2. RUDD'S BUSH-WARBLER (Apalis ruddi)

Gargett, V. 1982. The food requirement of a Black Eagle chick. Ostrich 53:178-181.

A hand-reared Black Eagle Aquila verreauxii chick that was fed on hyraxes, Heterohyrax brucei and Procavia capensis, ate 29 626 g from hatching to flying at 93 days. For 18 days after its first flight it ate 5 910 g, (mean 328 g per day). The eaglet's food intake increased in cold weather and decreased during a hot spell.     

Shiny cowbird Molothrus bonariensis egg size and chick growth vary between two hosts that differ markedly in body size

In birds, egg size affects chick growth and survival and it is an important component of reproductive success. The shiny cowbird Molothrus bonariensis is an extreme generalist brood parasite that uses hosts with a wide range of body masses. Survival of cowbird chicks decreases with host body mass, as competition for food with nestmates is more intense in large than in small hosts. We studied variation in shiny cowbird egg size and chick growth in two hosts that differ markedly in body size: the chalk‐browed mockingbird Mimus saturninus (70–75 g), and the house wren Troglodytes aedon (12–13 g). We analyzed: 1) if females parasitizing mockingbirds lay larger eggs than those parasitizing wrens, and 2) the association between egg size and chick growth. We experimentally controlled for time of parasitism and number of host chicks and evaluated growth rate of male and female parasite chicks. Shiny cowbirds parasitizing mockingbird nests laid larger eggs than those parasitizing wren nests. Chick body mass after hatching was positively associated with egg size until chicks were five days of age, but there was no association between egg size and growth rate, or asymptotic mass. There were no sexual differences in egg size or body mass at the time of hatching, but growth rate was higher in males than in females leading to sexual dimorphism in asymptotic mass. Differences in egg size between hosts and the effect of egg size on body mass after hatching support the hypothesis that different females are specialized in the use of hosts that differ in body mass.     

Egg size,fertility, hatchability,and chick survivability in captive California condors (Gymnogyps californianus)

California condors are one of the most endangered species native to the mainland United States and are subject of intense effort regarding captive breeding and reintroduction. We analyzed 20 years of California condor egg records from the wild and from three captive propagation facilities for fertility, hatchability, and chick survivability, along with changes in egg size due to multiple clutching. Overall annual mean percent of fertile eggs was 80.2%, hatchability was 87.3%, and chick survivability to ≥60 days was 95.5%. One egg-laying site had a significantly lower fertility rate (P ≤ 0.0001) than the other sites, which was probably due to pair incompatibility rather than any physiological factors. Egg volume of the first egg was significantly greater than both the second (t=6.73, P=0.0001) and third egg (t=6.62, P ≤ 0.0001) of the season, while the second egg had a significantly greater volume (t=3.20, P=0.0084) than the third egg. Chicks from the second egg (t=3.24, P=0.029) and third egg (t=7.94, P=0.0014) of the season were significantly smaller than chicks from the first egg of the season. The decrease in egg measures and chick hatch weight due to multiple clutching did not affect hatchability or chick survivability. There were significant positive relationships (P<0.0001) between fresh egg weight and chick hatch weight and between egg volume and chick hatch weight, as well as between fresh egg weight and egg volume. In spite of the decrease in fresh egg weight, egg volume and chick hatch weights, due to egg removal to stimulate double and sometimes triple clutching, the captive propagation program has been successful in producing birds for the restoration of this species. Zoo Biol 23:489–500, 2004. © 2004 Wiley-Liss, Inc.     

Reproductive measures and chick provisioning of Cory's Shearwater Calonectris diomedea borealis in the Azores

Capsule Timing of breeding influenced wing-length at fledging, and egg size may be an indicator of fledging weight and the amount of food received by chicks.

Aims To investigate chick growth, temporal patterns of chick food provisioning and the importance of indices of parental condition or quality, egg size and hatching date, to predict nestling body mass and wing-length at fledging, and compare breeding and chick feeding characteristics between colonies in the northeast Atlantic.

Methods A survey of Cory's Shearwater nests was carried out at Vila islet. A sample of 52 chicks, ringed and weighed at hatching, was selected to study chick growth and food provisioning.

