The function of habitat change during brood-rearing in the precocial Kentish plover <Emphasis Type="Italic">Charadrius alexandrinus</Emphasis>

Food availability is often variable during the breeding season. Parents with nonmobile, altricial young have no choice but to accept changes in local food availability, whereas in precocial animals, the parents may lead their young away from poor sites to areas that have rich resources and/or are safe from predators. We investigated the latter hypotheses in the Kentish plover Charadrius alexandrinus, a precocial shorebird that raises its young in two habitats: on lakeshore and in saltmarsh. Parents move with their broods from saltmarsh to lakeshore, especially late in the breeding season, and we hypothesized that lakeshores provide more food than the saltmarsh. Consistent with our hypotheses, plover chicks grew faster on the shore, and the difference in growth rates between the two habitats was amplified later in the breeding season. In addition, brood survival was higher on lakeshore than in saltmarsh and decreased with hatching date. Taken together, our results suggest that Kentish plover parents increase their reproductive success by switching brood-rearing habitats strategically.     

How did the Great Auk raise its young?

The extant auks show three strategies of chick rearing – precocial (chicks leave the nest site when a few days old), intermediate (young raised to a mass of around 20% of adult mass) and semi‐precocial (young raised to a mass of around 65% of adult mass). It is not known which strategy the extinct Great Auk used. In this paper, we investigate this issue by a novel combination of a time and energy budget model and phylogenetic comparison. The first approach indicates that for reasonable estimates of the equation parameters, the Great Auk could have followed an intermediate strategy. For a limited range of parameters, the Great Auk could have followed the semi‐precocial strategy. Phylogenetic comparison shows that it is unlikely that the Great Auk followed a precocial strategy. The results suggest that the Great Auk followed an intermediate strategy as does its presumed closest extant relative the Razorbill.     

Role of predation in the decline of kiwi,

Kiwi have declined markedly in abundance and range since human settlement of New Zealand. Three of the four species are still extant in mainland forests, despite decades of co-existence with various introduced mammals. Little spotted kiwi is now probably confined to offshore islands. The role of introduced mammals in these population declines was evaluated by measuring the survival rates of adults, eggs and chicks of brown kiwi (A. mantelli) and great spotted kiwi (A. haastii) in mainland forests. Mortality rates of adults ranged from 5%-16% and did not differ significantly between species or sexes. Overall, 14 out of 209 adult kiwi died during 159.6 radio-tracking years. Predators definitely caused five of these deaths. Sixty-nine (68%) of 102 eggs from 77 nesting attempts by 48 pairs failed to hatch. Predators probably caused about 10% of egg failures. Only three of 49 chicks probably survived to adulthood, indicating a juvenile mortality rate of about 94%. Predators killed at least 8% of chicks, 45% of juveniles, and possibly as many as 60% of all young kiwi. Ferrets and dogs were the main predators of adult kiwi, possums and mustelids were the main egg predators, while stoats and cats were largely responsible for the deaths of young kiwi. Population models show that northern brown kiwi are currently declining at 5.8% per annum. This decline could be halted by cutting the current predation rates on young kiwi by about 34% to 33%.     

Foraging in precocial chicks of the black-tailed godwit Limosa limosa: vulnerability to weather and prey size

Self-feeding precocial development is associated with high energy requirements and potentially vulnerable to short-term reductions in food availability, yet few studies have investigated development of foraging in precocial chicks and its sensitivity to environmental conditions. We studied time budgets and foraging behaviour during the 25-d prefledging period in the insectivorous chicks of a grassland shorebird, the black-tailed godwit Limosa limosa . Until 8–10  d old, parental brooding was the main determinant of chicks' daily foraging time. Brooding decreased with age and temperature and increased during rainfall. Foraging time increased to 70–90% of the daylight period in chicks older than a week, during which distances of 3–12  km  d⁻¹ were covered. Chicks took 98% of their arthropod prey from the grassland vegetation. Prey ingestion rates increased in the first week and slowly declined thereafter, modified by wind speed, temperature and time of day. Chicks in poor body condition were brooded more than chicks growing normally and hence had less feeding time, potentially leading to a negative condition spiral under adverse conditions. However, we found no effect of condition on prey ingestion rate that would preclude recovery when conditions improve. Combining behavioural observations with data on energy expenditure revealed that mean prey size was small (1–4.5  mg), necessitating a high feeding rate, but increased notably after 7–10  d of age. This coincided with a decrease in walking speed, suggesting that chicks fed more selectively. Prey of older chicks approached the upper limit of sizes available in exploitable densities in the grassland vegetation, and this enhances the chicks' sensitivity to variation in prey availability due to weather and agricultural practice.     

