Incubation feeding by helpers influences female nest attendance in the green woodhoopoe,<Emphasis Type="Italic"> Phoeniculus purpureus</Emphasis>

In many bird species, the female alone incubates the eggs, but the male provides her with some food during the incubation period. In cooperatively breeding species, helpers might be expected to assist the breeding male in provisioning the female, but their contribution has been generally ignored. Here, I show that in the green woodhoopoe, Phoeniculus purpureus, the breeding male and helpers of both sexes bring food to the incubating female. The helpers did not increase the overall amount of provisioning the female received: groups of all sizes delivered a similar amount of food per hour. Helpers did, however, reduce the workload of the breeding male. Male and female helpers provisioned incubating females equally, as expected in a species where both sexes are likely to derive equal benefits from their helping behaviours. Female nest attendance was positively related to the level of group provisioning, but only in the short term. Thus, the female-nutrition hypothesis, which suggests that incubation feeding can provide an important source of energy to the breeding female, is supported in the green woodhoopoe.Communicated by I. Hartley     

Cooperative breeding in purple gallinules: the role of helpers in feeding chicks

Summary The participation of breeders and helpers in the feeding of 21 broods of chicks was studied in a population of cooperatively breeding purple gallinules (Porphyrula martinica). In the breeding group, all birds over the age of 2 months fed chicks. Female breeders fed chicks at the highest rate, followed by male breeders and adult helpers, old juvenile helpers, and young juvenile helpers. The amount of food breeders fed chicks was independent of the number of helpers in the breeding group. However, breeders made fewer feeding visits when they had helpers. Male and female breeders spent similar amounts of time feeding chicks. Helpers spent significantly less time feeding chicks than did breeders. As helpers grew older they fed chicks at a faster rate, made more feeding visits and spent more time feeding chicks. Analysis of variance was used to determine which variables explained the variation in the brood feeding rate (amount of food delivered to an entire brood during one observation). Age of chicks had a significant nonlinear effect, and size of brood and number of helpers had significant linear effects on the brood feeding rate. Chicks in groups with helpers received more food and were accompanied for longer periods of time than chicks in groups without helpers; either or both of these factors may have led to increased chick survival.     

Flock size,food dispersion,and the feeding behavior of crossbills

Summary Feeding rates of five captive red crossbills (Loxia curvirostra) were measured when they were foraging alone, and in flocks of two or four on three seed dispersion patterns. On the most strongly clumped seed dispersion, individuals had higher mean feeding rates and the smallest probability of starvation when in flocks of two than when alone or in flocks of four. Individuals in flocks of four had higher feeding rates on the weakly clumped seed dispersion than on the uniform and more clumped seed dispersions; there were no food finding benefits gained from flocking on the uniform seed dispersion and aggression increased as food became more clumped. Most recent work has assumed that flocking results in higher feeding rates only because time spent vigilant is reduced. Crossbills, however, did not visit more cones per unit time as flock size increased, as would be expected if less time was spent vigilant. Thus, any reductions in vigilance as flock size increased were countered by increases in other behaviours, such as those related to aggression. Consequently, the higher mean feeding rates of crossbills in flocks than when solitary is not attributable to reduced vigilance. The increase in mean and the decline in variance of feeding rates occurred because crossbills in flocks found good patches earlier, and possible by spending less time assessing poor patches.     

Seasonal anorexia in the male red-sided garter snake,<Emphasis Type="Italic"> Thamnophis sirtalis parietalis</Emphasis>

Many animals show seasonal shifts in behaviors that coincide with breeding, migration, or hibernation. These behavioral shifts provide ideal opportunities to study the regulation of behavior. The red-sided garter snake (Thamnophis sirtalis parietalis) spends 8 months of the year inactive in underground hibernacula, 1 month breeding, and 3 months feeding to build up enough energy stores to survive the following winter. Although they emerge from 8 months of hibernation with severely depleted energy reserves, they do not feed until weeks later, after the breeding season. We tested the hypothesis that this lack of feeding during the breeding season is due to a shift in behavior rather than the distribution of food and potential mates. Male garter snakes were given a series of choices between pursuing a breeding or feeding opportunity. The proportion of tests in which males selected feeding over breeding gradually increased throughout the study period, reaching almost 100% in the final tests. Males also were given opportunities to feed and court at the beginning and end of the study. Males initially refused food and courted females, but when retested at the end of the study they fed and did not court females. Thus aphagia during the breeding season is due at least in part to an endogenous shift in behavior.Communicated by P. Weatherhead     

Is asynchronous hatching adaptive in herring gulls (Larus argentatus)?