Results Hatching success (51%) was much lower than fledging success (87%). Both hatching date and egg size contributed to explain wing-length at fledging, but hatching date, which was negatively correlated with wing-length at fledging, had the most important contribution (22%). There was some indication that egg size may explain variation in fledging weight and the amount of food received by chicks. Food delivery and feeding frequency of chicks varied throughout the chick development stage and three phases were distinguished: (1) 0–29 days, the highest feeding frequency values and a linear increase in food delivery; (2) 30–69 days, an oscillation in food delivery and medium feeding frequencies; (3) 70–90+ days, a sharp decrease in both food delivery and feeding frequency.

Conclusion Variation in food availability did not seem sufficient to override the overall importance of indices of parental quality in determining reproductive measures and chick provisioning. Breeding and feeding characteristics were similar between colonies in the northeast Atlantic, with variability in chick provisioning higher further south.     

Provisioning and growth in chicks of Wilson's storm-petrels (Oceanites oceanicus) on King George Island, South Shetland Islands

We present data on chick growth and chick feeding in Wilson's storm-petrel (Oceanites oceanicus) in a colony on King George Island, South Shetland Islands. Chicks were repeatedly weighed and the weight differences over 24 h were corrected for metabolic loss in order to obtain an estimation of meal sizes. Chicks were fed on 93% of the nights (n=688 nights). The average meal size for a single feeding was 8.5 g. Chicks received on average 1.2 feedings per night. These results are compared with data for this species from other locations. There was a trend for increased meal sizes from northern to southern populations, parallel to an increase in the adult mass, indicating that Wilson's storm-petrels carry optimal meal sizes according to their body size and may take advantage of increased food abundance by increasing feeding frequencies. We describe chick growth and discuss the influence of egg size, hatching date and feeding frequency on chick growth. The egg size had a positive influence on tarsus growth and body mass of chicks. Later-hatched chicks started wing growth and finished mass growth at a younger age and reached lower peak masses, indicating that late chicks may adapt to the restricted breeding season in their Antarctic breeding grounds by a more rapid development, but will fledge with a lower degree of development and less resources. Accepted: 22 May 2000     

Feeding experience and relative size modify the begging strategies of nestlings

The offspring of birds and mammals use a combination of movementsand vocalizations, known as begging, to solicit food from theirparents. A widespread interpretation of begging is that itconstitutes an honest signal of offspring need. But we knowthat in the house sparrow (Passer domesticus) the intensityof begging calls reflects the past experience of offspringin addition to their need. Here we show that this result generalizesto other species. An experiment with hand-reared magpies (Pica pica) and great spotted cuckoos (Clamator glandarius) indicates that the begging strategies depend on the past experience ofchicks and the composition of their brood. In asynchronoustwo-magpie broods, both chicks begged at the same intensitywhen the large chick obtained food more easily than its sibling,but the large chick begged at higher intensity when it was easier for the smaller chick to obtain food. Cuckoo chicks beggedat higher intensity than magpies.     

1. A CENSUS AND STUDY OF BIRDS IN ALBANY OPEN THORNBUSHVELD

Williams, A. J. &; Cooper, J. 1983. The Crowned Cormorant: breeding biology, diet, and offspring reduction strategy. Ostrich 54:213-219.

Crowned Cormorants Phalacrocorax coronatus were studied at Dassen and Marcus Islands. The most frequent clutch was three eggs. Egg size varied within clutches with first-laid eggs being largest and heaviest and subsequent eggs progressively smaller and lighter, The mean laying interval was 2,2 days, the mean laying-to-hatching interval was 23,0 days, and the hatching interval was one day. The normal incubation period was 22.4 days. The weight of hatchings was related to the position of the originating egg in the laying sequence. Chicks were fed within 24 h of hatching. Chick development is described over the first 35 days. One chick could fly at 35 days. Hatching success was 48,2%. Hatching success was greatest in second-laid eggs, least in last-laid eggs. The mean number of chicks hatched at a nest was two. Mean diving time was 23,5 s. Most food was fish, particularly klipfish Clinidae and pipefish Syngnathus, 60–160 mm long. The number of offspring produced can be related to food availability by interaction of difference in egg size, hatching asynchrony, and the preferential feeding by adults of the strongest-begging chick. There is a trend towards producing two chicks, normally those from the first two eggs to be laid.     