Interspecific infanticide deters predators

It is well known that young, small predator stages are vulnerable to predation by conspecifics, intra-guild competitors or hyperpredators. It is less known that prey can also kill vulnerable predator stages that present no danger to the prey. Since adult predators are expected to avoid places where their offspring would run a high predation risk, this opens the way for potential prey to deter dangerous predator stages by killing vulnerable predator stages. We present an example of such a complex predator–prey interaction. We show that (1) the vulnerable stage of an omnivorous arthropod prey discriminates between eggs of a harmless predator species and eggs of a dangerous species, killing more eggs of the latter; (2) prey suffer a minor predation risk from newly hatched predators; (3) adult predators avoid ovipositing near killed predator eggs, and (4) vulnerable prey near killed predator eggs experience an almost fourfold reduction of predation. Hence, by attacking the vulnerable stage of their predator, prey deter adult predators and thus reduce their own predation risk. This provides a novel explanation for the killing of vulnerable stages of predators by prey and adds a new dimension to anti-predator behaviour.     

PTERYLOGRAPHY, PLUMAGE DEVELOPMENT AND MOULT OF JAPANESE QUAIL COTURNIX C. JAPONICA IN CAPTIVITY

Japanese Quail were kept in small cages under controlled conditions of temperature and light, and their pterylography and moult are described. There are 10 primaries, 14 secondaries and corresponding numbers of greater upper and lower wing coverts. The alula has four feathers and the tail from five to six pairs of feathers. There is an apterium in the dorso-pelvic tract similar to that in other quail genera. The arrangement of feathers in the ventral and cervical tracts appears to differ from that described for some North American quail.
The chicks hatch with a covering of natal down. Pre-juvenile moult can be seen when the chicks are three days old. Juvenile body plumage is complete in about 30 days; the sides of the face, around the eyes, are the last places to acquire feathers. The tenth and last juvenile primary to grow is mature when the chicks are 41 days old.
The moult in which the juvenile plumage is replaced overlaps the post-natal moult and in part of the ventral tract natal down is replaced by the first adult feathers. This makes it possible to sex the quail at 14 days old. The first adult moult is complete, in the body tracts, by the time the birds are five to six weeks old. The dropping of juvenile primaries commences at about three weeks old and ceases when about eight weeks old. Only from three to six primaries are replaced; most birds studied replaced five. The significance of this difference from other Galliformes is discussed; it is thought to be associated with the species' migratory behaviour. Quail which remained in the controlled laboratory environment did not undergo any further moult. All birds moulted when both temperature and light period were reduced and most birds moulted when the light period alone was reduced. Adult birds housed in small cages in an unheated, unlit shed underwent a complete moult between August and December in which all primaries were replaced. This moult took 8–14 weeks to complete.     

Prey attack and predators defend: counterattacking prey trigger parental care in predators

That predators attack and prey defend is an oversimplified view. When size changes during development, large prey may be invulnerable to predators, and small juvenile predators vulnerable to attack by prey. This in turn may trigger a defensive response in adult predators to protect their offspring. Indeed, when sizes overlap, one may wonder "who is the predator and who is the prey"! Experiments with "predatory" mites and thrips "prey" showed that young, vulnerable prey counterattack by killing young predators and adult predators respond by protective parental care, killing young prey that attack their offspring. Thus, young individuals form the Achilles' heel of prey and predators alike, creating a cascade of predator attack, prey counterattack and predator defence. Therefore, size structure and relatedness induce multiple ecological role reversals.     

