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.     

INVERSE RELATIONSHIP BETWEEN FUNCTIONAL MATURITY AND EXPONENTIAL GROWTH RATE OF AVIAN SKELETAL MUSCLE: A CONSTRAINT ON EVOLUTIONARY RESPONSE

In this study, we investigate whether a tissue-level constraint can explain the general inverse relationship between growth rate and precocity of development in birds. On the whole, altricial (dependent) chicks grow three to four times faster than the less dependent, more able chicks of precocial species of similar adult mass. We suggest that an antagonism between growth and acquisition of mature function in skeletal muscle constrains postnatal growth and development in most species of birds. Altricial species, represented by European starlings in this study, hatch with skeletal muscle having low capacity for generating force but grow rapidly. Conversely, precocial species (northern bobwhite quail and Japanese quail), hatch with relatively mature skeletal muscle, especially in their legs, but grow more slowly. As development proceeds in all species, exponential growth rates decrease as muscles acquire adult levels of function. Among four variables associated with muscle function, exponential growth rate (EGR) was negatively correlated with pyruvate kinase activity (glycolysis), potassium concentration (electrical potential), and dry weight fraction (contractile proteins) in both pectoral and leg muscles but not with citrate synthase activity (aerobic metabolism) in either set of muscles. For pectoral muscle, these variables accounted for 87% of the total variation in EGR in all three species combined despite a twofold difference in growth rates between the starling and quail. EGRs of leg muscle (51% of variation accounted for) were less than predicted by the pectoral-muscle equation in quail during the early part of the postnatal period and in starlings during the late postnatal period. This result would not contradict a growth rate/maturity constraint hypothesis if EGRs were down-regulated for allometric or other considerations.     

Energetics of growth in semi-precocial shorebird chicks in a warm environment: the African black oystercatcher, Haematopus moquini

We studied prefledging growth, energy expenditure and time budgets of African Black Oystercatcher, Haematopus moquini, chicks on Robben Island, Western Cape, South Africa. The aim of the study was to investigate the effect of parental feeding on the growth and energetics of semi-precocial shorebird chicks. Chicks reached mean fledging mass, 463 g, in 40 days. The growth rate coefficient of African Black Oystercatcher chicks was 2% below the predicted value for a shorebird species of their body mass, but it was smaller than that of other precocial and semi-precocial shorebirds to date. Resting metabolic rate (RMR, measured through respirometry), daily metabolisable energy (DME), defined as daily energy expenditure (DEE, measured with doubly labelled water) plus energy deposited into tissue (E(tis)), and total metabolisable energy (TME) of African Black Oystercatcher chicks were similar to those expected for a species of their body size. DEE was not influenced by weather (ambient temperature, operative temperature and wind speed), therefore, variations in DEE may be explained by body mass alone. The relative RMR of the African Black Oystercatcher was greater, their TME was approximately the same, their average daily metabolisable energy (ADME) was less, and they spent less time foraging (short periods of parental feeding) and more time inactive than three precocial species in the Western Cape. Therefore, the semi-precocial mode of development of African Black Oystercatcher chicks reduced energy costs from thermoregulation and activity, and they were able to grow relatively faster than precocial, self-feeding shorebird species in similar climatic conditions. The growth rate coefficient of African Black Oystercatcher chicks was smaller than that of Eurasian Oystercatcher, Haematopus ostralegus, chicks, which may be a consequence of differences in body size and latitudinal effects.     

Mechanisms promoting higher growth rate in arctic than in temperate shorebirds

We compared prefledging growth, energy expenditure, and time budgets in the arctic-breeding red knot (Calidris canutus) to those in temperate shorebirds, to investigate how arctic chicks achieve a high growth rate despite energetic difficulties associated with precocial development in a cold climate. Growth rate of knot chicks was very high compared to other, mainly temperate, shorebirds of their size, but strongly correlated with weather-induced and seasonal variation in availability of invertebrate prey. Red knot chicks sought less parental brooding and foraged more at the same mass and temperature than chicks of three temperate shorebird species studied in The Netherlands. Fast growth and high muscular activity in the cold tundra environment led to high energy expenditure, as measured using doubly labelled water: total metabolised energy over the 18-day prefledging period was 89% above an allometric prediction, and among the highest values reported for birds. A comparative simulation model based on our observations and data for temperate shorebird chicks showed that several factors combine to enable red knots to meet these high energy requirements: (1) the greater cold-hardiness of red knot chicks increases time available for foraging; (2) their fast growth further shortens the period in which chicks depend on brooding; and (3) the 24-h daylight increases potential foraging time, though knots apparently did not make full use of this. These mechanisms buffer the loss of foraging time due to increased need for brooding at arctic temperatures, but not enough to satisfy the high energy requirements without invoking (4) a higher foraging intake rate as an explanation. Since surface-active arthropods were not more abundant in our arctic study site than in a temperate grassland, this may be due to easier detection or capture of prey in the tundra. The model also suggested that the cold-hardiness of red knot chicks is critical in allowing them sufficient feeding time during the first week of life. Chicks hatched just after the peak of prey abundance in mid-July, but their food requirements were maximal at older ages, when arthropods were already declining. Snow cover early in the season prevented a better temporal match between chick energy requirements and food availability, and this may enforce selection for rapid growth.     