Brood reduction in the Blue-eyed Shag Phalacrocorax atriceps

Brood reduction is common in a population of Blue-eyed Shags on Signy Island, South Orkney Islands. This paper describes possible adaptations which may reduce the brood. In clutches of three, the last egg was smaller, and hatched 2.4 days later than its siblings. Whilst 78–84% of first and second ('A' & 'B') chicks fledged, only 11 % of 'C' chicks did. In a sample of artificially synchronized broods chick survival was as high as in normal asynchronously hatching broods, but there were more cases of total brood loss. The age at which the C chick died was related inversely to the length of the A-C hatching interval. Relative differences in sibling weights were highest during the first 12 days, when most of the C chick deaths occurred. At this age the daily food requirements of each brood of three was one-tenth that of each brood of two just prior to fledging. It is suggested that C chicks were unable to compete effectively for a food supply which was limited by the parents, rather than by the environment. The asymptotic weight attained by A chicks was inversely related to brood size, and was greater than that of B or C chicks. Normal asynchronous broods produced at least one heavy (A) chick and one medium weight (B) chick, whilst in synchronized broods the asymptotic weight attained was similar to that of B chicks in normal broods.     

Consequences of sex-specific growth on sibling competition in black-headed gulls: a sexually-size dimorphic species with scramble competition

Biased mortality of the larger sex during the early developmental period has been reported for a number of size-dimorphic bird species. This can partly be explained by the fact that growing to larger size renders the larger sex more vulnerable to food shortage. However, since sibling rivalry is often size-dependent, chicks of the larger sex should have a competitive advantage. This raises the question as to why the larger sex does not always benefit from its size in sibling competition. We studied sibling competition in the black-headed gull (Larus ridibundus), a sexually-size dimorphic species with male-biased mortality. We manipulated the natural brood sex ratio and placed one male chick in direct competition with one female chick while concurrently controlling for differences in age, size and laying order. Male chicks outgrew their female siblings by 15% in asymptotic body mass and did not suffer from enhanced mortality. Female chicks tended to be more alert when the parents returned to the nest and were more persistent in gull-typical begging displays. Females were more likely to get the first food item, but they did not get more food, possibly due to a size-mediated dominance over the non-monopolizable regurgitated food. Thus, it is unlikely that sex differences in competitiveness significantly contribute to male-biased mortality in black-headed gulls. The previously reported male-biased mortality is more likely due to a disadvantage of a higher food demand and a higher sensitivity towards low egg quality, as has been shown in previous studies.     

Supplementary feeding increases Common Buzzard Buteo buteo productivity but only in poor‐quality habitat

Temporal heterogeneity in the effects of food supply during the breeding season on the productivity of the Common Buzzard Buteo buteo was investigated in a supplementary feeding experiment. Pairs were fed artificially (1) before egg‐laying, (2) after chicks hatched and (3) continuously throughout the season, and compared with (4) unfed controls. Pairs fed before egg‐laying had marginally larger clutches than those not fed, but lay date, egg volume and weight, brood size and hatching success were unaffected. Territorial quality had far greater effects, with pairs nesting in low‐quality habitats (bog, scrub and semi‐natural grassland) laying later and having lower hatching success, smaller broods and fewer fledglings than those in more productive agricultural landscapes. Supplementary feeding after egg hatching neutralized the negative effect of poor habitat, resulting in fed birds having significantly more fledglings. This study emphasizes the importance of food availability when provisioning chicks in suboptimal habitats and has implications for the success of diversionary feeding in reducing game losses to Buzzards.     

Habitat and diet of young grouse broods: resource partitioning between Capercaillie (Tetrao urogallus) and Black Grouse (Tetrao tetrix) in boreal forests

