Jeroen Reneerkens

Baseline and Stress‐Induced Levels of Corticosterone during Different Life Cycle Substages in a Shorebird on the High Arctic Breeding Grounds

After a migratory flight of several thousand kilometers to their high arctic breeding grounds, red knots (Calidris canutus islandica, Scolopacidae) showed high baseline concentrations of plasma corticosterone (58 ng/mL). Such high baseline corticosterone levels may be conditional for the right behavioral and metabolic adjustments to environmental and social stresses that shorebirds experience on arrival in an unpredictable tundra breeding environment. Despite the high baseline levels of corticosterone, red knots still showed a marked stress response during the postarrival period, with corticosterone concentrations increasing significantly during a 60‐min period of confinement. Baseline levels of corticosterone declined as the breeding season progressed. Red knots with brood patches, that is, birds that had completed egg laying and commenced incubation, had a reduced adrenocortical response to the stress of confinement compared with red knots with no, or with half‐developed, brood patches. This is consistent with the idea that birds breeding in extreme environments with short breeding seasons may exhibit a decreased adrenocortical response to stressful events to prevent high corticosterone concentrations from inducing interruptions of reproductive behavior.

Seasonally Changing Preen-Wax Composition: Red Knots' (Calidris Canutus) Flexible Defense Against Feather-Degrading Bacteria

Abstract During incubation, ground-breeding sandpipers such as Red Knots (Calidris canutus) create a warm, humid microclimate in the nest, conditions that favor the growth of feather-degrading bacteria present in their plumage. Just before incubation, the composition of waxes secreted by the uropygial gland of Red Knots and other sandpipers changes quickly and completely from a mixture of only monoesters to a mixture of only diesters. We hypothesized that the change in composition of the preen wax helps protect the plumage against feather-degrading bacteria. We tested the hypothesis by studying growth of the feather-degrading bacterium Bacillus licheniformis, which we found in the plumage of Red Knots. The removal of preen waxes from feathers resulted in faster bacterial degradation, confirming earlier findings that preen wax inhibits growth of feather-degrading bacteria. However, the degradation rate of feathers with preen wax based on diesters did not differ from that of feathers with preen wax based on monoesters. We suggest that preen waxes protect feathers by forming a physical barrier to microbes rather than through chemical properties of the waxes.

Exposing the structure of an Arctic food web.

How food webs are structured has major implications for their stability and dynamics. While poorly studied to date, arctic food webs are commonly assumed to be simple in structure, with few links per species. If this is the case, then different parts of the web may be weakly connected to each other, with populations and species united by only a low number of links. We provide the first highly resolved description of trophic link structure for a large part of a high-arctic food web. For this purpose, we apply a combination of recent techniques to describing the links between three predator guilds (insectivorous birds, spiders, and lepidopteran parasitoids) and their two dominant prey orders (Diptera and Lepidoptera). The resultant web shows a dense link structure and no compartmentalization or modularity across the three predator guilds. Thus, both individual predators and predator guilds tap heavily into the prey community of each other, offering versatile scope for indirect interactions across different parts of the web. The current description of a first but single arctic web may serve as a benchmark toward which to gauge future webs resolved by similar techniques. Targeting an unusual breadth of predator guilds, and relying on techniques with a high resolution, it suggests that species in this web are closely connected. Thus, our findings call for similar explorations of link structure across multiple guilds in both arctic and other webs. From an applied perspective, our description of an arctic web suggests new avenues for understanding how arctic food webs are built and function and of how they respond to current climate change. It suggests that to comprehend the community-level consequences of rapid arctic warming, we should turn from analyses of populations, population pairs, and isolated predator–prey interactions to considering the full set of interacting species.