Catabolic capacity of the muscles of shorebird chicks: maturation of function in relation to body size

Newly hatched precocial chicks of arctic shorebirds are able to walk and regulate their body temperatures to a limited extent. Yet, they must also grow rapidly to achieve independence before the end of the short arctic growing season. A rapid growth rate may conflict with development of mature function, and because of the allometric scaling of thermal relationships, this trade-off might be resolved differently in large and small species. We assessed growth (mass) and functional maturity (catabolic enzyme activity) in leg and pectoral muscles of chicks aged 1-16 d and adults of two scolopacid shorebirds, the smaller dunlin (Calidris alpina: neonate mass 8 g, adult mass 50 g) and larger whimbrel (Numenius phaeopus; neonate mass 34 g, adult mass 380 g). Enzyme activity indicates maximum catabolic capacity, which is one aspect of the development of functional maturity of muscle. The growth rate-maturity hypothesis predicts that the development of catabolic capacity should be delayed in faster-growing muscle masses. Leg muscles of both species were a larger proportion of adult size at hatching and grew faster than pectoral muscles. Pectoral muscles grew more rapidly in the dunlin than in the whimbrel, whereas leg muscles grew more rapidly in the whimbrel. In both species and in both leg and pectoral muscles, enzyme activities generally increased with age, suggesting increasing functional maturity. Levels of citrate synthase activity were similar to those reported for other species, but l-3-hydroxyacyl-CoA-dehydrogenase and pyruvate kinase (PK) activities were comparatively high. Catabolic capacities of leg muscles were initially high compared to those of pectoral muscles, but with the exception of glycolytic (PK) capacities, these subsequently increased only modestly or even decreased as chicks grew. The earlier functional maturity of the more rapidly growing leg muscles, as well as the generally higher functional maturity in muscles of the more rapidly growing dunlin chicks, contradicts the growth rate-maturity function trade-off and suggests that birds have considerable latitude to modify this relationship. Whimbrel chicks, apparently, can rely on allometric scaling of power requirements for locomotion and the thermal inertia of their larger mass to reduce demands on their muscles, whereas dunlin chicks require muscles with higher metabolic capacity from an earlier age. Thus, larger and smaller species may adopt different strategies of growth and tissue maturation.     

Effects of water discharge on fledging time,growth, and survival of piping plovers on the Missouri River

River flow management and modification is a global issue, and its effects on river-dependent organisms are pervasive. Flow modification can directly affect avian species through mortality or habitat loss, but less is known about indirect and sublethal effects of flow modification on reproductive output in these species. Young birds are more vulnerable to predation between hatching and fledging than after flight is achieved, but tradeoffs must be made to balance growth and survival. Predation pressure appears to be a significant factor affecting the time to fledging in altricial birds, but less is known about this threat for precocial birds. Birds reaching fledging earlier should have greater rates of survival to migration because their predator escape repertoire includes flight at an earlier age. We evaluated the effect of varying outflows from the Gavins Point Dam on the growth, age at fledging, and survival of piping plover (Charadrius melodus) chicks on the Missouri River (2006–2009). The study was characterized by 2 relatively high flow years (2006 and 2009) and 2 relatively low flow years (2007 and 2008). We used success rate in recapturing chicks in capture–mark–recapture models as an index for fledging. We attempted to recapture all chicks (n = 1,099) by hand every 3–4 days throughout the season to acquire morphological measurements. Models indicated that as flows from the dam increased, age at fledging increased. We also found that increasing flows were associated with decreasing daily survival rates (β_flow = −2.401, 95% CI: −4.351 to −0.452). Flow was also negatively related to chick mass gain, but we found less evidence for an effect on wing-chord length. Increased flows covered wet-substrate foraging habitat, and likely affected plover reproductive output directly through chick survival and indirectly through decreased growth and increased fledging times. © The Wildlife Society, 2013     

