Diving behaviour differs between incubating and brooding Brünnich's guillemots, Uria lomvia

Bird-borne data loggers were used to investigate the foraging strategies of Brünnich's guillemots breeding in a colony in the North Water Polynya: the flight performance and diving activity of incubating birds were compared to those of chick-rearing individuals. No significant differences were recorded between the bird groups in the potential foraging range. Conversely, clear differences were revealed between incubating and chick-rearing birds in diving behaviour. Chick-rearing birds were generally foraging at a significantly greater depth, and spent significantly higher proportions of the time submerged, than brooding individuals. Despite these differences, the estimated average daily energy expenditure of chick-rearing Brünnich's guillemots was only about 6% higher than that during incubation.     

Stage-dependent foraging in breeding black-legged kittiwakes Rissa tridactyla: distinguishing behavioural responses to intrinsic and extrinsic factors

Different phases of the annual cycle in birds and mammals are often associated with characteristic and recurrent foraging behaviours. The extent to which stage‐dependent changes in foraging behaviour are caused by intrinsic or extrinsic factors is unclear. We controlled for the effects of extrinsic factors by synchronising groups of incubating and chick‐rearing black‐legged kittiwakes Rissa tridactyla. Synchrony amongst incubators and rearers was achieved experimentally by switching eggs between nests. Behavioural responses to the treatment varied between the sexes. Male kittiwakes with prolonged‐incubation made fewer foraging trips but of greater duration compared to those rearing chicks resulting in no change in the time spent on trips between the two groups. Females with prolonged‐incubation carried out fewer trips than those rearing chicks but trip duration did not differ between the two stages which resulted in prolonged‐incubating birds spending a lower percentage time on trips. In contrast, foraging ranges did not differ between prolonged‐incubation and chick‐rearing birds for either sex. This suggests that extrinsic factors, such as food availability and distribution determine kittiwake foraging locations and ranges, whereas intrinsic factors, reflected in parental duties, constrain nest attendance. Female prolonged‐incubators invested lower levels of parental effort, in terms of daily energy expenditure, compared to chick‐rearers whereas males did not show stage‐related differences in energy expenditure. This provides evidence that incubation could be an energetically cheaper stage although under normal conditions this difference may be masked by temporal variation in environmental factors. We conclude that while conditions differ between the incubation and chick rearing stages for kittiwakes at this colony, they are not the main factors prompting changes in stage‐related foraging patterns. Intrinsic factors such as sex differences, or behaviours required for each stage of the annual cycle, rather than extrinsic factors related to seasonal environments, are likely to be the main proximate cause of recurring changes in behaviour between breeding stages.     

Early development and differentiation of the Laysan albatross (Phoebastria immutabilis (Rothschild, 1893): Procellariiformes)

Bird incubation is subdivided into two phases: differentiation (embryonic phase) and growth (fetal phase). Most birds have a relatively short incubation period (20–30 days) with the phase transition occurring midway through the incubation period. The Laysan albatross (Phoebastris immutabilis) is a large pelagic bird with a long incubation period. The purpose of this study was to document the differentiation phase with the aim of ascertaining the impact of a lengthened incubation on embryonic development. Eighty‐two previously collected albatross embryos were examined, measured, and staged. The albatross was found to develop more slowly than smaller birds, with a rate similar to other long‐incubating birds. Legs and wings grow at similar rates but exhibit variation in growth among their anatomical components. While the albatross embryos shared some morphological stages with chickens, they were more similar to ducks and pelicans. Special features of the albatross not shared with the Gallianserae (chickens and ducks) included an alligator‐like curved tail, narial tubes, and a cloacal bulge. Further examination of other larger pelagic birds with long incubation periods are needed to determine the uniqueness of the Laysan albatross embryonic development. Although much embryonic phase growth was documented in the postnatal period, little is known about the later, fetal phase in Laysan albatross. Future studies should involve examination of later (post day 32) fetuses. J. Morphol. 277:1231–1249, 2016. © 2016 Wiley Periodicals, Inc.     