From nestling calls to fledgling silence: adaptive timing of change in response to aerial alarm calls

Young birds and mammals are extremely vulnerable to predators and so should benefit from responding to parental alarm calls warning of danger. However, young often respond differently from adults. This difference may reflect: (i) an imperfect stage in the gradual development of adult behaviour or (ii) an adaptation to different vulnerability. Altricial birds provide an excellent model to test for adaptive changes with age in response to alarm calls, because fledglings are vulnerable to a different range of predators than nestlings. For example, a flying hawk is irrelevant to a nestling in a enclosed nest, but is dangerous to that individual once it has left the nest, so we predict that young develop a response to aerial alarm calls to coincide with fledging. Supporting our prediction, recently fledged white-browed scrubwrens, Sericornis frontalis, fell silent immediately after playback of their parents' aerial alarm call, whereas nestlings continued to calling despite hearing the playback. Young scrubwrens are therefore exquisitely adapted to the changing risks faced during development.     

Field determination of age in male great bustards (Otis tarda) in spring

During a long-term study of individually marked, free-living male great bustards captured as chicks and radio-tracked through several years in Spain, we studied the development with age of two secondary sex traits, the moustachial feathers and the neck plumage pattern. Juvenile males acquired full adult plumage between their fourth and seventh years. The main changes occurred at the neck, coinciding with the onset of sexual maturity. The grey colour typical of immature males turned to ivory white around the fourth to fifth spring, and a gradual increase was appreciated in adults in the brightness of the white colour of the upper neck and in the contrast between this and a progressively more intense chestnut brown at the neck base. Based on these changes, we proposed four neck plumage patterns that can be used to differentiate male age classes during the mating period. The development of moustachial feathers showed more interindividual variability and was not as useful as the neck plumage to estimate male age.     

ADAPTATION, CONSTRAINT, AND COMPROMISE IN AVIAN POSTNATAL DEVELOPMENT

1. This paper discusses factors that influence the evolution of growth rate and determine its variation among species of birds. Growth rate is related to evolutionary fitness through the use of time, energy, and nutrients. In addition, balances between factors favouring rapid growth and those favouring slow growth may be investigated directly by experiment and by comparative observation. 2. David Lack (1968) proposed that the growth rate of the young is the optimum balance between selection for rapid growth to reduce the vulnerable period of development and selection for slow growth to reduce the energy requirements of the young. 3. To test Lack's hypothesis, the growth rates of birds, estimated by fitting sigmoid equations to curves relating weight to age, were surveyed widely from the literature. Among all species examined, growth rate was inversely related to adult weight. Among birds of similar size, most variation in growth rate was related to the degree of maturity of the neonate. Altricial chicks, which depend upon their parents for food and warmth, grow more rapidly than precocial chicks, which are self-sufficient shortly after hatching. Lack's hypothesis, which predicts a direct relationship between growth rate and mortality rate, was not supported. 4. I propose that the key to understanding variation in growth rate among birds lies in the balance between rate of cell proliferation or cell growth, on one hand, and acquisition of mature function, on the other. This idea is consistent with principles of cellular and developmental biology. It is supported by comparisons of (a) the neonates of different species, (b) the individual over the course of the developmental period, and (c) tissues whose use is acquired at different stages of development, wherein more mature individuals or tissues grow more slowly than those with less developed function. 5. Species of birds that are classified as semi-precocial develop precocially but grow rapidly. Although these seemingly violate the general rule relating growth rate to precocity, a closer inspection of their development reveals that they too support the rule. In the Common Tern, the legs, which are the key organ in precocial development, grow at the expected slow rate. The body as a whole grows rapidly because the growth increment of the legs is small and their growth is completed quickly. 6. Growth rates of precocial birds do not decrease abruptly at hatching. This points more to gradual tissue differentiation than to the pattern of procurement and allocation of energy as the primary control for growth rate. 7. Precocious development is favoured when the chicks are capable of self-feeding or when food supplies are distant from the next site and travelling time between one and the other is long. Precocity of the neonates frees both parents to feed at a distant food source. 8. Some species having diets with low levels of protein or other nutrients may grow slowly in order to match nutrient requirements to their availability in the diet. This pattern is indicated especially among the Procellariiformes, which feed an oily diet to their young, and also among tropical fruit-eating birds. 9. Some tropical, pelagically-feeding sea-birds that rear only one offspring at a time may not be able to procure food sufficient to support rapid chick growth. Alternative explanations for slow growth among these species include difficulty in obtaining essential nutrients and more precocious development of activity than in related species having more rapid growth.     