Ontogenetic development of the ophthalmic rete in relation to brain cooling in chickens and pigeons

The brain cooling capacity of the altricial pigeon increases during posthatching growth at a higher rate than that of the precocial duck and chicken. To determine if this difference between the altricial and the precocial modes of development can be related to growth rates of the vascular heat exchanger involved in brain cooling (the ophthalmic rete), we performed a morphometric analysis of this structure during the post-hatching maturation of the three species. The number of vascular units in the rete did not change during development but differed significantly among species. The retia continued to grow in length and diameter in an exponential relation with body mass at similar rates in all species. The surface area of the retial arteries, which reflects the area available for countercurrent heat exchange, also increased exponentially with body mass, but without significant differences among the three species. However, the effectiveness of the rete in brain cooling, as indicated by the degree of brain cooling per unit of heat-exchange area in the rete, was higher in the altricial pigeon than in the precocial chicken and duck. It is concluded that the posthatching morphometric changes in the ophthalmic rete (rete ophthalmicum) are important for the development of brain cooling capacity, but cannot solely explain differences in brain cooling between growing altricial and precocial birds. These differences are most likely related to differences in the maturation of the central thermoregulatory control system and the peripheral effector mechanisms among the two groups of birds.     

Embryonic Calls as Care-soliciting Signals in Budgerigars,Melopsittacus undulatus

Embryonic calls occur 1–3 d before hatching among precocial and some altricial birds. In precocial species, calls may synchronize hatching among siblings or, in semi-precocial species, elicit parental attention to, and often thermoregulation of, the hatching egg. Much less is known about the functional significance of calls in fully altricial species. In this study, naturalistic observations and laboratory experiments were used to document factors affecting calling and the parental responses to calls in one altricial species, the budgerigar Melopsittacus undulatus. Budgerigar chicks hatch asynchronously and vocalize 24–48 h before hatching. Embryonic calling rates increase at higher egg temperatures, and also as embryos near hatching. Parents easily locate a calling egg in their clutch, even among a large brood of much older, vocalizing nestlings. Furthermore, they actively assist in the last stages of hatching by helping to break the shell along the crack in the egg. Both observational and experimental evidence suggests that embryonic vocalizations are distinctive signals that increase parental attention and care, and may stimulate hatching assistance to a calling egg.     

On the optimal growth of the alimentary tract in avian postembryonic development

It was shown in the mathematical model described elsewhere that when growth rate of the chicks is maximized and not constrained by the food availability, the optimal relationship between body mass and alimentary tract mass should conform to a single straight line, or two-, or three-segmented straight lines. Here, we present the data on growth of 11 bird species, and we test the model using the mass of intestines as an indicator of growth of the alimentary tract. The results support the predictions of the model for altricial species and contradict them for precocials. Since precocial species examined here were not food-limited, we suggest that the lack of optimal growth of their alimentary tract is inherent to their mode of development. This may account for their lower growth rate, as compared to altricials. The existence of the optimal growth of the alimentary tract in altricial nestlings suggests that under natural conditions the food is much more abundant than it is generally assumed.     

Mode of development and interspecific avian brood parasitism

Avian interspecific brood parasites differ considerably in theircommitment to parasitism; 87 species are obligate brood parasites,whereas 35 species are known to be facultative brood parasites.This variation is strongly related to mode of development. Obligateparasitism is found almost exclusively in altricial species,whereas facultative interspecific parasitism is predominantin precocial birds. We propose that the association betweenmode of development and form of parasitism reflects a fundamentaldifference between altricial and precocial birds in the relativebenefits of emancipation from parental care after laying. Weargue that altricial brood parasites obtain such a large increasein realized fecundity by avoiding the costs of parental carethat obligate parasitism is favored over facultative parasitism.In contrast, precocial brood parasites gain relatively littlein terms of increased fecundity via obligate parasitism, andmuch of this increase could potentially be gained by facultativeparasitism. Thus, obligate interspecific brood parasitism willnot be favored in precocial birds. Three factors influence thisdifference between altricial and precocial species: (1) altricialbirds have relatively more energy and nutrients with which tolay additional eggs, (2) altricial birds can produce more eggsfor the same amount of energy and nutrients, and (3) altricialbirds realize a greater relative gain in fecundity for eachadditional egg laid. We suggest further that facultative interspecificparasitism in birds may originate simply through a carry overof intraspecific parasitism; 29 of 33 facultative interspecificparasites also parasitize conspecifics. Facultative parasitismof other species would provide a greater range of potentialhost nests and could be maintained as an evolutionarily stableend point by the same mechanisms that maintain intraspecificbrood parasitism. [Behav Ecol 1991;2:309–318]     

