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...
Birds with uniparental incubation may face a time allocation problem between incubation and feeding. Eggs need regular warming to hatch successfully, but the parent must leave the nest to feed and safeguard its own survival. Time allocation during incubation is likely to depend on factors influencing egg cooling rates, parental energy requirements and feeding intake rate. How this allocation problem is resolved was subject of this study on arctic‐breeding shorebirds. We compared incubation rhythms between four uniparental shorebird species differing in size and expected to find both species differences and weather effects on the organisation of incubation. Attentive behaviour and responses to variation in weather showed a remarkable consistency across species. All species alternated feeding bouts (recesses) with brooding bouts throughout the day. Recesses were concentrated in the warmer parts of the day, while recess duration showed little diurnal variation. Despite continuous daylight, a pronounced day‐night rhythmicity was apparent. The four species in this study spent a similar proportion (13–19%) of the time off their nest. After correction for weather effects, the number of recesses was largest in the smallest species, while recess duration was longest in the largest species. Total recess time per day increased on cold days through an increase of mean recess length, while the number of recesses decreased. Comparing our observations to predictions derived from criteria that birds might use to organise their attentive behaviour, showed that the limits are set by parental requirements, while the energy stores of adults provide some leeway for short‐term adjustments to environmental variability. If breeding birds trade off feeding time against incubation time, energy stores are expected to be influenced by weather. We expected uniparental species to be more likely to show weather effects on condition than biparentals, as in the latter ‘off duty’ time is much larger and independent of weather. This prediction was tested by comparing energy stores in two uniparental species and a biparental congener. While body mass of uniparental incubators decreased after a period with low temperatures, body mass of the biparental species did not.
We modeled the relationship between egg flotation and age of a developing embryo for 24 species of shorebirds. For 21 species, we used regression analyses to estimate hatching date by modeling egg angle and float height, measured as continuous variables, against embryo age. For eggs early in incubation, we used linear regression analyses to predict hatching date from logit-transformed egg angles only. For late incubation, we used multiple regression analyses to predict hatching date from both egg angles and float heights. In 30 of 36 cases, these equations estimated hatching date to within four days of the true hatching date for each species. After controlling for incubation duration and egg size, flotation patterns did not differ between shorebirds grouped by mass (≥100 g or <100 g) or taxonomy (Scolopacidae versus Charadriidae). Flotation progressed more rapidly in species in which both adults incubate the clutch versus species in which only one adult incubates the clutch, although this did not affect prediction accuracy. We also pooled all continuous data and created a generalized regression equation that can be applied to all shorebird species. For the remaining three species, we estimated hatching date using five float categories. Estimates of hatching date using categorical data were, overall, less accurate than those generated using continuous data (by 3%–5% of a given incubation period). Our equations were less accurate than results reported in similar studies; data collected by multiple observers and at multiple sites, as well as low sample sizes for some species, likely increased measurement error. To minimize flotation method prediction error, we recommend sampling in early incubation, collecting both egg angle and float height data in late incubation, and developing site- and species-specific regression models where possible.
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