Factors affecting fledging weight of Adélie penguin (Pygoscelis adeliae) chicks: a modeling study

An individual-based model is developed to examine mechanisms that potentially underlie the observed constancy in fledging weight (2.8-3.2 kg) of Adélie (Pygoscelis adeliae) penguin chicks, in spite of large variability in the abundance of Antarctic krill (Euphausia superba), the primary food source. The model describes the energetic requirements of the chick, with growth resulting from the difference between assimilated energy and respiration. Parameterizations of these metabolic processes are based upon experimental and field observations. Ingestion of Antarctic krill by the chick is dependent on the frequency of food delivery to the chick by the adults, which is based on measured foraging times. The mass, size, and size frequency distribution of Antarctic krill fed to the chick are specified using empirical data. The energy content of the Antarctic krill provided to the chick is taken to be constant or allowed to vary with size. The simulations show that food availability is most critical in the latter portion of chick development, when growth rates and food demands are high. Low food availability during this time must be compensated by either feeding chicks with larger krill of higher caloric value or by increased assimilation efficiency. Periods when small krill with lower caloric value dominate require more frequent feeding of the chicks in order to attain their observed fledging weight. Thus, although the total food energy given to the chick is the main factor determining chick growth, the distribution of food availability relative to chick size (i.e., different net growth rates) and food quality are also factors influencing the fledging weight of penguin chicks. The simulations provide insight into the compensating effects of food delivery, food quality, and metabolic processes that allow Adélie penguin chicks to reach their observed fledging weight in spite of considerable environmental variability in food supply.     

Validating growth and development of a seabird as an indicator of food availability: captive‐reared Caspian Tern chicks fed ad libitum and restricted diets

ABSTRACT For seabirds raising young under conditions of limited food availability, reducing chick provisioning and chick growth rates are the primary means available to avoid abandonment of a breeding effort. For most seabirds, however, baseline data characterizing chick growth and development under known feeding conditions are unavailable, so it is difficult to evaluate chick nutritional status as it relates to foraging conditions near breeding colonies. To address this need, we examined the growth and development of young Caspian Terns (Hydroprogne caspia), a cosmopolitan, generalist piscivore, reared in captivity and fed ad libitum and restricted (ca. one‐third lower caloric intake) diets. Ad libitum‐fed chicks grew at similar rates and achieved a similar size at fledging as previously documented for chicks in the wild and had energetic demands that closely matched allometric predictions. We identified three general characteristics of food‐restricted Caspian Tern chicks compared to ad libitum chicks: (1) lower age‐specific body mass, (2) lower age‐specific skeletal and feather size, such as wing chord length, and (3) heightened levels of corticosterone in blood, both for baseline levels and in response to acute stress. Effects of diet restriction on feather growth (10–11% slower growth in diet‐restricted chicks) were less pronounced than effects on structural growth (37–52% slower growth) and body mass (24% lower at fledging age), apparently due to preferential allocation of food resources to maintain plumage growth. Our results suggest that measurements of chick body mass and feather development (e.g., wing chord or primary length) or measurement of corticosterone levels in the blood would allow useful evaluation of the nutritional status of chicks reared in the wild and of food availability in the foraging range of adults. Such evaluations could also inform demography studies (e.g., predict future recruitment) and assist in evaluating designated piscivorous waterbird conservation (colony) sites.     

Limited effects of egg size on chick growth in the chinstrap Penguin Pygoscelis antarctica

We studied the effect of egg size on chick growth in the chinstrap penguin under natural non-manipulative, conditions. The influence of egg size on chick mass at hatching, 15 and 45 days of age was analysed controlling for the effect of hatching date, as hatching date has been repeatedly associated with chick growth in this species. Egg volume explained 20% of the variation in the body mass of chicks at hatching. Its effect disappeared as chicks grew, being statistically not significant at the age of 15 and 45 days. Egg volume asymmetry also had no role in determining chicks' growth asymmetries. Accepted: 6 June 1998     

Accelerated growth rates in late‐hatched Rhinoceros Auklet Cerorhinca monocerata chicks depend on food conditions and growth stage: an experimental approach

In some bird species, the survival of chicks hatching later in the season is lower than those hatched earlier due to increased risk of predation and a seasonal decline in feeding conditions. To reduce these risks, it might be advantageous for late‐hatched chicks to grow faster and hence fledge at younger age. In this experimental study, the growth rates of early‐ and late‐hatched Rhinoceros Auklet Cerorhinca monocerata chicks were compared under average and poor food supplies in captivity. Controlling for potentially confounding effects of chick mass at 10 days old, chick age and nest‐chamber temperature, late‐hatched chicks had higher wing growth rate than early‐hatched chicks before attaining the minimum wing length required for fledgling under both average and poor food supplies. After attaining the minimum wing length, however, late‐hatched chicks had a lower fledging mass, indicating a potential cost that could diminish the early advantage of fast wing growth.     