Hatching asynchrony commonly induces a size hierarchy among siblings and the resultant competition for food between siblings can often lead to starvation of the smallest chicks within a brood. We created herring gull (Larus argentatus) broods with varying degrees of hatching synchrony by manipulating the timing of incubation while maintaining the originally laid eggs. The degree of hatching asynchrony affected sibling size hierarchy at the time of hatching of the last-hatched ”c-chick.” In unmanipulated broods, there was no disadvantage of being a c-chick. However, when asynchrony was experimentally increased, we found reduced survival of the c-chick only in the exaggerated asynchronous experimental group. The effects were observable only during the first 10 days of chick life. We recorded no cases of the chicks dying of starvation. Furthermore, behavioral observations indicated that there was no sibling competition, and no selective feeding of larger sibs in the study colony. We propose that the observed lower survival rates of c-chicks in exaggerated asynchronous broods resulted from their lesser motor abilities, affecting their chances of escaping predators. Fledging success for the whole colony was generally high and almost half of all pairs fledged all three chicks, which is indicative of a good feeding environment. We argue that normal hatching asynchrony is a favorable solution in a good feeding environment, but that increased asynchrony reduces breeding success. We do not view asynchrony in the herring gull as an adaptation for brood reduction and propose instead that it may come about because there has been selection for incubation to start before clutch completion. Received: 14 April 1999 / Received in revised form: 20 October 1999 / Accepted: 23 January 2000     

Proximate control of siblicide in cattle egrets: a test of the food-amount hypothesis

Avian sibling aggression plays an important role in controlling food distribution among nest mates. The food-amount hypothesis predicts that the amount of food obtained by siblings acts as a proximate cue controlling their level of aggression. The importance of this cue to cattle egrets (Bubulcus ibis) in Oklahoma was evaluated by studying parental feeding rates, food distribution among nestlings, and nestling aggression in three-chick broods [A-(oldest), B-, and C-chicks]. We compared nests in which siblicide occurred to nests where all three chicks survived (nonsiblicide nests). Relative to nonsiblicide nests, siblicide nests received decreasing amounts of food as the day of siblicide approached. Similarly, aggression levels of all three chicks in siblicide nests increased significantly compared with nonsiblicide nests, with most of the aggressive behavior involving B- and C-chicks. As a result, on a per-day basis, food amount and aggression exhibited a strong, negative correlation, thus providing support for the food-amount hypothesis. Though the total amount of food delivered to siblicide nests declined on the days prior to siblicide, the amount of food received by A-chicks in these nests did not. Relative to nonsiblicide nests, A-chicks in siblicide nests received an increased share (percent of volume) of the food delivered at the expense of C-chicks, with the percent remaining the same for B-chicks. The percentage of food B-chicks received the day after siblicide increased significantly from that received the day before siblicide, with no change occurring in either the amount or percentage of food received by A-chicks; clearly, B-chicks were the beneficiary of their aggressive actions that resulted in the death of C-chicks. We provide evidence supporting the hypothesis that aggression is proximately linked to food amount, possibly by a relatively simple cue such as hunger. Received: 25 November 1994/Accepted after revision: 10 February 1996     

Benefits of large broods by higher chick survival and better territories in a precocial shorebird