Variable energetic responses to clutch size manipulations in white-throated dippers Cinclus cinclus

We used the doubly-labelled water technique to measure daily energy expenditure (DEE) of a free-living uniparental incubator, the white-throated dipper Cinclus cinclus , in Scotland. DEE was 205±8 (s.e.m.) kJ d⁻¹ for 17 females incubating their natural clutch sizes, equivalent to 3.2±0.1×basal metabolic rate (BMR). To investigate the influence of clutch size on the energy budget, we measured the DEE of 14 females with clutches increased or reduced by a single egg. Birds with reduced clutch sizes had an energy expenditure with a mean and variance that did not differ from those of birds with unmanipulated clutches. Enlarging the clutch led to an increase in energy expenditure to over 4×BMR for some individuals but not for others, resulting in greater variance in energy expenditure for birds with enlarged clutches. Individual variation in energy expenditure could not be fully explained by environmental conditions, by patterns of behaviour or clutch size. Incubating females received a maximum of only 4 kJ d⁻¹ (2% of DEE) from provisioning by the male, and mobilised up to 6 kJ d⁻¹ (3% of DEE) from reserves. Females spent 2.9±0.2 h (n=20) away from the nest each day, so a foraging rate of 95 kJ h⁻¹ was required during incubation recesses to balance DEE. This 'required foraging rate' is double previous estimates of the maximum rates of energy acquisition for birds of this size. We suggest that the greater likelihood of a raised energy expenditure associated with larger clutches, combined with the difficulties in maintaining energy supplies, may constitute a constraint on avian clutch size.     

Foraging movements of the Shy Albatross Diomedea cauta breeding in Australia; implications for interactions with longline fisheries

Satellite telemetry was used to identify the foraging zones of Shy Albatrosses Diomedea cauta breeding at two sites off Tasmania, Australia (Albatross Island in western Bass Strait and Pedra Branca to the south) to assess their level of interaction with longline fisheries. Adult birds from both colonies fed locally both in and outside the breeding season. Breeding birds from Albatross Island foraged over the Australian continental shelf or slope waters off northwest Tasmania, while those from Pedra Branca foraged between the colony and the southeastern edge of the continental shelf. The distances travelled by the birds and the duration of their foraging trips varied during the breeding cycle and tended to decrease as eggs approached hatching. Adults which were tracked near the end of the breeding season (March-April, n = 7 birds) deserted their chicks prematurely, and while dispersing further than incubating or brooding birds, they remained over the continental shelf and slope waters off southeast Australia. Home range analyses indicated 41% overlap between foraging zones of birds during successive breeding stages. Dispersal during the postbreeding period extended the foraging zones with less overlap between individuals (10% for Albatross Island and 19% for Pedra Branca). The recent contraction of the Japanese Southern Bluefin Tuna longline fishery to the south and east coasts of Tasmania has resulted in extensive overlap with adult Shy Albatrosses from Pedra Branca, but appears to pose a minimal threat to adult birds from Albatross Island. Coupled with the concomitant increase in the Australian domestic tuna longlining industry, adult Shy Albatrosses from southern Tasmania (Pedra Branca and the Mewstone) are vulnerable to incidental capture through out their annual cycle.     

Benefits of mass reduction for commuting flight with heavy food load in Leach’s storm-petrel,Oceanodroma leucorhoa

When rearing chicks, Leach’s storm-petrels (Oceanodroma leucorhoa) commute between foraging areas and breeding colonies with heavy food loads. At this time they should maximize the size of energy-supplying organs in response to increased energy expenditure but minimize total body mass to decrease the energetic cost of flight. Nineteen storm-petrels were killed to examine the changes in body composition and the masses of energy-supplying organs in birds that were incubating and rearing chicks. Parents lost a mean of 7.95 g in body mass between the stages of incubation and chick-rearing mainly via a loss of skin including subcutaneous adipose tissue, and a small fraction of heart and digestive organs, which are considered energy-supplying organs. This mass loss actually enables them to decrease flight cost by 14.4%. The benefits of decreasing flight costs by reducing total body mass are greater than if the energy-supplying organs of birds are enlarged only.