We compared habitat use and diets of young Capercaillie and Black Grouse broods in a boreal forest in southeast Norway. We used pointing dogs to search for broods (N = 83) in mature “natural” forest types and examined the crop content of 66 chicks 1–9 weeks old. We also measured the abundance of insects in the habitats where broods were found. Although overlapping substantially in both habitat and diets, there were notable differences: Capercaillie broods were more frequently recorded in bilberry-dominated forest types, whereas Black Grouse preferentially used pine bog forest, a more open habitat with little bilberry. Capercaillie chicks ate proportionally more insects, particularly lepidopteran larvae, and insects dominated their diet for a longer period of time (until age 28–29 days) than in Black Grouse (14–15 days). After reaching their peaks, the quantity of insects in the crops declined rapidly especially in Capercaillie, and in one of 2 years this occurred at a time when insects, including larvae, were still abundant in the habitats. Among plant foods, both species ate large amounts of Bilberry (Vaccinium myrtillus) and Bog Whortleberry (V. uliginosum). The main difference between species was a large proportion of both over-wintered and new, not yet ripe, berries of Cranberry (Oxycoccus quadripetalus) in Black Grouse, and a higher proportion of the forb Melampyrum sylvaticum in Capercaillie. The difference in diets reflected their differential use of habitats; the Vaccinium-preferred habitats of Capercaillie were richer in insects, particularly larvae, than the pine bog habitat preferred by Black Grouse. Because insects, especially larvae, comprised a larger proportion of the diet of Capercaillie chicks and chicks of this species need more food to sustain their rapid growth, Capercaillie is likely to be more sensitive to variation in insect food than Black Grouse. Also, by reducing the abundance of bilberry, the main host plant of larvae chick food, clearcutting forestry has negative effects on the brood habitat quality of both species.     

Chick growth and survival in gentoo penguins (Pygoscelis papua): effect of hatching asynchrony and variation in food supply

Summary In the gentoo penguin, Pygoscelis papua, we examined the effects of intra-clutch egg size differences and hatching asynchrony on differential chick growth and survival (including post-fledging survival), in five years for which indices of food supply were available. An initial size hierarchy within-broods at hatching was due to hatching asynchrony not intra-clutch egg size differences. In 1988 only (a poor food year), the weight advantage gained by the first-hatched (A) chick persisted to the end of brooding (30 days), with more second-hatched (B) chicks dying. There was no difference between A- and B-chick weights at fledging (60 days) or in overall chick survival between synchronous and asynchronous broods in any year. Postfledging survival (measured in one year) was not related to fledging weight or hatching order. These results provide only partial support for the hypothesis that gentoo penguins operate a brood reduction strategy to optimise chick survival in years of low food availability. We suggest that hatching asynchrony in gentoo penguins may result from selection to keep the first egg warm as soon as it is laid, due to extreme low ambient temperatures.     

Effects of intraclutch variation in egg size and hatching asynchrony on nestling development and survival in semi‐precocial Herring Gulls

Intraclutch egg size variation may non‐adaptively result from nutritional/energetic constraints acting on laying females or may reflect adaptive differential investment in offspring in relation to laying/hatching order. This variation may contribute to size hierarchies among siblings already established due to hatching asynchrony, and resultant competitive asymmetries often lead to starvation of the weakest nestling within a brood. The costs in terms of chick mortality can be high. However, the extent to which this mortality is egg size‐mediated remains unclear, especially in relation to hatching asynchrony which may operate concomitantly. I assessed effects of egg size and hatching asynchrony on nestling development and survival of Herring Gulls (Larus argentatus), where the smaller size and later hatching of c‐eggs may represent a brood‐reduction strategy. To analyze variation in egg size, I recorded the laying order and laying date of 870 eggs in 290 three‐egg clutches over a 3‐yr period (2010–2012). I measured hatchlings and monitored growth and survival of 130 chicks from enclosed nests in 2011 and 2012. The negative effect of laying date (β = ?0.18 ± SE 0.06, P = 0.002) on c‐egg size possibly reflected the fact that late breeders were either low quality or inexperienced females. The mass, size, and condition of hatchling Herring Gulls were positively related to egg size (all P < 0.0001). C‐chicks suffered from increased mortality risk during the first 12 d, identified as the brood‐reduction period in my study population. Although intraclutch variation in egg size was not directly related to patterns of chick mortality, I found that smaller relative egg size interactively increased differences in relative body condition of nestlings, primarily brought about by the degree of hatching asynchrony during this brood‐reduction period. Thus, the value of relatively small c‐eggs in Herring Gulls may lie in reinforcing brood reduction through effects on nestling body condition. A reproductive strategy Herring Gulls might have adopted to maintain a three‐egg clutch, but that also enables them to adjust the number of chicks they rear relative to the prevailing environmental conditions and to their own condition during the nestling stage.     