An Experimental Study of Predator Recognition in Great Tit Fledglings

Studies of naturally predator-naïve adult birds (finches on predator-free islands) and birds experimentally hand reared in isolation from predators indicate that birds can recognise predators innately; that is, birds show anti-predator behaviour without former experience of predators. To reduce predation risk efficiently during the vulnerable fledgling period, we would predict an innate response to be fully developed when the chicks leave the nest. However, 30-day-old naïve great tit fledglings ( Parus major ) did not respond differently to a model of a perched predator than to a similarly sized model of a non-predator. Although chicks showed distress responses such as warning calls and freezing behaviour, they did not differentiate between the stimuli. In contrast, wild-caught first-year birds (4 mo old) and adults responded differentially to the two stimuli. Lack of recognition of a perched predator might be one explanation for the high mortality rate found in newly fledged great tits. Our results imply that parental care is not only important for food provisioning, but also to reduce predation risk during the time when fledglings are most vulnerable.     

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.     

Comparative growth rates and oxygen consumption in young Galliformes

The high correlation between growth rate and adult body weight has been much more thoroughly documented for altricial birds than for precocial species. This paper gathers data from the literature for precocial Galliformes and also reports new growth data on six galliform species for analysis. The onset of homeothermic ability is investigated in Galliformes over a range of body size. The results confirm that (1) large species' chicks grow at a slower rate than those of smaller species, and (2) larger species' chicks can thermoregulate earlier than smaller species' chicks under cold stress situations. Published embryonic body weights are also analysed to determine when growth rate differences appear in the development of precocial species. No interspecific differences appeared in the relative growth rates of embryos, and therefore species body size does not appear to influence growth rate before hatching.     

Northern Bobwhite Chick Survival and Effects of Weather

Reliable vital rates for all life stages are necessary to identify limiting factors in wildlife populations and inform sound wildlife management. The difficulty associated with capturing and tagging precocial young, such as northern bobwhite chicks (Colinus virginianus), and linking variation in recruitment to ecological conditions has contributed to significant knowledge gaps in their population ecology. During 1999–2017, we captured and patagial-tagged ≤12-day-old bobwhite chicks (n = 3,576) and estimated their survival from time of capture (Jun–Sep) to fall and winter (Nov and Jan) recapture and winter (Feb) recovery. We used Burnham's model implemented in Program MARK to integrate mark-recapture and dead-recovery (via harvest) data to estimate survival for the 19-year study. By including weather covariates, we also evaluated explicit hypotheses related to temperature and precipitation effects on chick survival. We found inter-annual, intra-annual, and intra-seasonal variation in chick survival with an average annual daily survival estimate of 0.9887 (95% CI = 0.9321, 0.9918). Precipitation amount and number of precipitation (>0.635 cm) events during the first 4 weeks post-tagging decreased the daily survival rate of chicks curvilinearly. Average minimum daily temperature and maximum daily temperature effects on survival were negligible, but an interaction between minimum temperature and cumulative precipitation during the first 14 days post-tagging affected survival. We recommend population modelers incorporate intra- and inter-annual variation in chick survival to improve predictions. Observed variation in chick survival rates portends a significant opportunity to improve population management for bobwhites and other game birds. © 2019 The Wildlife Society     

Comparative analyses of the influence of developmental mode on phenotypic diversification rates in shorebirds

Phenotypic diversity is not evenly distributed across lineages. Here, we describe and apply a maximum-likelihood phylogenetic comparative method to test for different rates of phenotypic evolution between groups of the avian order Charadriiformes (shorebirds, gulls and alcids) to test the influence of a binary trait (offspring demand; semi-precocial or precocial) on rates of evolution of parental care, mating systems and secondary sexual traits. In semi-precocial species, chicks are reliant on the parents for feeding, but in precocial species the chicks feed themselves. Thus, where the parents are emancipated from feeding the young, we predict that there is an increased potential for brood desertion, and consequently for the divergence of mating systems. In addition, secondary sexual traits are predicted to evolve faster in groups with less demanding young. We found that precocial development not only allows rapid divergence of parental care and mating behaviours, but also promotes the rapid diversification of secondary sexual characters, most notably sexual size dimorphism (SSD) in body mass. Thus, less demanding offspring appear to facilitate rapid evolution of breeding systems and some sexually selected traits.     