Unexpected diversity in socially synchronized rhythms of shorebirds

The behavioural rhythms of organisms are thought to be under strong selection, influenced by the rhythmicity of the environment. Such behavioural rhythms are well studied in isolated individuals under laboratory conditions, but free-living individuals have to temporally synchronize their activities with those of others, including potential mates, competitors, prey and predators. Individuals can temporally segregate their daily activities (for example, prey avoiding predators, subordinates avoiding dominants) or synchronize their activities (for example, group foraging, communal defence, pairs reproducing or caring for offspring). The behavioural rhythms that emerge from such social synchronization and the underlying evolutionary and ecological drivers that shape them remain poorly understood. Here we investigate these rhythms in the context of biparental care, a particularly sensitive phase of social synchronization where pair members potentially compromise their individual rhythms. Using data from 729 nests of 91 populations of 32 biparentally incubating shorebird species, where parents synchronize to achieve continuous coverage of developing eggs, we report remarkable within- and between-species diversity in incubation rhythms. Between species, the median length of one parent’s incubation bout varied from 1–19 h, whereas period length—the time in which a parent’s probability to incubate cycles once between its highest and lowest value—varied from 6–43 h. The length of incubation bouts was unrelated to variables reflecting energetic demands, but species relying on crypsis (the ability to avoid detection by other animals) had longer incubation bouts than those that are readily visible or who actively protect their nest against predators. Rhythms entrainable to the 24-h light–dark cycle were less prevalent at high latitudes and absent in 18 species. Our results indicate that even under similar environmental conditions and despite 24-h environmental cues, social synchronization can generate far more diverse behavioural rhythms than expected from studies of individuals in captivity. The risk of predation, not the risk of starvation, may be a key factor underlying the diversity in these rhythms.

Trans-Equatorial Migration Routes, Staging Sites and Wintering Areas of a High-Arctic Avian Predator: The Long-tailed Skua ( Stercorarius longicaudus )

The Long-tailed Skua, a small (<300 g) Arctic-breeding predator and seabird, is a functionally very important component of the Arctic vertebrate communities in summer, but little is known about its migration and winter distribution. We used light-level geolocators to track the annual movements of eight adult birds breeding in north-east Greenland (n = 3) and Svalbard (n = 5). All birds wintered in the Southern Hemisphere (mean arrival-departure dates on wintering grounds: 24 October-21 March): five along the south-west coast of Africa (0–40°S, 0–15°E), in the productive Benguela upwelling, and three further south (30–40°S, 0–50°E), in an area extending into the south-west Indian Ocean. Different migratory routes and rates of travel were documented during post-breeding (345 km d−1 in late August-early September) and spring migrations (235 km d−1 in late April) when most birds used a more westerly flyway. Among the different staging areas, a large region off the Grand Banks of Newfoundland appears to be the most important. It was used in autumn by all but one of the tracked birds (from a few days to three weeks) and in spring by five out of eight birds (from one to more than six weeks). Two other staging sites, off the Iberian coast and near the Azores, were used by two birds in spring for five to six weeks. Over one year, individuals travelled between 43,900 and 54,200 km (36,600–45,700 when excluding staging periods) and went as far as 10,500–13,700 km (mean 12,800 km) from their breeding sites. This study has revealed important marine areas in both the south and north Atlantic Ocean. Sustainable management of these ocean basins will benefit Long-tailed Skuas as well as other trans-equatorial migrants from the Arctic.

Do Uniparental Sanderlings Calidris alba Increase Egg Heat Input to Compensate for Low Nest Attentiveness

Birds breeding in cold environments regularly have to interrupt incubation to forage, causing a trade-off between two mutually exclusive behaviours. Earlier studies showed that uniparental Arctic sandpipers overall spend less time incubating their eggs than biparental species, but interspecific differences in size and ecology were potential confounding factors. This study reports on a within-species comparison of breeding schedules and metal egg temperatures in uni- and biparental sanderlings (Calidris alba) in Northeast Greenland in relation to ambient temperature. We recorded incubation schedules with nest temperature loggers in 34 sanderling clutches (13 uniparentals, 21 biparentals). The temperature of a metal egg placed within the clutch of 17 incubating birds (6 uniparentals, 9 biparentals) was measured as an indicator of the heat put into eggs. Recess frequency, recess duration and total recess time were higher in uniparentals than in biparentals and positively correlated with ambient temperatures in uniparentals only. Uniparental sanderlings maintained significantly higher metal egg temperatures during incubation than biparentals (1.4°C difference on average). Our results suggest that uniparental sanderlings compensate for the lower nest attendance, which may prolong the duration of the incubation period and negatively affect the condition of the hatchlings, by maintaining a higher heat flux into the eggs.