Testing the relationship between clutch size and brood size in the Coot (Fulica atra)

The time between egg laying and chick fledging is of crucial importance for the survival of young birds. I analyzed breeding output at consecutive phases of growth of young Coots (Fulica atra) relative to the clutch size and laying date. Considering the specific breeding biology of the Coot, I tested whether chick survival reveals clutch size-dependent variability. Clutch size did not affect hatching success; it only affected brood size, and that merely temporarily. During the first 20 days after hatching, i.e. during the time of the highest chick mortality, birds with larger clutches lost chicks at a higher rate. As a result, the number of fledged chicks was independent of the initial number of chicks, and pairs with different clutch sizes had a similar number of fledglings. The laying date had no effect. This pattern of age-related chick survival points to the greater role of the type of chick growth (semi-precocial) and behavior in their survival.     

Maternal effects as drivers of sibling competition in a parent–offspring conflict context? An experimental test

Maternal effects occur when the mother's phenotype influences her offspring's phenotype. In birds, differential allocation in egg yolk components can allow mothers to compensate for the competitive disadvantage of junior chicks. We hypothesize that the parent–older chick conflict peaks at intermediate conditions: parents benefit from the younger chick(s) survival, but its death benefits the older chick in terms of growth and survival. We thus expect maternal compensation to follow a bell‐shaped pattern in relation to environmental conditions. We studied a black‐legged kittiwake (Rissa tridactyla) population where previous results revealed increased allocation of yolk testosterone in younger as compared to older chicks in intermediate conditions, in line with our theoretical framework. We therefore predicted a maternally induced increase in aggressiveness, growth, and survival for younger chicks born in intermediate environmental conditions. Controlling for parental effects and chick sex, we manipulated food availability before egg laying to create a situation with intermediate (Unfed group) and good (Fed group) environmental conditions. Within each feeding treatment, we further created experimental broods where the natural hatching order was reversed to maximize our chances to observe an effect of feeding treatment on the younger chicks' aggressiveness. As predicted, we found that chick aggressiveness was higher in younger chicks born from the Unfed group (i.e., in intermediate environmental conditions), but only when they were put in a senior position, in reversed broods. Predictions on growth and survival were not confirmed. Mothers thus seem to favor the competitiveness of their younger chick in intermediate conditions via egg yolk components, but our study also suggests that hatching asynchrony need to be small for maternal compensation to be efficient. We emphasize the need for further studies investigating other chick behaviors (e.g., begging) and focusing on the relative role of different yolk components in shaping parent–offspring conflict over sibling competition.     

4. SOME BIRDS OBSERVED AT LAKE MENTZ

Randall, R. M. &; Randall, B. M. 1900. The hard-shell4 diet of African Black Oystercatcher chicks at St Croix Island, South Africa. Ostrich 53:157-163.

The hard-shelled diet of African Black Oystercatcher Haematopus moquini chicks was established by monitoring piles of food remains brought to the chicks. Mussels were the dominant prey species constituting about 90% numerically and about 95% by mass. Limpets were the next most abundant prey item, while false limpets, keyhole limpets and whelks were present in small numbers. Transects down the intertidal area showed that African Black Oystercatchers ignored some abundant species such as barnacles and small gastropods. They took most prey species in the same relative proportion that they were represented in transects, but there was evidence of selection for limpets and avoidance of false limpets and whelks. The modal size class of mussels in feeding piles was greater than in transects.     

Effect of food deprivation on dominance status in blue-footed booby (Sula nebouxii) broods

A pecking hierarchy is normally established in the usual two-chickbrood of the blue-footed booby (Sula nebouxii). The senior (first-hatched)chick dominates its smaller sibling and receives a greater shareof parentally provided food. Experimental broods were createdby putting together two unrelated junior chicks of die sameage in a vacated foster nest The state of the chicks was manipulatedby a period of controlled artificial feeding so that each chickunderwent a different level of food deprivation. The resultingdominance relationship depended on the relative food deprivationlevel of the chicks: the hungrier chick normally became dominant.However, die effect of hunger was occasionally overruled bysize difference: when die hungrier chick was much smaller thanits foster sibling, it was unable to gain dominance over itslarger companion. Dominance status is likely to have greatervalue for die hungrier chick, while die cost of fighting shouldbe lower for die larger chick. These results conform to dieevolutionarily stable strategy predicted for games widi asymmetricpayoff and differences in resource holding power.     