     

Prevalence of avian pox virus and effect on the fledging success of Laysan Albatross

ABSTRACT.   Avian pox virus ( Poxvirus avium ) is a mosquito-borne disease that occurs worldwide in a variety of bird species, but little is known about its prevalence or effect on seabirds. We monitored prevalence of pox virus and its effect on fledging success of Laysan Albatross ( Phoebastria immutabilis ) on Oahu, Hawaii, from 2003 to 2007. Pox prevalence in albatross chicks averaged 88% in years with high rainfall and 3% in years with low rainfall. Diagnosis of pox virus was clinically confirmed in two birds by Muscovy Duck ( Cairina moschata ) fibrolast cultures. Severity of infection ranged from small wart-like nodules and lesions on the bill, face, eyes, tarsus, and feet, to large tumorous growths that completely covered both eyes and caused deformation of the bill and skull. Most chicks recovered from infection, and the fledging rate in pox epizootic years (82%) did not differ from that in years with low pox prevalence (80%) or the average fledging rate on Midway Atoll (86%). Three chicks with severe infections were resighted as healthy adults on Kauai and Oahu in 2007, confirming postfledging survival of at least some birds. The high recovery rate, fledging success, and postfledging survival indicate that Laysan Albatross have strong immunity to avian pox virus.     

Expensive Traditions: Energy Expenditure of Aleutian Geese in Traditional and Recently Colonized Habitats

ABSTRACT In an effort to reduce goose depredation at a traditional spring migratory stopover site, private landowners implemented a coordinated hazing plan to scare Aleutian cackling geese (Branta hutchinsii leucopareia) from private lands to adjacent public pastures that were cultivated and set aside specifically for geese. Coincidentally, some Aleutian geese began using a new stopover site 150 km farther south in their spring migratory range; numbers at the new site continue to increase. We tested the idea that when their ability to acquire resources deteriorates geese are likely to seek improved foraging conditions, especially during spring migration when individuals strive to maximize nutrient stores and minimize energy expenditure. We quantified measures of goose foraging performance in traditional and new spring staging sites by calculating foraging opportunity, foraging effort, body condition, and daily energy expenditure. Geese staging at the site with higher levels of human disturbance had less foraging opportunity and, despite increased foraging effort and more nutritious food-plants at the site, birds there experienced an elevated energy expenditure and poorer body condition than birds at the new stopover site. Reduced foraging time and increased energy expenditure at the traditional spring staging site may have triggered the colonization process. Suitability assessment of habitat for migratory geese should include measures of foraging opportunity, disturbance risks, and daily energy expenditure in addition to quantity and quality of foods.     

中国鸟类一新纪录种--黑背信天翁(鹱形目,信天翁科)

近期在福州江智明先生的鸟类标本收藏中发现1只1956年3月采自福建沿海的黑背信天翁Diomedea immutabilis标本,为中国鸟类新纪录.     

Restoration of body mass in King Penguins after egg abandonment at a critical energy depletion stage: early vs late breeders

In fasting‐incubating seabirds, it has been proposed that egg abandonment and refeeding should be induced when a low body mass (BM) threshold is attained, thus ensuring adult survival at the expense of immediate breeding. In the context of life‐history trade‐offs in long‐lived birds, we have tested this hypothesis by comparing short‐term survival and restoration of BM in King Penguins Aptenodytes patagonicus that abandoned their egg to those that were relieved normally by their mate at the end of the first incubation shift. Since King Penguins have an extended laying period, the possible influence of seasonal factors was also examined by comparing early and late breeders. Forty incubating males were experimentally forced to fast until egg abandonment by preventing relief by the female. At egg abandonment of both early and late breeding males, BM was below the BM threshold, fasting duration was eight days (about 30%) longer than for relieved birds, and plasma uric acid level was elevated (signature of increased body protein catabolism, phase III of fasting). All abandoning birds survived and came back from sea at a BM similar to that of relieved penguins. The duration of the foraging trip of abandoning early breeders was the same as that of relieved birds, and some abandoning birds engaged in a new breeding attempt. Abandoning late breeders, however, made foraging trips twice as long as those of relieved males. This difference can be explained by time constraints rather than nutritional constraints, abandoning early breeders having enough time left in the breeding season to engage in a new breeding attempt in contrast to abandoning late breeders. These observations lend support to the suggestion that not only BM but also an internal clock intervene in the decision to engage in breeding or not. By preventing a lethal energy depletion ashore and by acting at a fasting stage where the capacity to restore BM at sea is unaffected, abandonment at a low body condition threshold plays a major role in the trade‐off between adult penguin survival and reproduction.     