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     

Energetic costs of raising Pygoscelid penguin chicks

Energy requirements of resting Adélie (Pygoscelis adeliae), Gentoo (P. papua) and Chinstrap (P. antarcticd) penguin chicks were determined with respect to body mass via respirometry in Antarctica. Resting metabolic rates of all Pygoscelid penguin chicks were similar (ANOVA p=0.91) and best described by E=0.0096 M^0.98 (n=24, r²=0.97), where E is power (W) and M is mass (g). Using the results obtained here and data published in the literature, I determined a) the amount of food needed from hatching to fledging as 29.8, 31.7 and 56.4 kg per chick for Adélie, Chinstrap and Gentoo penguins, respectively and b) the average amount of food left daily to the parent after feeding the brood throughout the breeding period. Parents keep only a minimum of food for themselves just prior to the time when chicks begin forming crèches. Thereafter, nest relief intervals are increased, and the amount of food parents can keep for themselves rises. The results of both models are discussed with respect to available data on Pygoscelid penguin food requirements.     

Time‐lapse cameras reveal latitude and season influence breeding phenology durations in penguins

Variation in the phenology of avian taxa has long been studied to understand how a species reacts to environmental changes over both space and time. Penguins (Sphenicidae) serve as an important example of how biotic and abiotic factors influence certain stages of seabird phenology because of their large ranges and the extreme, dynamic conditions present in their Southern Ocean habitats. Here, we examined the phenology of gentoo (Pygoscelis papua) and chinstrap penguins (Pygoscelis antarctica) at 17 sites across the Scotia arc, including the first documented monitoring of phenology on the South Sandwich Islands, to determine which breeding phases are intrinsic, or rather vary across a species range and between years. We used a novel method to measure seabird breeding phenology and egg and chick survival: time‐lapse cameras. Contrary to the long‐standing theory that these phases are consistent between colonies, we found that latitude and season had a predominant influence on the length of the nest establishment, incubation, and guard durations. We observe a trend toward longer incubation times occurring farther south, where ambient temperatures are colder, which may indicate that exposure to cold slows embryo growth. Across species, in colonies located farther south, parents abandoned nests later when eggs were lost or chicks died and the latest record of eggs or chicks in the nest occurred earlier during the breeding period. The variation in both space and time observed in penguin phenology provides evidence that the duration of phases within the annual cycle of birds is not fundamental, or genetic, as previously understood. Additionally, the recorded phenology dates should inform field researchers on the best timing to count colonies at the peak of breeding, which is poorly understood.     

Hand-Rearing,Release and Survival of African Penguin Chicks Abandoned Before Independence by Moulting Parents

The African penguin Spheniscus demersus has an ‘Endangered’ conservation status and a decreasing population. Following abandonment, 841 African penguin chicks in 2006 and 481 in 2007 were admitted to SANCCOB (Southern African Foundation for the Conservation of Coastal Birds) for hand-rearing from colonies in the Western Cape, South Africa, after large numbers of breeding adults commenced moult with chicks still in the nest. Of those admitted, 91% and 73% respectively were released into the wild. There were veterinary concerns about avian malaria, airsacculitis and pneumonia, feather-loss and pododermatitis (bumblefoot). Post-release juvenile (0.32, s.e.  = 0.08) and adult (0.76, s.e.  = 0.10) survival rates were similar to African penguin chicks reared after oil spills and to recent survival rates recorded for naturally-reared birds. By December 2012, 12 birds had bred, six at their colony of origin, and the apparent recruitment rate was 0.11 (s.e.  = 0.03). Hand-rearing of abandoned penguin chicks is recommended as a conservation tool to limit mortality and to bolster the population at specific colonies. The feasibility of conservation translocations for the creation of new colonies for this species using hand-reared chicks warrants investigation. Any such programme would be predicated on adequate disease surveillance programmes established to minimise the risk of disease introduction to wild birds.     