Effect of brood size and hatching sequence on prefledging mortality of Sandwich terns: why lay two eggs?

The mortality of Sandwich tern Sterna sandvicensis chicks held in enclosures was studied in colonies on Griend, in the Dutch Wadden Sea, from 1992 to 1999, and on Hirsholm, in the Danish Kattegat, in 1997. Survival of chicks until fledging was 73% for chicks hatching from first-laid eggs or single-egg clutches and 59–64% for partially hatched two-egg clutches, whereas 6% of second hatchlings survived until fledging. Less than 2% of all two-chick broods actually fledged two chicks. Because 18% of the two-egg clutches only hatched one egg, 7% of fledglings of two-egg clutches originated from a second-laid egg. In nests where both eggs hatched, the number of chicks was usually reduced soon after hatching. Within five days of hatching more than 50% of the second hatchlings died of starvation or were preyed upon. It seems that overproduction commonly occurs in Sandwich terns and that investment in a surplus egg mainly serves as an insurance mechanism. On Griend and Hirsholm, chick productivity of two-egg clutches was somewhat higher than for one-egg clutches. Undernourishment was an important cause of death, either directly by starvation or by selective predation of chicks in poor condition. This, in combination with earlier, studies suggests that Sandwich tern parents on Griend are exposed to severe food stress.     

Do shiny cowbird females adjust egg pecking behavior according to the level of competition their chicks face in host nests?

Interspecific brood parasites, like the shiny cowbird (Molothrus bonariensis), lay eggs in nests of other species. Shiny cowbird females peck and puncture eggs when they parasitize host nests. This behavior increases the survival of cowbird chicks when they have to compete for food with larger nestmates. However, cowbird chicks may benefit from smaller nestmates as they increase food provisioning by parents and the cowbird chicks secure most extra provisioning. We investigated whether egg-pecking behavior by female shiny cowbirds might be adjusted to the competition that their chicks face in host nests. We found that more host eggs are destroyed per cowbird egg laid in a larger-bodied host (chalk-browed mockingbird, Mimus saturninus, 70-75 g) than a smaller-bodied host (house wrens, Troglodytes aedon, 12-13 g). We also tested egg-pecking preferences in choice experiments with female cowbirds in captivity and found cowbirds presented with eggs in artificial nests pecked first and more frequently, and punctured more frequently the larger egg when this was a host egg, but not when this was a cowbird egg. Our results are partially consistent with the hypothesis that shiny cowbird females adaptively adjust their egg pecking behavior according to the competition that their chicks face in host nests.     

Geographical variation in egg size dimorphism in rockhopper penguins

All crested penguins present a unique reversed hatching asynchrony: the larger second-laid egg (B-egg) hatches before the smaller first-laid egg (A-egg). Although both eggs often hatch, the A-chick generally dies of starvation within days after hatching. However, within rockhopper penguins, the population at the Falkland Islands is unique in that some birds manage to raise both chicks. Although it has been suggested that the egg size dimorphism between A- and B-eggs may explain how long both eggs and chicks survive, this hypothesis has never been explicitly tested. We expect that both eggs are retained longer in the less dimorphic clutches than in the more dimorphic ones. In this paper, we have compiled egg measurements for three rockhopper penguin species (Eudyptes chrysocome, E. filholi and E. moseleyi) in order to compare the intra-clutch egg size dimorphism among these species. Furthermore, we have collected new data to compare egg size dimorphism between two populations of E. chrysocome (Falkland Islands versus Staten Island). A-egg volumes are more variable between species and populations than B-egg volumes. E. chrysocome and especially the population from the Falkland Islands produces the largest A-eggs and the least dimorphic eggs. Nevertheless, as differences in A-egg volumes between species and between the populations of Falkland Islands and Staten Island are stronger and more significant than differences in egg dimorphism, we suggest that A-egg volume, more than egg dimorphism, could be one of the factors influencing the prevalence of twins. A large A-egg and/or reduced egg dimorphism is probably necessary to enable rockhopper penguins to raise two chicks, but other reasons may also be involved which enable them to keep both eggs and chicks.