Feeding ecology of wintering terns in Guinea-Bissau

We studied the feeding ecology of Little Terns Sterna albifrons , Sandwich Terns S. sandvicensis and Royal Terns S. maxima in the Archipélago dos Bijagós (11°40'N, 15°45'W) in Guinea-Bissau (West Africa) during the winter of 1992/1993. More than 95% of all prey taken by these terns were roundfish, ranging in weight from 0.3 to 40 g. Birds usually fed alone, but sometimes they were observed feeding in mixed-species flocks consisting of 15–200 individuals. Capture rate ( n fish per hour foraging) in these flocks was higher than that of solitary birds. However, smaller fish were caught by birds foraging in flocks, so food intake rate (g/h) did not differ between solitary and flock-feeding birds. The relationships between foraging behaviour of the three tern species and abiotic factors, such as time, tide and water clarity, have been investigated. Capture rate of Royal Terns increased with water clarity. For Little Terns and Sandwich Terns, food intake rate was lower in the most turbid waters compared to clearer waters. There was very little foraging activity during high tide. For Little Terns and Royal Terns, food intake rate was about twice as high during receding and low tides as during an incoming tide. Food intake rate averaged 8 g/h in Little Terns, 60 g/h in Sandwich Terns and 45 g/h in Royal Terns. With a rough model, we estimate the maximum rate of daily energy expenditure of terns wintering in the tropics at 3 × BMR (defined as energy expenditure of inactive bird at thermoneutrality in a post-absorptive state during the resting phase of the daily cycle). From an energetic viewpoint, wintering Sandwich Terns in Guinea-Bissau seem to have an easy living.     

Bringing Home the Trash: Do Colony-Based Differences in Foraging Distribution Lead to Increased Plastic Ingestion in Laysan Albatrosses?

When searching for prey, animals should maximize energetic gain, while minimizing energy expenditure by altering their movements relative to prey availability. However, with increasing amounts of marine debris, what once may have been ‘optimal’ foraging strategies for top marine predators, are leading to sub-optimal diets comprised in large part of plastic. Indeed, the highly vagile Laysan albatross (Phoebastria immutabilis) which forages throughout the North Pacific, are well known for their tendency to ingest plastic. Here we examine whether Laysan albatrosses nesting on Kure Atoll and Oahu Island, 2,150 km apart, experience different levels of plastic ingestion. Twenty two geolocators were deployed on breeding adults for up to two years. Regurgitated boluses of undigestable material were also collected from chicks at each site to compare the amount of plastic vs. natural foods. Chicks from Kure Atoll were fed almost ten times the amount of plastic compared to chicks from Oahu despite boluses from both colonies having similar amounts of natural food. Tracking data indicated that adults from either colony did not have core overlapping distributions during the early half of the breeding period and that adults from Kure had a greater overlap with the putative range of the Western Garbage Patch corroborating our observation of higher plastic loads at this colony. At-sea distributions also varied throughout the year suggesting that Laysan albatrosses either adjusted their foraging behavior according to constraints on time away from the nest or to variation in resources. However, in the non-breeding season, distributional overlap was greater indicating that the energy required to reach the foraging grounds was less important than the total energy available. These results demonstrate how a marine predator that is not dispersal limited alters its foraging strategy throughout the reproductive cycle to maximize energetic gain and how this has led to differences in plastic ingestion.     

Foraging strategies of incubating and brooding king penguins Aptenodytes patagonicus

For oceanic birds like king penguins, a major constraint is the separation of foraging areas from the breeding colony, largely because swimming increases foraging costs. However, the relationship between foraging strategy and breeding stage has been poorly investigated. Using time-depth recorders, we studied the diving behaviour of two groups of king penguins that were either incubating or brooding chicks at Crozet Islands (Southern Indian Ocean) at the same period of the year. Although birds with chicks had the highest predicted energy demand, they made foraging trips half as long as incubating birds (6 vs. 14 days) and modified their time and depth utilisation. Birds with chicks dived deeper during daylight (mean maximum depth of 280 m vs. 205 m for those incubating). At night, birds with chicks spent twice as much time diving as those incubating, but birds at both stages never dived beyond 30 m. Movements to greater depths by brooding birds are consistent with the vertical distribution of myctophid fish which are the main prey. As chick provisioning limits trip duration, it is suggested that it is more efficient for parents to change their diving patterns rather than to restrict their foraging range. Received: 23 June 1997 / Accepted: 3 November 1997     