Common loon parents defend chicks according to both value and vulnerability

In many territorial breeders, conspecifics that intrude during the chick‐rearing period pose a threat to survival of young. Defense of young from intruders is costly to parents, so it is likely that intense selective pressure has shaped chick defense so as to maximize parental fitness. We simulated territorial intrusion by exposing adult common loons Gavia immer and their chicks to a decoy and used mixed models to investigate responses. We tested two hypotheses: 1) the value hypothesis, which holds that parents should defend large broods of offspring more strongly because of the greater potential fitness benefits they offer, and 2) the vulnerability hypothesis, which predicts vigorous defense of young offspring, whose small size and limited mobility render them vulnerable to sudden attacks from intruders that approach under water. Under natural conditions, parents spent over 80% of their time within 20 m of chicks younger than two weeks (‘young chicks’) but 66% or less of their time close to chicks four weeks or older (‘old chicks’). Parents of young chicks associated less with the decoy but yodelled and penguin danced more during decoy trials than did parents of old chicks, supporting the conclusion that the parents protected young chicks not by engaging intruders directly but by remaining close to chicks and using vocalization and display to keep intruders at a distance. While these findings lent clear support to the vulnerability hypothesis, the value hypothesis too was supported, as males with two‐chick broods were almost three times more likely to yodel than males with singleton chicks. Age of parents was not associated with any aspect of chick defense, but the paucity of known‐aged parents in the oldest age classes makes future investigation of age effects warranted.     

Satellite data identify decadal trends in the quality of Pygoscelis penguin chick‐rearing habitat

Pygoscelis penguins are experiencing general population declines in their northernmost range whereas there are reported increases in their southernmost range. These changes are coincident with decadal‐scale trends in remote sensed observations of sea ice concentrations (SIC) and sea surface temperatures (SST) during the chick‐rearing season (austral summer). Using SIC, SST, and bathymetry, we identified separate chick‐rearing niche spaces for the three Pygoscelis penguin species and used a maximum entropy approach (MaxEnt) to spatially and temporally model suitable chick‐rearing habitats in the Southern Ocean. For all Pygoscelis penguin species, the MaxEnt models predict significant changes in the locations of suitable chick‐rearing habitats over the period of 1982–2010. In general, chick‐rearing habitat suitability at specific colony locations agreed with the corresponding increases or decreases in documented population trends over the same time period. These changes were the most pronounced along the West Antarctic Peninsula where there has been a rapid warming event during at least the last 50 years.     

Tracking day and night provides insights into the relative importance of different wader chick predators

Poor reproductive success driven by nest and chick predation severely limits the population recovery of waders breeding on lowland wet grassland. Managing predation requires knowledge of the predators and because these can be grouped into nocturnal or diurnal hunters, detecting the timing of predation can help assess their relative impacts. Wader nest studies investigating the timing of egg predation have identified nocturnal mammals, primarily Red Foxes Vulpes vulpes, as the most important nest predators, but quantifying predator importance for highly mobile wader chicks is more difficult. Manual radiotelemetry can detect whether chicks are alive but cannot detect the time of predation, and predator identity can be determined only in the few cases where remains are recovered. As an alternative we used automatic radio tracking stations (ARTS) to constantly record the signals and predation timing of 179 radiotagged Lapwing Vanellus vanellus chicks, combining this with manual telemetry, inference about predator identity from predated remains and site‐level Fox, mustelid and avian predator activity monitoring. This approach succeeded in detecting the time of predation for 60% of the 155 chicks that were predated. Diurnal chick predation accounted for a larger number of predation events, but nocturnal predation was more intensive in terms of predation likelihood per hour. Mammalian predation during both day and night had a larger impact on chick survival than did avian predation. Raptors were primarily responsible for predation by birds and Foxes for predation by mammals, with Foxes also having a larger influence on daily chick predation rates than other predators. Chick predation increased seasonally, implying that earlier‐hatching breeding attempts are more likely to be successful. Higher Fox, raptor and mustelid activity resulted in higher proportions of chicks being predated by those predators, so quantifying the activity of those three predator groups on a site could be a quicker alternative to studying chicks when investigating which predator species to target with site‐specific predation management.     

Great Spotted Cuckoo Nestlings but not Magpie Nestlings Starve in Experimental Age‐Matched Broods

Nestlings of non‐evicting avian brood‐parasites have to compete for food with foster parents' own nestlings. The outcome of these competitive contests is determined mainly by body size differences between parasitic and host nestlings. As part of the coevolutionary arms race between brood parasites and their hosts at the nestling stage, it has been reported that some host foster parents discriminate against parasitic chicks and are reluctant to feed them. Here, by experimentally creating size‐matched broods of different composition (only magpie Pica pica chicks, only great spotted cuckoo Clamator glandarius chicks or mixed broods), we show that great spotted cuckoo chicks starved in 20.2 per cent (17 of 84) of the parasitized magpie nests even in absence of size asymmetries, while in none (0 of 72) of the nests a magpie chick starved. As far as we know, this is the first record of non‐evictor brood parasitic nestlings starving without being smaller than their host nestmates in a frequently used host species. Nest composition had no effect on chick starvation. The cuckoo nestling starved even in two of the nests occupied by only one cuckoo chick. Our results could be explained by (1) magpies being reluctant to feed cuckoo chicks; (2) parasitic chicks receiving lower‐quality food items or cuckoo nestlings being sensitive to some particular component of the diet (e.g. cereal grains); and (3) the existence of cuckoo chick discrimination ability by magpie foster parents.     

