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.

ABSTRACT. Of all climatic zones on earth, Arctic areas have experienced the greatest climate change in recent decades. Predicted changes, including a continuing rise in temperature and precipitation and a reduction in snow cover, are expected to have a large impact on Arctic life. Large numbers of birds breed on the Arctic tundra, and many of these, such as shorebirds and passerines, feed on arthropods. Their chicks depend on a short insect population outburst characteristic of Arctic areas. To predict the consequences of climate change for reproduction in these birds, insight into arthropod phenology is essential. We investigated weather-related and seasonal patterns in abundance of surface-active arthropods during four years in the tundra of NW Taimyr, Siberia. The resulting statistical models were used to hindcast arthropod abundance on the basis of a 33-year weather dataset collected in the same area. Daily insect abundance was correlated closely with date, temperature, and, in some years, with wind and precipitation. An additional correlation with the number of degree-days accumulated after 1 June suggests that the pool of potential arthropod recruits is depleted in the course of the summer. The amplitude of short-term, weather-induced variation was as large as that of the seasonal variation. The hindcasted dates of peak arthropod abundance advanced during the study period, occurring seven days earlier in 2003 than in 1973. The timing of the period during which birds have a reasonable probability of finding enough food to grow has changed as well, with the highest probabilities now occurring at earlier dates. At the same time, the overall length of the period with probabilities of finding enough food has remained unchanged. The result is an advancement of the optimal breeding date for breeding birds. To take advantage of the new optimal breeding time, Arctic shorebirds and passerines must advance the start of breeding, and this change could affect the entire migratory schedule. Because our analyses are based on a single site, we cannot conclude that this is a general pattern for the entire Arctic. To investigate the generality of this pattern, our approach should be applied at other sites too.Key words: climate change, arthropods, Siberia, timing of breeding, phenology, Arctic birds RÉSUMÉ. De toutes les zones climatiques de la Terre, ce sont les régions de l'Arctique qui ont enregistré le plus grand changement climatique au cours des dernières décennies. Les changements prévus, qui comprennent notamment l'augmentation continue des températures et des précipitations de même que la diminution de la couverture de neige, devraient avoir de grandes incidences sur la vie de l'Arctique. De grandes quantités d'oiseaux se reproduisent sur la toundra de l'Arctique, et beaucoup d'entre eux, tels que les oiseaux de rivage et les passériformes, se nourrissent d'arthropodes. Leurs oisillons dépendent d'une courte affluence de population d'insectes, ce qui est caractéristique des régions arctiques. Afin de prévoir les co...

The doubly labeled water (DLW) method was validated against respiration gas analysis in growing precocial chicks of the black-tailed godwit (Limosa limosa) and the northern lapwing (Vanellus vanellus). To calculate the rate of CO2 production from DLW measurements, Lifson and McClintock's equations (6) and (35) were employed, as well as Speakman's equation (7.17) (all single-pool models). The average errors obtained with the first two equations (+7.2% and -11.6%, respectively) differed significantly from zero but not the error obtained with Speakman's equation (average: -2.9%). The latter error could be reduced by taking a fractional evaporative water loss of 0.13, instead of the value of 0. 25 recommended by Speakman. Application of different two-pool models resulted in relative errors of the DLW method of -15.9% or more. After employing the single-pool model with a fractional evaporative water loss value of 0.13, it was found that there was no relationship between the relative growth rate of the chick and the relative error of the DLW method. Recalculation of previously published results on Arctic tern (Sterna paradisaea) chicks revealed that the fit of the validation experiment could be considerably improved by employing a single-pool model and assuming a fractional evaporative water loss of 0.20 instead of the value of 0.50 taken originally. After employing the value of 0.20, it was found that there was no relationship between the relative growth rate of the chick and the relative error of the DLW method. This suggests that isotope incorporation into new body substances does not cause a detectable error. Thus, the DLW method seems to be applicable in young birds growing as fast as 20% d-1, after making adjustments for the fractional evaporative water loss. We recommend Speakman's equation (7.17) for general use in growing birds when evaporation is unknown.