When reproductive success is constant in one breeding phase, different tactics that increase variation in reproductive success among individuals may evolve in other phases. For instance, in shorebirds, which usually have a limited clutch size of four eggs, variation in reproductive tactics among individuals is expected either before egg-laying (e.g. diverse mating systems) or after hatching of the young (e.g. diverse parental care). In this paper, I studied the pied avocet (Recurvirostra avosetta), a shorebird with a modal clutch size of four eggs, to test whether post-hatch chick adoption as an alternative tactic can be linked to increased variation in annual reproductive success. When predation was high, naturally adopting pairs produced more filial fledglings than did pairs not adopting chicks and not losing chicks to adoption. The number of filial fledglings increased with the number of adopted young, possibly through diluting the chances of predation on filial young. Experimental chick addition did not lead to more fledged young due to low brood integrity as shown by the frequent loss of chicks from some experimental broods. When predation was low, larger broods occupied feeding territories with higher prey abundance than smaller broods, possibly due to their dominance over smaller ones. Pairs that lost chicks to adoption (donors) fledged as many filial young in their broods as did non-adopters/non-donors, whereas the total number of donors’ filial fledglings, including those raised in adopting broods, approached that of adopters. These findings show, for the first time, that post-hatch alternative reproductive tactics can lead to variation in annual reproductive success and to higher success for some pairs even in species where past adaptations limit variation in reproductive success in a certain phase of reproduction.     

An experimental study of paternal behavior in red-winged blackbirds

Summary The effect of brood size and female nesting status on male parental behavior was investigated in red-winged blackbirds Agelaius phoeniceus using brood size manipulation experiments. Male redwings allocated parental effort on the basis of brood size and nestling age. Males began assisting females only at nests with at least three offspring older than three days. Female nesting status had no singificant influence on male parental care. When females were unable to meet a brood's demand for food, males assisted females with nestling feeding. Females did not reduce the amount of food delivered to nestlings when males assisted. The amount of food brought to nestlings by the male was additional to the amount of food provided by the female. Male assistance increased fledgling success. When female provisioning was sufficient to meet a brood's demand for food males did not assist. The value of male parental care varied inversely with the ability of the female to meet nestling food demands. The ability of unassisted females to provide sufficient food and to raise a brood of nestlings successfully appeared to be influenced by resource abundance.     

Transition to independence and evidence of extended parental care in the gentoo penguin (<Emphasis Type="Italic">Pygoscelis papua</Emphasis>)

We used radio telemetry and observations to study the activity patterns and behavior of gentoo penguin chicks at Admiralty Bay, King George Island, South Shetland Islands in 2005 during their “fledging period”; defined as the time between a chick’s first trip to sea and its final dispersal from the breeding colony. Gentoo penguins exhibited delayed dispersal of young and extended parental provisioning, behaviors not observed in other Pygoscelis species. Chicks took their first trip to sea at a mean age of 70 days of age, before finally departing the colony at a mean age of 82 days. During this fledging period, individual chicks made an average of five trips to sea. Trip duration increased significantly as chicks aged, with trips to sea becoming similar to literature values of adult foraging trips in both timing and duration. Behavioral observations and mass dynamics confirmed that many chicks were still being fed during this fledging period, with parental feeding behaviors most often observed in the late afternoon to evening hours. We hypothesize that these behaviors provide an opportunity for chicks to gain experience at sea prior to dispersal and might allow them to develop foraging skills before they are completely independent.     

Dominance in feeding territories relates to foraging success and offspring growth in brown skuas <Emphasis Type="Italic">Catharacta antarctica lonnbergi</Emphasis>

Territoriality should lead to strict dominance, as territory holders typically control access to resources and exclude others from their use. In feeding territories, dominance should be reflected in foraging success and ultimately in reproduction differences; however, these successive links have rarely been made explicit. Therefore, we investigated a population of brown skuas Catharacta antarctica lonnbergi, in which only part of the breeding population occupied feeding territories within penguin colonies. We identified the dominance hierarchy and determined the foraging success of the participants in fights for access to penguin carcasses within the territories. Furthermore, we monitored offspring growth from parents with and without feeding territories. Our results indicated a clear dominance hierarchy with territorial birds in their own territory dominating over territorial breeders from other territories, non-territorial breeders and non-breeding birds. However, territory owners could not completely exclude others from access to food. Foraging success was positively related to dominance scores: The dominant territory owners received 63% of a carcass, whereas non-territorial pairs could get less than 10%. The link between foraging success and offspring development was less clear: Although male chicks of non-territorial parents suffered from lower growth rates and, thus, delayed fledging, there were no such differences in female chicks. Territoriality in skuas did not imply a complete occupation of food, but guaranteed optimal growth conditions for offspring. Non-territorial individuals were forced to search for alternative resources, and the restricted access to the preferred food resulted in inferior conditions for offspring development, making this foraging strategy less rewarding.     