Breeding on the extreme edge: Modulation of the adrenocortical response to acute stress in two High Arctic passerines

ABSTRACT Arctic weather in spring is unpredictable and can also be extreme, so Arctic‐breeding birds must be flexible in their breeding to deal with such variability. Unpredictability in weather conditions will only intensify with climate change and this in turn could affect reproductive capability of migratory birds. Adjustments to coping strategies are therefore crucial, so here we examined the plasticity of the adrenocorticotropic stress response in two Arctic songbird species—the snow bunting (Plectrophenax nivalis) and Lapland longspur (Calcarius lapponicus)—breeding in northwest Greenland. Across the breeding season, the stress response was strongest at arrival and least robust during molt in male snow buntings. Snow bunting females had higher baseline but similar stress‐induced corticosterone levels compared to males. Modification of the stress response was not due to adrenal insensitivity, but likely regulated at the anterior pituitary gland. Compared to independent nestlings and adult snow buntings, parental‐dependent chicks had a more robust stress response. For Lapland longspurs, baseline corticosterone was highest at arrival in both male and females, and arriving males displayed a higher stress response compared to arriving females. Comparison of male corticosterone profiles collected at arrival in Greenland (76°N) and Alaska (67–71°N;) reveal that both species have higher stress responses at the more northern location. Flexibility in the stress response may be typical for birds nesting at the leading edges of their range and this ability will become more relevant as global climate change results in major shifts of breeding habitat and phenology for migratory birds. J. Exp. Zool. 323A: 266–275, 2015.

Effects of food abundance and early clutch predation on reproductive timing in a high Arctic shorebird exposed to advancements in arthropod abundance

Summary Climate change may influence the phenology of organisms unequally across trophic levels and thus lead to phenological mismatches between predators and prey. In cases where prey availability peaks before reproducing predators reach maximal prey demand, any negative fitness consequences would selectively favor resynchronization by earlier starts of the reproductive activities of the predators. At a study site in northeast Greenland, over a period of 17 years, the median emergence of the invertebrate prey of Sanderling Calidris alba advanced with 1.27 days per year. Yet, over the same period Sanderling did not advance hatching date. Thus, Sanderlings increasingly hatched after their prey was maximally abundant. Surprisingly, the phenological mismatches did not affect chick growth, but the interaction of the annual width and height of the peak in food abundance did. Chicks grew especially better in years when the food peak was broad. Sanderling clutches were most likely to be depredated early in the season, which should delay reproduction. We propose that high early clutch predation may favor a later reproductive timing. Additionally, our data suggest that in most years food was still abundant after the median date of emergence, which may explain why Sanderlings did not advance breeding along with the advances in arthropod phenology.

Egg hatchability in high Arctic breeding wader species Charadriiformes is not affected by determining incubation stage using the egg flotation technique

Capsule By following the fate of both floated and non-floated eggs from individual clutches of four Arctic wader (shorebird) species, we show that the use of the flotation method for determining incubation stage of eggs (which involves both the submersion and handling of eggs) does not negatively affect hatchability.

Within-population variation in mating system and parental care patterns in the Sanderling (Calidris alba) in northeast Greenland

ABSTRACT Sandpipers and allies (Scolopacidae) show an astounding diversity in mating and parental care strategies. Comparative studies have tried to interpret this variation in terms of phylogenetic constraints and ecological shaping factors. In such analyses, mating and parental care systems are necessarily discretely classified at the species level. The few available descriptive studies on breeding strategies of the Sanderling (Calidris alba) came to variable conclusions, which, in turn, were inconsistently used in these comparative studies. We provide empirical data on mating and parental care patterns in Sanderlings studied during six summers in northeast Greenland. In 135 clutches, we determined parental care from incubation profiles using temperature loggers and confirmed that both uniparental incubation by both sexes and biparental incubation (45 and 90 clutches, respectively) occurred. We used microsatellite-based parentage analyses to describe the degree of extrapair mating. In 48 completely assayed families, we found 6 cases of polygamy (4 cases of polyandry, 2 cases of polygyny) that involved both uniparental and biparental clutches. This implies substantial variation in the patterns of mating and parental care, defying categorical assignments even at the local level. We conclude that the classification of mating strategy and parental care pattern for the Sanderling has been rather coarse, and that comparative analyses have not taken the observed intrapopulation variability into account. Because sandpipers show such variable reproductive behavior, between and within species, more detailed descriptive studies using parentage analyses are required to revisit previous statements about the intensity of sexual selection, including sexual size dimorphism, in shorebirds. In view of the great variability, methods of comparison will need elaboration too.