Brood amalgamation in the Bristle-thighed Curlew Numenius tahitiensis: process and function

Alloparental care in birds generally involves nonbreeding adults that help at nests or breeding adults that help raise young in communal nests. A less often reported form involves the amalgamation of broods, where one or more adults care for young that are not their own. We observed this phenomenon among Bristle-thighed Curlew Numenius tahitiensis broods in western Alaska during 1990–1992. Amalgamation of broods generally involved the formation of temporary and extended associations. Temporary associations were formed by the incidental convergence of broods soon after they left their nests. During this period, parents defended distinct brood-rearing areas, were antagonistic to conspecifics and remained together for less than 3 days. Extended associations formed when chicks were 1–2 weeks old. Here, parents and their broods occupied distinct, but adjacent, brood-rearing areas and moved around as a unit. Whether a brood participated in either temporary or extended associations or remained solitary appeared to depend on brood density in the immediate area and on hatching date. When chicks were 3–4 weeks old, aggregations of up to ten broods formed wherein young mixed and parents defended a common brood-rearing area. All broods (n = 48) that survived to fledging joined such aggregations. Alloparental care involved only antipredator defence and was not associated with activities such as feeding and brooding. Most female parents abandoned their broods shortly after the young could fly and when aggregations were forming. The female parent of a pair always deserted its young before or on the same day as the male parent and, in every aggregation, one or two males continued to tend young for about 5 days longer than other male parents. In most cases, adults deserted the young 2–6 days before the young departed the area when about 38 days old. Bristle-thighed Curlews also formed temporary associations with American and Pacific Golden Plover Pluvialis dominica and Pluvialis fulva, Whimbrel Numenius phaeopus, Bar-tailed Godwit Limosa lapponica, Western Sandpiper Cal-idris mauri and Long-tailed Skua Stercorarius longicaudus. Curlews and other larger bodied species commonly attack-mobbed predators together, whereas smaller bodied species generally gave alarm calls and circled the predators. For all species, the intensity of antipredator defence by attending adults gradually decreased as young became older and aggregations formed. We suggest that amalgamation of broods among Bristle-thighed Curlew enhances predator defence, aids in the process of flock formation for migrating young, and allows females and some males to desert their young earlier.     

Load Lightening in Southern Lapwings: Group‐Living Mothers Lay Smaller Eggs than Pair‐Living Mothers

Females of some cooperative‐breeding species can decrease their egg investment without costs for their offspring because helpers‐at‐the‐nest compensate for this reduction either by feeding more or by better protecting offspring from predation. We used the southern lapwing (Vanellus chilensis) to evaluate the effects of the presence of helpers on maternal investment. Southern lapwings are cooperative (some breeding pairs are aided by helpers), chick development is precocial, thus adults do not feed the chicks, and adults offer protection from predators through mobbing behaviors. We tested whether southern lapwing females reduced their reproductive investment (i.e. load‐lightening [LL] hypothesis) or increased their investment (i.e. differential allocation hypothesis) when breeding in groups when compared with females that bred in pairs. We found that increased group size was associated with lower egg volume. A significant negative association between the combined egg nutritional investment (yolk, protein, and lipid mass) and group size was observed. Chicks that hatched from eggs laid in nests of groups were also smaller than chicks hatched in nests of pairs. However, there was no relationship between the body mass index of chicks, or clutch size and group size, which suggests that such eggs are, simply, proportionally smaller. Our results support the LL hypothesis even in a situation where adults do not feed the chicks, allowing females to reduce investment in eggs without incurring a cost to their offspring.     