Ecological mismatches are moderated by local conditions for two populations of a long‐distance migratory bird

Ecological mismatches between reproductive events and seasonal resource peaks are frequently proposed to be a key driver of population dynamics resulting from global climate change. Many local populations are experiencing reduced reproductive success as a consequence of mismatches, but few mismatches have led to species‐level population declines. To better understand this apparent paradox, we investigated the breeding phenology and chick survival of two disjunct populations of Hudsonian godwits Limosa haemastica breeding at Churchill, Manitoba and Beluga River, Alaska. Only one population experienced a mismatch: godwits bred nearly one week after the onset of the invertebrate peak at Churchill because of asynchronous climatic change occurring throughout their annual cycle. However, chicks were not uniformly affected by the mismatch — growth rates and survival of young chicks were not correlated with invertebrate abundance, but older chicks tended to suffer lower survival rates on days of low invertebrate abundance. Ecological mismatches thus resulted in a complex array of consequences, but nonetheless contributed to reductions in chick survival. In contrast, godwits at Beluga River hatched their chicks just before the invertebrate peak, such that the period of highest energetic need coincided with the period of highest invertebrate abundance. As a result, growth rates and survival of godwit chicks were unaffected by invertebrate abundance. Godwits at Beluga River were able to properly time their reproduction because of predictable rates of climatic change and strong selection imposed by high predation on late‐hatched chicks. Taken together, our results suggest that population‐specific, local‐scale selection pressures play a critical role in determining the degree and severity of ecological mismatches. The potential for global climate change to induce species‐level population declines may therefore be mediated by the spatial variation in the selection pressures acting across a species’ range.     

Chick‐provisioning behaviour of two shearwaters breeding in south‐eastern Australia

The chick‐provisioning behaviour of the short‐tailed shearwater Puffinus tenuirostris and the wedge‐tailed shearwater Puffinus pacificus was investigated in a mixed colony on Montague Island, New South Wales, Australia, over two breeding seasons. This colony is located at opposite edges of the breeding distribution of the two species. Frequent weighing techniques were used to determine chick feeding frequency, feed timing, meal size, chick weight loss and indices of food conversion efficiency of the chicks. Short‐tailed shearwater parents fed their chicks larger more infrequent meals than wedge‐tailed shearwater parents. Short‐tailed shearwater chicks demonstrated higher food conversion efficiencies and lower weight loss than wedge‐tailed shearwater chicks, indicating either differences in diet or metabolic rates. The feeding frequency in wedge‐tailed shearwaters also fluctuated more widely than for short‐tailed shearwaters over the two breeding seasons. Despite the fact that the timing of the breeding cycle on Montague Island is almost identical for the two species, these differences in chick provisioning are probably a result of differences in prey type and location, so they may help explain variations in annual breeding success and limits to the distribution of the two species.     

A comprehensive demographic assessment of the endangered Fiordland crested penguin Eudyptes pachyrhynchus

ABSTRACT

Many species of Eudyptes penguin have shown substantial population declines and in response, there have been efforts to identify the key demographic parameters. Here, we present the demographic parameters of one of the least well known and the least abundant species of crested penguin, the endangered Fiordland crested penguin Eudyptes pachyrhynchus. A population study incorporating mark–recapture, nest occupancy and breeding success was conducted over 16 years at several sites in the northern half of the range of the species. Survival probabilities were calculated using standard Cormack–Jolly–Seber models and the Burnham Live and Dead model. The annual probability of true survival for banded birds and apparent survival for birds with transponders were both estimated at 89% during their adult years, which is similar to that reported for Eudyptes penguin species inhabiting more southerly latitudes. Annual juvenile survival was assessed for Fiordland crested penguins until their first return at 77%. The mean breeding success (0.61?±?0.02 chicks/pair) was higher than is observed for other crested penguin species, except the southern rockhopper penguin, which may be due to having lower A-egg ejection rates and higher rates of fledging two chicks per pair. Breeding success was related to the niche of predators present.     