Evaluating the prudence of parents: daily energy expenditure throughout the annual cycle of a free-ranging bird, the macaroni penguin Eudyptes chrysolophus

We measured daily energy expenditure (DEE) continuously for a whole year in a free ranging bird, the macaroni penguin Eudyptes chrysolophus . We combined these measurements with concurrently recorded foraging behaviour, and literature information on body mass and dietary factors to estimate prey consumption rates and foraging success. DEE was at a maximum during late chick-rearing but was equally high during all other active phases of the breeding season. DEE was approximately 4×resting metabolic rate, which accords with established theory and suggests a common 'energetic ceiling' throughout the summer period. However, whether this represents a maximum in physiological capacity, or a rate which optimises fitness is still unclear. Rates of prey consumption and foraging success followed different patterns from daily energy expenditure. Daily prey consumption was high as the penguins prepared for long fasts associated with moulting and incubation but relatively low during chick-rearing, when foraging areas were restricted and foraging success lower. It appears that the energy intake of macaroni penguins is subject to extrinisic or environmental constraints rather than to intrinsic physiological limits.     

Foraging strategy of the little auk Alle alle throughout breeding season – switch from unimodal to bimodal pattern

Energy and time allocation differs between incubation and chick‐rearing periods, which may lead to an adjustment in the foraging behaviour of parent birds. Here, we investigated the foraging behaviour of a small alcid, the little auk Alle alle during incubation and compared it with the chick‐rearing period in West Spitsbergen, using the miniature GPS (in Hornsund) and temperature loggers (in Magdalenefjorden). GPS‐tracking of 11 individuals revealed that during incubation little auks foraged 8–55 (median 46) km from the colony covering 19–239 (median 120) km during one foraging trip. Distance from the colony to foraging areas was similar during incubation and chick‐rearing period. During incubation 89% of foraging positions were located in the zone over shallower parts of the shelf (isobaths up to 200–300 m) with sea surface temperature below 2.5°C. Those environmental conditions are preferred by Arctic zooplankton community. Thus, little auks in the Hornsund area restrict their foraging (both during the incubation and chick‐rearing period) to the area under influence of cold, Arctic‐origin water masses where its most preferred prey, copepod Calanus glacialis is most abundant. The temperature logger data (from 4 individuals) indicate that in contrast to the chick‐rearing period, when parent birds alternated short and long trips, during the incubation they performed only long trips. Adopting such a flexible foraging strategy allows little auks to alter their foraging strategy to meet different energy and time demands during the two main stages of the breeding.     

Night conditions affect morning incubation behaviour differently across a latitudinal gradient

Intermittently incubating birds alternate between sessions of egg warming and recesses for foraging during the day, but stay on the nest continuously at night. Hence, energy costs of nocturnal incubation (which increase during longer and colder nights) cannot be replenished until the next day. Night conditions might therefore be expected to affect morning incubation behaviour the day after. We tested this prediction by exploring latitudinal and seasonal trends in characteristics of the first recess in Eastern Bluebirds Sialia sialis over a 1400-km latitudinal gradient in the continental USA. The time from civil dawn to leaving the nest (latency) increased with latitude early in the breeding season but decreased with latitude late in the season. Birds breeding at higher latitudes also took longer first recesses throughout the season, which led to a larger drop in nest temperature. At the local scale, birds rose earlier after longer nights if the night was also cold, but night length did not predict latency following warm nights. The first recess was longer if the night was warmer, probably because birds could replenish reserves at lower risk of low egg temperature. Our study shows that characteristics of the night led to behavioural changes in features of early morning incubation that were evident at both continental and local scales. These responses also affected nest temperature. Hence, night conditions carry over to incubation behaviour the following morning, which in turn may impose thermal constraints on embryonic development.     

