyandry and interspecific brood parasitism in cuckoos.     

Reproductive success and individual variation in feeding frequency of House Sparrows (Passer domesticus)

Parental care is assumed to be closely associated with individual differences in reproductive success. We investigated how feeding frequencies varied among parents and how this affected the subsequent reproductive success in insular populations of House Sparrows Passer domesticus in northern Norway. Female parents fed their offspring more than male parents did, and the feeding rates were positively related to the feeding rates of the partner. A positive relationship between feeding rates and bill depth was present in females. In males, the feeding rates were negatively related to total badge size and positively related to visible badge size, after the effect of other variables had been taken into account. A non-linear convex relationship between feeding frequency and hatch day was present in males, which could reflect either the seasonal change in weather conditions or the seasonal variation in food availability. For both sexes, feeding frequencies increased with increasing brood size, but at the same time the average feeding rate per nestling decreased with increased brood sizes. Finally, our results indicate that the amount of parental investment, measured as feeding rates during the nestling stage, may have a positive long-term influence both on the number of fledglings that recruit as well as the probability that fledglings survive until recruitment.     

Predation by sparrowhawks Accipiter nisus on male and female pied flycatchers Ficedula hypoleuca in relation to their breeding behaviour and foraging

Bright plumage, song display, and aggressive resource defence in males may cause higher predation on males than on females during the breeding season. However, in birds, higher predation on females is sometimes observed. Parental investment may be high in females (egg-laying, incubation and feeding of offspring), which might lead to a high risk of predation. We studied predation by sparrowhawks Accipiter nisus in relation to behaviour in pied flycatchers Ficedula hypoleuca where breeding males are more conspicuous than females in plumage and behaviour. Male pied flycatchers generally occupied more exposed perches than females. Females were more mobile and foraged more than males, especially prior to and during incubation. During the incubation and nestling stages, when predation on the sexes could be directly compared, sparrowhawks took about the same number of male and female pied flycatchers. During incubation, however, females spent about 77% of the day in the nest and were 4.7 times more vulnerable than males per unit of time available (i.e. outside the nest). A comparison with the chaffinch Fringilla coelebs , where hawks took more females than males, indicates that timing of breeding, foraging behaviour and parental roles of males and females affect predation risk.     

Individual Consistency in Parental Effort Across Multiple Stages of Care in the House Sparrow, Passer domesticus

In many avian species in which biparental care is provided to offspring, substantial variation exists within members of each sex in the level of effort contributed to various forms of parental care. Questions remain as to whether individuals that contribute more toward one parental activity also contribute more toward other activities in which they participate. We examined the contributions of male and female house sparrows (Passer domesticus) to three forms of parental care: incubation, nestling provisioning, and nest defense, and compared the investments made by individuals at each stage of care relative to other same‐sexed parents. In both males and females, nestling feeding rates were positively associated with time spent incubating, but no relationships were found between measures of nestling feeding and nest defense. The predictability of an individual's feeding behavior based on earlier incubation efforts may make incubation a good stage for individuals to evaluate the parental abilities of their partners.     

Costs and benefits of parental care in eastern kingbirds

We manipulated brood sizes of eastern kingbirds (Tyrannus tyrannus)to measure the costs and benefits of parental care and to testwhether kingbirds showed evidence of individual optimizationof reproductive effort. We found that the number of feedingtrips (trips/h) increased and that per capita feeding rates(trips/nestling/h) declined as brood size increased. The declinein per capita feeding rates was mostly due to high feeding rateto broods of one: parents made roughly equal number of tripsto feed each nestling in broods of two to five. Nonetheless,nestling mass declined with brood size, probably because largebroods were fed more small prey. Nestling condition (mass adjustedfor structural size) differed only between broods of one andfive. After controlling for effects of brood size, feeding rateshad no supplementary influence on either nestling size or condition,but productivity and feeding rate were positively and significantlyrelated. Adult male condition did not vary with brood size,manipulated brood size, or total feeding rate, but declinedas the pair's per capita feeding rates increased. In addition,males that returned to breed were in better condition beforeleaving for migration than those that failed to return. Femalecondition tended to decline, and the probability of returningto breed dropped when broods were enlarged. However, femalecondition was independent of the probability of returning. Ourresults show that high feeding rates were costly, but that theycarried benefits (greater productivity). Some evidence for individualoptimization of reproductive effort existed: variability innestling and adult female condition were better explained bychanges in brood size than by the actual number of young inthe nest. However, most evidence supported the alternative thatincreased brood size was equally costly for all birds     

Breeding biology and provisioning of nestling snow buntings in the Canadian High Arctic

We examined basic breeding biology, as well as the effects of parental sex, brood size, nestling age, and nest habitat on foraging distances and parental food delivery rates of snow buntings (Plectrophenax nivalis) on Devon Island, NT, Canada, in the summers of 2003–2006. Clutch sizes and initiation dates were similar to those found in previous studies, although nest density was much higher. Feeding rates and foraging distances of buntings differed depending on the size and age of the brood, but were similar between nest habitats. Unlike in earlier studies, we found that male buntings made fewer feeding visits, but traveled longer distances to forage for food than females. Whether these differences between the sexes relate to reducing competition for prey, or to differing sex roles in parental care due to the poorly insulated nest cavities of this species, remains to be determined.     