Precocial development of locomotor performance in a ground-dwelling bird (Alectoris chukar): negotiating a three-dimensional terrestrial environment

Developing animals are particularly vulnerable to predation. Hence, precocial young of many taxa develop predator escape performance that rivals that of adults. Ontogenetically unique among vertebrates, birds transition from hind limb to forelimb dependence for escape behaviours, so developmental investment for immediate gains in running performance may impair flight performance later. Here, in a three-dimensional kinematic study of developing birds performing pre-flight flapping locomotor behaviours, wing-assisted incline running (WAIR) and a newly described behaviour, controlled flapping descent (CFD), we define three stages of locomotor ontogeny in a model gallinaceous bird (Alectoris chukar). In stage I (1–7 days post-hatching (dph)) birds crawl quadrupedally during ascents, and their flapping fails to reduce their acceleration during aerial descents. Stage II (8–19 dph) birds use symmetric wing beats during WAIR, and in CFD significantly reduce acceleration while controlling body pitch to land on their feet. In stage III (20 dph to adults), birds are capable of vertical WAIR and level-powered flight. In contrast to altricial species, which first fly when nearly at adult mass, we show that in a precocial bird the major requirements for flight (i.e. high power output, wing control and wing size) convene by around 8 dph (at ca 5% of adult mass) and yield significant gains in escape performance: immature chukars can fly by 20 dph, at only about 12 per cent of adult mass.     

Aggression by adult South American Terns toward conspecific chicks

ABSTRACT Infanticide and nonfatal aggression by adults toward unfamiliar chicks have been widely reported in colonial birds, and can be an important cause of chick mortality. We studied intraspecific aggression by adults toward chicks at a South American Tern (Sterna hirundinacea) colony in Patagonia, Argentina, during 2005 to characterize this behavior, evaluate its relationship with nesting density, chick age and microhabitat characteristics, and assess its effect on breeding success. Of 111 chicks in the study area, 45% were attacked at least once. Chicks older than 9 d posthatching were more likely to be attacked than younger chicks, and unattended chicks were more likely to be attacked than guarded chicks (88 vs. 12%). Chicks were also attacked more often when in their own territories (76% of cases), but were less likely to be attacked in territories with more vegetation cover and high‐quality shelters (i.e., vegetation with characteristics that prevented adults from reaching chicks). The number of aggression events was not related to nest density. At least 8% of the chicks in our study area died as a result of adult intraspecific aggression. Our results indicate that intraspecific aggression by adult South American Terns toward chicks is relatively common in the Punta Loma colony and should not be underestimated as a factor affecting their breeding success.     

Weather,microclimate, and energy costs of thermoregulation for breeding Adélie Penguins

Summary We measured meteorological conditions and estimated the energy costs of thermoregulation for young and adult Adélie Penguins (Pygoscelis adeliae) at a breeding colony near the Antarctic Peninsula. Air temperatures averaged < 5°C and strong winds were frequent. Operative temperatures (T_e) for adults ranged from –8 to 28°C, averaging 5–6°C, for the period from courtship to fledging of chicks. The average energy cost of thermoregulation (C_th) for adult penguins was equivalent to 10–16% of basal metabolism. C_th comprised about 15% of the estimated daily energy budget (DEB) of incubating adults, but only about 1% of the DEB of adults feeding chicks. The T_e's for chicks older than 14 days ranged from 0 to 31°C, averaging 8.0 C. The C_th for downy chicks ranged from about 31% of minimal metabolic rate (MMR) in 1 kg chicks to about 10% of MMR in 3 kg chicks. Between initial thermal independence (age 12–14 days) and the cessation of parental feeding (age 35–40 days), chicks use about 10–11% of assimilated energy for thermoregulation. C_th is equivalent to about 17% of the MMR of fledglings during their 2–3 week fast. We observed no indication of thermal stress (i.e., conditions in which birds cannot maintain stable T_b) in adults and no indication of cold stress in any age class. However, on clear, calm days when air temperature exceeds 7–10°C for several hours, downy chicks are vulnerable to lethal hyperthermia.