GROWTH RATES OF BIRDS IN THE HUMID NEW WORLD TROPICS

The growth curves of 40 species of lowland neotropical birds were fitted by logistic equation. The birds were mostly from Panama, Trinidad and Surinam. The growth constants of the fitted equations (asymptote A and growth rate K) were compared within and among species, and with previously published data on temperate species. Growth parameters of tropical passerines are about as variable within species as they are within temperate species. In both cases, variation in A and K between broods is greater than it is within broods. Panamanian birds breed during the dry-wet transition and conditions for growth apparently improve as rainfall increases. Asymptotes of growth curves are higher, and mortality within broods lower, as the breeding season progresses. Asynchronous hatching and the reduction of brood-size by selective starvation of young is a prominent phenomenon during the early part of the breeding season. Several instances are reported, however, of young persisting in nests with inadequate feeding and greatly subnormal weights. Slowed development under conditions of poor nutrition may be adaptive in the tropics if periods of low food availability are short and allow the possibility of recovery from undernourishment. As a group, neotropical lowland passerines (30 species) grow 23% more slowly than a sample of 51 temperate passerines. Variation of growth rates among these tropical species is similar to variation among temperate species, and it is related to adult body-size the length of the nestling period. Young of tropical and temperate species attain similar asymptotes, relative to adult body-weight, by the end of the nestling period. Hypotheses are advanced which might explain the slower growth rate of tropical species, and tested to the extent available data permits. (1) Because brood-size can be changed only by adding or removing whole young, changes in growth rate could provide finer adjustment of the energy requirements of the young to the feeding capacity of the parents. This model predicts different means and variances for growth rate within groups of species with different clutch-sizes, predictions not supported by available data. (2) Growth rate is shown to increase the maximum energy requirement of a nestling only if K exceeds some value determined by the energy requirement of the young, growth rate should vary in proportion to the level of basal maintenance metabolism. In a small sample of tropical species, rates of basal metabolism were 25% lower than in a comparable sample of temperate species. These data therefore support the hypothesis, although the cause of the lower metabolic rate of the tropical nestlings is not known. (3) Daily periods of hypothermia could reduce the energy requirement of the young and at the same time reduce their growth rate; but observations of body temperatures of tropical nestlings are contrary to this hypothesis. (4) The short day-length of tropical climates reduces the time during which young can assimilate energy relative to their energy expenditures. This model predicts that tropical nestlings would have less productive energy available, (consistent with their reduced growth rates), but it also predicts that arctic birds should grow faster than temperate species, which is not confirmed by available data. (5) The low nitrogen content of fruits may cause the slow growth of a few strictly frugivorous species (Oilbird and Bearded Bellbird), but among other tropical species growth rate is not correlated with the estimated proportion of fruit in the diet.     

Chick growth rates in Charadriiformes: comparative analyses of breeding climate, development mode and parental care

The rate at which a young bird grows is highly diverse across taxa. We investigated the influences of ecological variables on growth rates of shorebirds, gulls and their allies (order Charadriiformes) using comparative analyses of 68 species. We investigated three hypotheses: (1) exposure to cold temperatures results in reduced growth rate due to the increased energy expenditure required for thermoregulation, (2) fast growth rates allow offspring to complete development in habitats with short periods of fair ecological conditions, and (3) parental feeding allows the offspring to grow faster than self-feeding offspring. Charadriiform species are suitable for testing these hypotheses, because they breed in diverse environmental conditions that include Arctic and Antarctic habitats, temperate zones and tropics, and in some species the offspring feed themselves (precocial) whereas in others they are fed by the parents (semi-precocial). First, we tested the influence of ambient temperature on growth rate and we found that species breeding in cold habitats had faster growth rates than species breeding in warm temperatures. The relationship between growth rate and ambient temperature was not significantly different between precocial and semi-precocial offspring. Second, we tested the influence of the length of the breeding season on growth rate, and we found that species with shorter breeding seasons had faster growth rates than species with longer breeding seasons. Finally, we show that precocial offspring grew slower than semi-precocial offspring, and this relationship remains significant when the influences of ambient temperature and breeding season length are statistically controlled. In conclusion, our work, using phylogenetically corrected models, confirms that ambient environment and developmental mode of young together influence the growth of Charadriiform chicks, and their effects are additive.     