Females compensate for moult‐associated male nest desertion in Hooded Warblers

Uniparental offspring desertion occurs in a wide variety of avian taxa and usually reflects sexual conflict over parental care. In many species, desertion yields immediate reproductive benefits for deserters if they can re‐mate and breed again during the same nesting season; in such cases desertion may be selectively advantageous even if it significantly reduces the fitness of the current brood. However, in many other species, parents desert late‐season offspring when opportunities to re‐nest are absent. In these cases, any reproductive benefits of desertion are delayed, and desertion is unlikely to be advantageous unless the deserted parent can compensate for the loss of its partner and minimize costs to the current brood. We tested this parental compensation hypothesis in Hooded Warblers Setophaga citrina, a species in which males regularly desert late‐season nestlings and fledglings during moult. Females from deserted nests effectively doubled their provisioning efforts, and nestlings from deserted nests received just as much food, gained mass at the same rate, and were no more likely to die from either complete nest predation or brood reduction as young from biparental nests. The female provisioning response, however, was significantly related to nestling age; females undercompensated for male desertion when the nestlings were young, but overcompensated as nestlings approached fledging age, probably because of time constraints that brooding imposed on females with young nestlings. Overall, our results indicate that female Hooded Warblers completely compensate for male moult‐associated nest desertion, and that deserting males pay no reproductive cost for desertion, at least up to the point of fledging. Along with other studies, our findings support the general conclusion that late‐season offspring desertion is likely to evolve only when parental compensation by the deserted partner can minimize costs to the current brood.     

Provisioning behaviour at the nest in single-parent versus biparental nests and male versus female parents in the common redstart (<Emphasis Type="Italic">Phoenicurus phoenicurus</Emphasis>)

We analysed video-sequences of undisturbed parental provisioning behaviour on 12 nests of common redstart (Phoenicurus phoenicurus). In 4 of the 12 nests, chicks were fed by a single parent only. We compared provisioning rate of chicks, time spent on the nest and food allocation rules between nests with uniparental and biparental care and between male and female parents in biparental nests. In nests with a single parent, the frequency of feeding visits per parent was higher than in biparental nests. As a result, the rate of food provisioning of chicks was similar in uniparental and biparental nests. The food allocation rules did not differ between uniparental and biparental nests. In biparental nests, male and female provisioning behaviour was similar though with two exceptions: males had a strong preference for feeding chicks in front positions in the nest and females spent a longer time on the nest after feeding. We conclude that single common redstart parents are able to compensate fully for the absence of the other parent through increased provisioning efforts, and that in biparental nests, males and females contribute equally to the provisioning of the young.     

Effect of altitude on male parental expenditure in Mountain Bluebirds ( Sialia currucoides ): are higher-altitude males more attentive fathers?

Male investment of time and energy in caring for offspring varies substantially both between and within bird species. Explaining this variation is of long-standing interest to ornithologists. One factor that may affect male care is breeding site altitude, through its effects on climate. The harsher, less predictable abiotic conditions at higher altitudes are hypothesized to favour increased male investment of time and energy in offspring care. We tested this hypothesis by comparing male parental behaviour in Mountain Bluebirds (Sialia currucoides) nesting at 1500 and 2500 m a.s.l. in the Bighorn Mountains of Wyoming, USA. We compared rates of prey delivery to nestlings at these two altitudes at two times: 1–2 days after hatching, when females spend much of their time brooding young, and 12–13 days later, when brooding has ended and nestling energy demands are peaking. High-altitude males fed nestlings 18 and 28% more often than low-altitude males early and later in the nestling stage, respectively, but only the difference in late-stage feeding rates were significant. Like males, females at the high site also fed nestlings significantly more often than females at the low site later in the nestling stage (45% difference in feeding rates). Consequently, the proportion of all feeding trips made by males at the high site (40%) did not differ significantly from that at the low site (44%). Parents at the high altitude may feed nestlings more often to compensate for their greater thermoregulatory costs. Parents may also be attempting to assist nestlings in storing fat and/or attaining a large size and effective homeothermy as quickly as possible to enhance nestling ability to survive bouts of severe weather which are common at high altitudes.