Seasonal changes in the diet of a critically endangered seabird and the importance of trawling discards

Pelagic seabirds obtain food from oceans where the availability of their prey changes rapidly both seasonally and spatially. Here, we investigated changes in the trophic habits of the critically endangered Balearic shearwater (Puffinus mauretanicus) through the breeding season and tested for dietary differences between sexes and age classes. We analysed δ¹⁵N and δ¹³C values in blood of adults during the pre-incubation, incubation and chick-rearing periods and of their chicks. Using a two-isotope mixing model, we estimated dietary contributions based on isotope values from potential prey species which included small pelagic species available naturally and demersal fish species available only from trawling discards. Balearic shearwaters showed clear isotopic and dietary variation through the breeding season. During pre-incubation, breeding adults appeared to exploit demersal fish, whereas during the incubation and chick-rearing period, they fed mainly on pelagic anchovies (Engraulis encrasicolus) and pilchards (Sardina pilchardus). Similarly, chicks were fed mainly with anchovies, a resource with a high energetic value. This variation in the dietary habits of adult shearwaters during the breeding season was probably related to both natural and fishery-induced seasonal changes in the availability of potential prey species within their main feeding grounds. However, changes in the nutritional requirements of the shearwaters could also play an important role. Indeed, diet differed between sexes during pre-incubation: females fed less on trawling discards and more on small pelagic fish than males. This sexual segregation in diet could be the consequence of higher nutritional requirements of females during this period. Our study reveals the differential importance of both trawling discards and small pelagic fish species for a pelagic seabird depending on the breeding period and illustrates the importance of considering the entire breeding season when making inferences about the importance of specific prey in seabird dietary studies.     

Food delivery and sibling competition in experimentally even-aged broods of the cattle egret

Summary The age of nestlings was made even in 12 nests of cattle egrets (Bubulcus ibis) by exchanging young chicks or eggs in a breeding colory in Japan. The growth of chicks in such synchronously hatching broods (SHBs) grew almost as fast as first-hatched chicks in 38 control (asynchronously hatching) broods (AHBs). Four last-hatched chicks in AHBs but no chicks in SHBs died of starvation. The frequency of parental nest-visits with food, that of begging food by chicks, and food mass eaten by chicks were greater in SHBs than in AHBs during the first half of the nestling period and similar thereafter. Dominance rank within each SHB was formed through ritualistic fights among siblings. It was correlated with neither growth rates nor winning ratios in food contests, but in some SHBs the subordinate chicks were attacked more frequently by dominant siblings during food contests than in AHBs. The most subordinate chick in one SHB died as a direct result of such attacks. Sibling aggressions were more frequent in SHBs than in AHBs.     

Paternal expenditure is related to brood sex ratio in polygynous great reed warblers

In many polygynous animals, parents invest more heavily in individual sons than in daughters. However, it is unclear if these differences in investment are a consequence of sex differences in the demand of offspring related to sexual size dimorphism or a consequence of parental manipulation. Here, we report on parental food delivery frequency in relation to brood size and brood sex ratio in a wild population of polygynous great reed warblers Acrocephalus arundinaceus. We used the polymorphic microsatellite loci on the Z chromosome to sex chicks. We found that paternal feeding frequency (times/h per nest) increased not with brood size, but with the proportion of males in the brood, although the demand per nest was more closely related to brood size than to brood sex ratio. Additionally, the increase in rate of paternal feeding frequency in relation to the brood sex ratio was much higher than the increase in rate of nestling food demands. Maternal feeding frequency was independent of both brood size and brood sex ratio. These results strongly suggest that fathers preferentially invest in their sons. We propose that parents can afford sex-biased parental care in animals in which food provisioning is enough for all offspring to survive. Received: 22 January 1996/Accepted after revision: 30 June 1996     