Strategies of mass change in breeding birds

Birds experience strikingly different patterns of mass change during breeding. In some species with uniparental incubation, the incubating parents (mostly females) lose mass and attain their lowest point when the chicks hatch (Incubatory mass loss strategy = IML). In species with shared incubation, or with intense incubation feeding, the incubating parents maintain or increase in body mass without reducing attentiveness levels (Incubatory mass constancy = IMC). In other species, uniparental incubation with IMC is associated with reduced levels of attentiveness. The mobilization of fat stores or its preservation are involved in the two strategies. IML leads to mass increases after hatching of the young, while the opposite is true for IMC. The non-incubating sex in uniparental species does not experience significant mass changes during breeding. Fasting endurance, predation risk and mode of development are proposed as the main selective factors determining strategy. Latitude, climate, diet and food availability interact with the main factors. From a revision of the literature we can deduce that IML is mainly found among large birds with precocial development, while IMC is typical of smaller species with altricial development. Proportional mass losses are positively correlated with body mass in IML-species, as well as in precocial species, where body mass explains more than 70% of the variation in proportional mass change. Incubation periods increase with body mass in uniparental IMC-species, but not in IML-species. IML is thus associated to fast embryonic growth in large species. Successful raising of highly-dependent young in altricial and semialtricial species apparently depends on one of the parents retaining reserves until hatching. Subsequent mass losses may be necessary to maintain the brooding parent through a period when nestlings require heat, insulation and food. Patterns of mass change are not mere consequences of reproductive stress but the outcome of adaptive compromises between different selective factors and constitute an important aspect of the breeding biology and life history of birds.     

Benefits and costs of rapid growth in common murre chicks Uria aalge

While accelerated growth can be advantageous to nestling birds, there may be a tradeoff between rapid growth and resistance to food shortages. Common murres Uria aalg e are colonial seabirds that benefit from reproductive synchrony. Individuals that lay eggs late should benefit if they produce chicks capable of growing quickly and fledging synchronously with their neighbors. In this study, we controlled food provisioning of captive-hatched common murre chicks from a single subcolony and examined differences in growth between early-hatched individuals and their later-hatched neighbors. We assessed potential costs of rapid growth by comparing growth of chicks fed ad libitum with their growth under food restricted conditions. Chicks that hatched later were heavier, ate more and gained body mass more quickly than chicks that hatched earlier. Late-hatched chicks grew quickly enough to reach the same mass as their early-hatched neighbors in five days. However, chicks that grew more quickly under ad libitum food conditions grew more slowly when food was restricted. We conclude that murres that lay eggs late may synchronize their reproduction with early-laying neighbors by producing rapidly growing chicks. However, the ability to compensate for late hatching by growing quickly can be costly when food becomes limited.     

Catch-up growth in Japanese quail (Coturnix Japonica): relationships with food intake, metabolic rate and sex

The effects of early environmental conditions can profoundly affect individual development and adult phenotype. In birds, limiting resources can affect growth as nestlings, but also fitness and survival as adults. Following periods of food restriction, individuals may accelerate development, undergoing a period of rapid “catch-up” growth, in an attempt to reach the appropriate size at adulthood. Previous studies of altricial birds have shown that catch-up growth can have negative consequences in adulthood, although this has not been explored in species with different developmental strategies. Here, we investigated the effects of resource limitation and the subsequent period of catch-up growth, on the morphological and metabolic phenotype of adult Japanese quail (Coturnix japonica), a species with a precocial developmental strategy. Because males and females differ in adult body size, we also test whether food restriction had sex-specific effects. Birds that underwent food restriction early in development had muscles of similar size and functional maturity, but lower adult body mass than controls. There was no evidence of sex-specific sensitivity of food restriction on adult body mass; however, there was evidence for body size. Females fed ad lib were larger than males fed ad lib, while females subjected to food restriction were of similar size to males. Adults that had previously experienced food restriction did not have an elevated metabolic rate, suggesting that in contrast to altricial nestlings, there was no metabolic carry-over effect of catch-up growth into adulthood. While Japanese quail can undergo accelerated growth after re-feeding, timing of food restriction may be important to adult size, particularly in females. However, greater developmental flexibility compared to altricial birds may contribute to the lack of metabolic carryover effects at adulthood.     