A dynamic model of short-term energy management in small food-caching and non-caching birds

The survival of small birds in winter is critically dependenton the birds' ability to accumulate and maintain safe levelsof energy reserves. In some species, food caching facilitatesenergy regulation by providing an energy source complementaryto body fat. We present a dynamic optimization model of short-term,diurnal energy management for both food-caching and non-caching birds in which only short-day, winter conditions are considered.We assumed that birds can either rest, forage and eat, forageand cache, or retrieve existing caches (the two latter optionsare available only to caching birds). The model predicted thatwhen there is variability in foraging success (here modeledstrictly as within-day variability), both caching and non-caching birds should increase their fat reserves almost linearly inthe morning slowing down toward late afternoon, a result consistentwith field data but different than the result of a previousdynamic program. Non-cachers were predicted to carry higherfat levels than cachers especially when the variability inforaging success is high. Probability of death for non-caching birds was predicted to be higher than that for cachers, especiallyat higher levels of variability in foraging success. Amongcaching birds, an increase in number of caches and fat reserveswas also predicted if: (1) mean foraging success was decreased,(2) variability in foraging success was increased, and (3)energy expenditure at night was increased over our baselineconditions. Under the conditions simulated in our model, birdswere predicted to cache only if cache half-life (i.e., timeinterval over which 50% of the caches are forgotten or lostto pilferage) exceeded 2.5 days, indicating that low pilferagerate and long memory favor more caching. Finally, we showedthat such daily patterns of energy management do not necessarilyrequire relaxing assumptions about mass-dependent predationrisk.     

Barn Swallow Hirundo rustica parents work harder when foraging conditions are good

In altricial birds, the great effort involved in supplying food to nestlings can create trade‐offs in the allocation of resources between the current brood and parental self‐maintenance. In poor foraging conditions, parents have to adjust their energy expenditure in relation to the increased foraging costs. However, intra‐specific variation in parental energy expenditure has rarely been evaluated in the context of these trade‐offs. Here, we quantified the daily energy expenditure (DEE) of parent Barn Swallows Hirundo rustica during the nestling period in relation to foraging conditions while controlling for differences in brood size and nestling age. DEE varied substantially with environmental conditions, increasing by 10 kJ/day per 5 °C in ambient temperature, and by 11 kJ/day per hour in day length. Parent birds did not compensate for a poor aerial insect supply on cool days, but reduced their DEE. Parents only slightly buffered a negative energy balance during chick provisioning with stored body reserves. They did not sacrifice their own energy demands to keep up a high energy flow to the brood when foraging conditions were poor. Instead they worked harder when foraging conditions allowed a surplus intake, fully compensating for their additional efforts, and made maximum use of the rich food supply, allowing the brood to accrue body reserves to compensate for low food intake on cold days. This strategy of energy management may have evolved in the context of the adaptation to the aerial foraging mode and to the ephemeral nature of aerial food resources.     

Do T3 levels in incubating eiders reflect the cost of incubation among clutch sizes?

Complete development of avian eggs requires external heat, inducing in most species an energetic cost of incubation for the parents. Triiodothyronine (T(3)) has been implicated in the control of the metabolic rate and is decreased during fasting in most bird species. This raises the question of the regulation of T(3) during reproduction when incubation (thus heat production) is associated with fasting (and energy sparing). In this study, plasma concentrations of T(3) were studied for different clutch sizes in incubating, as well as in nonincubating, fasting female eiders. Our results show that the T(3) levels decrease during fasting in nonincubating birds, whereas they were maintained during the incubation fast. T(3) levels increased in female eiders at hatching. The plasma T(3) level did not vary among natural clutch sizes in eiders but did so when manipulated. T(3) levels increased when eggs were added (to a maximum of six eggs, i.e., the biggest natural clutch size) or removed (to two eggs, i.e., the smallest natural clutch size). Our results suggest that (1) high T(3) levels during incubation may participate to a threshold of heat production and incubation metabolic rate in eiders despite the fact that they are fasting; (2) since T(3) is associated with the energy expenditure in birds, incubating an enlarged or reduced clutch size may lead to a higher energetic cost of incubation in eiders; and (3) the energy demand of the ducklings at hatching is probably important, as the female T(3) concentrations are then at their highest levels. Thus, any modification of the natural clutch size leads to a rise in the T(3) level of the incubating female, suggesting an additional cost of incubation. Knowing that there is no variation of T(3) levels among natural clutch sizes, this study suggests that a female eider produces a number of eggs corresponding to the energy she can invest in incubation.