Brood reduction in response to manipulated brood sizes in the common swift (Apus apus)

Following a brood size manipulation experiment on the common swift (Apus apus) in which different levels of parental effort were created, brood reduction occurred in all five nests manipulated to four chicks and in two of the five manipulated to three young. This provided an opportunity for looking in detail at the parental investment decisions concerning allocation strategies between parent and young during the process of brood reduction. The data recorded here were analysed on a visit-by-visit basis regarding changes in parental and chick body masses, the mass of prey delivered and the estimated mass of parental self-feeding. The results show that food delivery rates did not increase in proportion to the number of chicks in the broods. This reduction in delivery rates per chick resulted in lower chick masses and ultimately in the death of one or more chicks in the larger broods. The resulting low parental body masses for adult birds feeding larger broods suggested that these parents could not have raised all their young without risking their own survival. There was a tendency for parents from nests that experienced brood reduction to feed themselves instead of allocating most of the food gathered to the young. After brood reduction, parents regained lost body mass and their young fledged at masses only slightly lower than normal. The differential allocation decisions regarding parental self-feeding which resulted in brood reduction were largely responsible for keeping the parents in good enough condition to care for the surviving nestlings. Therefore, this study clearly demonstrates that brood reduction can operate through the differential allocation of food between parent and young in a way that can have adaptive consequences for the survival of parents and their young. Correspondence to: T.L.F. Martins     

Group-living in cliff swallows as an advantage in avoiding predators

Summary Cliff swallows (Hirundo pyrrhonota) in SW Nebraska, USA, nest in colonies and associate in groups away from their colonies. The degree to which group-living in this species affords advantages in the avoiding of predators was examined. The distance from the colony at which a snake predator was detected increased with colony size. In flocks away from the colonies, group vigilance increased, but the time that each individual spent vigilant decreased, with flock size. As a result, birds in large flocks had more time for preening and mud-gathering. Cliff swallows did not effectively mob predators and thus were unable to deter predators regardless of group size. Nesting within each colony was highly synchronous, but when the effects of ectoparasites on nesting success were removed, individuals nesting during the peak breeding period were no more successful than those nesting before or after the peak. This suggests that swamping of predators is unlikely in cliff swallow colonies. Nests at the edges of colonies were more likely to be preyed upon than nests nearer the center, suggesting that colonial nesting conferred some selfish herd benefits. Overall reproductive success did not vary with colony size. While cliff swallows receive some anti-predator benefits by living in groups, the avoidance of predators is probably not a major selective force for the evolution of coloniality in this species.     

Provisioning rules and chick competition in asynchronously hatching common terns (Sterna hirundo)

Interactions between nestling birds and their parents are models for examining parent–offspring communication and sibling competition. Most studies have focused on species where young are restricted to a nest. However, offspring of many species are mobile and fed by parents for an extended period post-hatch. These chicks mobility may provide an opportunity to examine the role of signalling and physical competition on parental feeding decisions. We examined parental provisioning rules in relation to offspring behaviour and hatching order (i.e., competitive ability) in a species with mobile young, the common tern. We determined that about 95% of feedings were directed to the first chick to reach the parent when it landed with food. We developed a probabilistic model to predict the likelihood of a chick reaching the parent first, and thus receiving food. Our model showed that begging intensity, feeding history, and the interaction between begging intensity and relative proximity to the parent best predicted which chick would arrive first. Increased begging was associated with arriving first significantly more when a chick was relatively further from the parent than when it was closer than its siblings. Independently of these factors, larger, earlier-hatched chicks were more likely to be fed than smaller, later-hatched chicks. Additional analyses showed that parents landed closer to more intensively begging chicks, however, increased begging did not explain the advantage of earlier-hatched chicks because begging intensity did not vary with hatching order. Instead, earlier-hatched chicks were more likely to outrun later-hatched siblings and reach the parent first.     