Eggshell structure, mode of development and growth rate in birds

The shell of an egg contributes to successful embryogenesis in many ways, such as through protection, respiration and water exchange. The shell is also the major source of calcium for the development of high-calcium consuming organs, e.g. the skeleton, muscles and brain. Some studies show, moreover, that growth rate may play a fundamental role in the pattern of skeletal development in birds: the faster the growth the less ossified the skeleton is at hatching. We predicted, therefore, that slow (precocial) and fast (altricial) growing bird species should lay eggs encased in shells with different structures adapted to support different rates of calcium removal by developing embryos. We tested this prediction by comparing the fine structure of the inner eggshell surface (mammillary layer) from 36 bird species belonging to 18 orders ranging from Struthioniformes to Passeriformes. Using scanning electron microscopy, we compared the mammillary layer of both non-incubated eggs and eggs at the time of hatching, i.e., before and after embryonic development and the accompanying calcium removal. The results were consistent with the prediction, i.e., the number of mammillary tips per unit of surface area was associated with mode of development and growth rate. The number was higher, and calcium removal was also more extensive, in shells from precocial bird species than in shells from altricial bird species.     

Sibling Growth Patterns in Great tits: Does Increased Selection on Last-hatched Chicks Favour an Asynchronous Hatching Strategy?

In birds, competitive abilities of siblings in relation to their sex and the magnitude of hatching delay are still poorly understood. We compared the sex-specific growth of the last-hatched, competitively disadvantaged chicks with that of synchronously hatched chicks in two successive years. Sons exhibited higher growth rates than daughters in a year with delayed onset of breeding, and this sex-related difference was more pronounced among the asynchronously hatched chicks. Females apparently do not selectively allocate more resources to the last-laid eggs because neonatal body mass and the growth rate of asynchronously hatched chicks did not differ between years, despite the fact that in one of the years, asynchronous chicks hatched from replaced eggs taken randomly from other nests and not from eggs laid last by the incubating female. Among chicks that survived, moderately asynchronous siblings grew at a lower rate than synchronous ones, whereas no difference in mass gain was revealed between synchronous and strongly asynchronous siblings irrespective of their contrasting competitive abilities. We suggest that selection of the fittest asynchronous chicks may improve the overall quality of asynchronous broods, thus favouring the maintenance of asynchronous hatching strategies in variable environments.     

Adaptive hatching hypotheses do not explain asynchronous hatching in Brown-headed Parrots Poicephalus cryptoxanthus

At the core of the suite of adaptive hatching hypotheses advanced to explain asynchronous hatching in birds is the assumption that if food is not limited then all the hatchlings will develop normally to adulthood. In this study Brown-headed Parrot Poicephalus cryptoxanthus chicks were hand fed and weighed on a daily basis. Their respective masses were compared using a reparamatised Richards growth model. The results show that although the second and third hatched chicks initially grow faster than the first, the final mass attained by the third chick is significantly less than either of its siblings. Adaptive hatching hypotheses do not explain this result. We propose that the third chick is opportunistic and that is has evolved as insurance against prior chicks surviving because of interspecific pressure on the breeding biology of the species.     

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.     

The effect of egg size on post-hatching development in the Razorbill: an experimental study

A recent experimental study reported that Thick-billed Murre Uria lomvia chicks that hatched from large eggs grew their wing feathers more quickly than did small-egg chicks. There is little evidence of this (or any other) egg-size effect on post-hatching development in other birds. Thick-billed Murres are marine birds of the family Alcidae that employ the unique "intermediate" developmental strategy: chicks go to sea after 15-30 days at the nest site, at <30% of adult mass, accompanied by their male parent. Rapid feather growth during the brief nestling period is critical to enable chicks to make the transition from life at the nest site to life at sea quickly and safely. At the Gannet Islands, Labrador, Canada, in 1996 and 1997, I tested whether egg size has the same effect on wing-feather growth in the Razorbill Alca torda , another of the intermediate auks. To control for underlying correlations between egg size and other parental attributes, eggs were switched randomly among pairs. As in other birds, egg size strongly predicted hatchling mass, and to less extent hatchling size (tarsus length), but had no effect on the rate at which nestlings gained mass. However, egg size had the same effect on wing growth in Razorbills as in Thick-billed Murres: the wings of large-egg chicks began rapid, linear growth sooner, indicating that early development of wing feathers was enhanced in large-egg chicks. Differences in wing length established in this manner persisted through the nestling period. Egg-size effects on feather growth have not been detected in experimental studies on other birds, suggesting that effects of the magnitude seen in Razorbills and Thick-billed Murres might reflect evolutionary priorities in the post-hatching development of intermediate auks.