Climate fluctuation effects on breeding of blue penguins (Eudyptula minor)

Abstract

El Niño and La Niña climate perturbations alter sea currents and food availability for seabirds in many areas of the world. This changes their breeding success and mortality. Blue penguin (Eudyptula minor) breeding success is dependent upon whether one or two clutches per season are laid, and the hatching and fledging success of these clutches. This study uses six years of data from five blue penguin breeding colonies, three from Taiaroa Head, Otago Peninsula and two from Oamaru, to examine whether annual variation in breeding success correlates with El Niño/La Niña perturbations. When La Niña conditions prevailed, penguins started breeding later, and there was a lower proportion of double breeders than in El Niño and normal years. The probability of a newly hatched chick surviving to fledging was also dependent on whether large‐scale climatic conditions prevailed, whereas hatching success and overall breeding success (number of fledged chicks per breeding pair) showed no correlation with climate perturbations.     

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.     

Ontogeny of chick behavior: a tool for monitoring the growth and development of lesser adjutant storks

We studied the behavioral development of seven lesser adjutant stork (Leptoptilos javanicus) chicks from hatching to fledging over three breeding seasons at the Bronx Zoo. We developed an ethogram and compared the rate at which behaviors appeared in relation to brood size, sex, and the conditions in which the chicks were raised by their parents. Although sample sizes were small, there seem to be sex‐related differences in the rate at which behaviors develop, with females developing more rapidly than males. Larger clutch size may be associated with slower growth rate because the single male developed faster than the two males in 2004. The slowest growth rate, observed in a single male chick in 1999, was most likely owing to nutritional deficiencies and other health complications. More research is needed, but these results can be used to help evaluate the age and health of lesser adjutant stork chicks in captivity and in the wild. Zoo Biol 26:533–538, 2007. © 2007 Wiley‐Liss, Inc.     

Relevance of forage fish in the diet of Magellanic penguins breeding in northern Patagonia,Argentina

We quantified the trophic niche of Magellanic penguins (Spheniscus magellanicus) breeding and moulting in Golfo San Jorge, Argentina, through conventional stomach content and stable isotope analysis. A total of 112 adults were flushed during the early and late chick stages of 2011 and 2012 at Isla Vernacci Norte, and at least 15 prey taxa were found, including fishes, cephalopods, crustaceans and polychaetes. Overall, Argentine anchovy (Engraulis anchoita) showed the highest contribution in terms of importance by mass (68.1–85.3%, depending on chick stage and year), except for the old chick stage in 2011 when the shortfin squid (Illex argentinus) was the main prey consumed (56.0%). Based on carbon and nitrogen isotopic values from a total of 256 blood samples, corresponding to young and old chicks and to adults of both sexes sampled throughout the incubation, chick and moult stages at the above mentioned colony and years, Bayesian mixing model outputs showed that Argentine anchovy was always the main prey (48–86%). Bayesian mixing model outputs obtained from adults of both sexes and their chicks during the late chick stage of 2013 at Isla Vernacci Norte, Isla Tova and Isla Leones also showed that Argentine anchovy was the main prey consumed. This is the first comprehensive assessment of Magellanic penguin diet composition in northern Patagonia, quantifying the relative contribution of prey in the diet of adults and chicks at different stages of the annual cycle and years, and confirms the relevance of a forage fish such as the Argentine anchovy in its trophic ecology.     

Darwin's Finch Begging Intensity Does Not Honestly Signal Need in Parasitised Nests

Parental care should be selected to respond to honest cues that increase offspring survival. When offspring are parasitised, the parental food compensation hypothesis predicts that parents can provision extra food to compensate for energy loss due to parasitism. Chick begging behaviour is a possible mechanism to solicit increased feeding from attending parents. We experimentally manipulated parasite intensity from Philornis downsi in nests of Darwin's small ground finch (Geospiza fuliginosa) to test its effects on chick begging intensity and parental food provisioning. We used in‐nest video recordings of individually marked chicks to quantify nocturnal parasite feeding on chicks, subsequent diurnal chick begging intensity and parental feeding care. Our video analysis showed that one chick per brood had the highest parasite intensity during the night (supporting the tasty chick hypothesis) and weakest begging intensity during the day, which correlated with low parental care and rapid death. We observed sequential chick death on different days rather than total brood loss on a given day. Our within‐nest video images showed that (1) high nocturnal larval feeding correlated with low diurnal begging intensity and (2) parent birds ignored weakly begging chicks and provisioned strongly begging chicks. Excluding predation, all parasite‐free chicks survived (100% survival) and all parasitised chicks died in the nest (100% mortality). Weak begging intensity in parasitised chicks, which honestly signalled recent parasite attack, was not used as a cue for parental provisioning. Parents consistently responded to the strongest chick in both parasitised and parasite‐free nests.