Seasonal variation in the effects of food availability on gametogenesis in the purple urchin (<Emphasis Type="Italic">Strongylocentrotus purpuratus</Emphasis>)

The amount of food and when it is available affect both the timing of reproduction and the number of gametes produced by purple sea urchins, Strongylocentrotus purpuratus. To investigate this further, the effects of food availability on feeding rates, gonad growth, and gamete development were examined in S. purpuratus collected from the Point Loma kelp forest near San Diego, California, USA (32.69° N, 117.26° W) in September (Fall) 2007, and February (Spring) and July (Summer) 2008, using laboratory mesocosms. Each seasonal laboratory feeding experiment lasted 3 months, and different levels of food availability were established with different feeding frequencies (from 1 to 7 days week⁻¹). Gonad tissues of male and female urchins were staged at the end of each experiment using histological analyses. Reduced food availability resulted in increased daily consumption rates, especially in the Fall when gamete development began. Food limitation at this time resulted in failure to produce viable gametes, suggesting there is a critical period early in gonad development when food limitation affects reproductive competency. Food limitation later in gonad development did not stop viable gamete production, although it did reduce gamete output.     

Age-dependent vigilance in winter aggregations of cooperatively breeding white-winged choughs (Corcorax melanorhamphos)

Summary Winter aggregation of cooperatively breeding white-winged choughs (Corcorax melanorhamphos) was examined to elucidate the advantages for different group members. Adults have higher vigilance burdens than juveniles when foraging in their regular group. Aggregation of groups in winter enables the older members to reduce vigilance and increase the time spent foraging. Vigilance was lowest and foraging time highest in aggregations of 20 individuals but the size of cooperatively breeding groups did not exceed fifteen. Despite increased foraging time, choughs did not consume more food items when aggregating.     

Preferential allocation of food by magpies Pica pica to great spotted cuckoo Clamator glandarius chicks

Adult magpies Pica pica provide parasitic great spotted cuckoo Clamator glandarius nestlings with a diet very similar to that fed to their own chicks. In both naturally and experimentally parasitized nests, great spotted cuckoo chicks were fed at a higher rate than magpie chicks in the same nest. This preferential allocation of food by magpie parents to great spotted cuckoo chicks is consistent with the supernormal stimulus hypothesis, because this result implies that cuckoo chicks provide stronger stimuli for parental care than host chicks. Great spotted cuckoo chicks receive most of the food brought to the nest by the foster parents, because they exploit a series of stimuli which jointly (or sometimes individually) operate as a supernormal stimulus. This hypothesis predicts that if any stimulus is masked, the efficiency of the cuckoo in eliciting parental care will decrease. Here, we analyze experimentally the effects of two of these stimuli, preferential feeding of large nestlings and of nestlings with conspicuous palatal papillae. Firstly, when we experimentally introduced one medium-sized (7–9 days) cuckoo chick into an unparasitized magpie nest where the largest magpie chick was 12–15 days old, the cuckoo did not receive significantly more food than the average or the largest magpie chick. Secondly, when unparasitized nests were experimentally parasitized with a cuckoo chick that had its gape painted to mimic that of magpie chicks, the parasitic cuckoo received less food than the average magpie chick.     

Parental effort in the California gull: tests of parent-offspring conflict theory

Summary Parental effort was studied among knownaged California gulls (Larus californicus) with a brood size of two. Results supported three of Triver's predictions from parent-offspring conflict theory. First, the amount of time parents withheld food from offspring increased with increasing offspring age. Second, older parents were less likely than younger parents to withhold food from offspring throughout the period of parental care. Third, older parents had a longer period of parental care compared to younger parents. Adult survival to future breeding seasons was inversely related to duration of parental care. Older parents, in association with their greater levels of parental effort, had a lower survival rate compared to that of younger parents.