Maarten Brugge

ENDOGENOUS CIRCANNUAL RHYTHMICITY IN BODY MASS, MOLT, AND PLUMAGE OF GREAT KNOTS (CALIDRIS TENUIROSTRIS)

Abstract Four Great Knots (Calidris tenuirostris) were kept for six years in a constant-temperature indoor aviary. For two of those six years, they were kept under photoperiodic conditions that mimicked natural changes in daylength for wild birds, followed by four years under a constant photoperiod (light:dark cycle 12:12 h). Under cyclical “natural” photoperiods, three of the four birds maintained cycles of body mass and contour and flight-feather molt somewhat comparable to that of free-living birds, though the multiple mass peaks characteristic of northward migration were replaced by a single period of high body mass; the mass peaks for southward migration appeared to be absent. Contour-feather molts between nonbreeding and breeding plumages were delayed, and the period of wing molt was longer than in free-living birds. Under constant photoperiods, clear circannual phenotype cycles were maintained. The length of the period with elevated body mass tripled but was partly compensated by a shortening of the duration of wing molt (which never coincided with high body masses). Nevertheless, total cycle lengths were >13 months. Perhaps most interestingly, under constant photoperiod, there was evidence that two components of what is normally considered an integrated phenotypic event, the prebasic molts of contour and wing feathers, were desynchronized. This suggests that the underlying organizational structure of traits is modular to some extent. Such modularity would increase the flexibility and versatility of the cyclic phenotype in evolutionary contexts. Ritmo del Ciclo Anual Endógeno del Peso Corporal, Muda y Plumaje en Calidris tenuirostris

Sex-specific winter distribution in a sexually dimorphic shorebird is explained by resource partitioning

Sexual size dimorphism (SSD) implies correlated differences in energetic requirements and feeding opportunities, such that sexes will face different trade-offs in habitat selection. In seasonal migrants, this could result in a differential spatial distribution across the wintering range. To identify the ecological causes of sexual spatial segregation, we studied a sexually dimorphic shorebird, the bar-tailed godwit Limosa lapponica, in which females have a larger body and a longer bill than males. With respect to the trade-offs that these migratory shorebirds experience in their choice of wintering area, northern and colder wintering sites have the benefit of being closer to the Arctic breeding grounds. According to Bergmanns rule, the larger females should incur lower energetic costs per unit of body mass over males, helping them to winter in the cold. However, as the sexes have rather different bill lengths, differences in sex-specific wintering sites could also be due to the vertical distribution of their buried prey, that is, resource partitioning. Here, in a comparison between six main intertidal wintering areas across the entire winter range of the lapponica subspecies in northwest Europe, we show that the percentage of females between sites was not correlated with the cost of wintering, but was positively correlated with the biomass in the bottom layer and negatively with the biomass in the top layer. We conclude that resource partitioning, rather than relative expenditure advantages, best explains the differential spatial distribution of male and female bar-tailed godwits across northwest Europe.

Seasonal Time Keeping in a Long-Distance Migrating Shorebird

Because of the complications in achieving the necessary long-term observations and experiments, the nature and adaptive value of seasonal time-keeping mechanisms in long-lived organisms remain understudied. Here we present the results of a 20-year-long study of the repeated seasonal changes in body mass, plumage state, and primary molt of 45 captive red knots Calidris canutus islandica, a High Arctic breeding shorebird that spends the nonbreeding season in temperate coastal areas. Birds kept outdoors and experiencing the natural photoperiod of the nonbreeding area maintained sequences of life-cycle stages, roughly following the timing in nature. For 6 to 8 years, 14 of these birds were exposed to unvarying ambient temperature (12 °C) and photoperiodic conditions (12:12 LD). Under these conditions, for at least 5 years they expressed free-running circannual cycles of body mass, plumage state, and wing molt. The circannual cycles of the free-running traits gradually became longer than 12 months, but at different rates. The prebreeding events (onset and offset of prealternate molt and the onset of spring body mass increase) occurred at the same time of the year as in the wild population for 1 or several cycles. As a result, after 4 years in 12:12 LD, the circannual cycles of prealternate plumage state had drifted less than the cycles of prebasic plumage state and wing molt. As the onset of body mass gain drifted less than the offset, the period of high body mass became longer under unvarying conditions. We see these differences between the prebreeding and postbreeding life-cycle stages as evidence for adaptive seasonal time keeping in red knots: the life-cycle stages linked to the initiation of reproduction rely mostly on endogenous oscillators, whereas the later stages rather respond to environmental conditions. Postbreeding stages are also prone to carryover effects from the earlier stages.

Understanding spatial distributions: negative density-dependence in prey causes predators to trade-off prey quantity with quality

Negative density-dependence is generally studied within a single trophic level, thereby neglecting its effect on higher trophic levels. The ‘functional response’ couples a predators intake rate to prey density. Most widespread is a type II functional response, where intake rate increases asymptotically with prey density; this predicts the highest predator densities at the highest prey densities. In one of the most stringent tests of this generality to date, we measured density and quality of bivalve prey (edible cockles Cerastoderma edule) across 50 km² of mudflat, and simultaneously, with a novel time-of-arrival methodology, tracked their avian predators (red knots Calidris canutus). Because of negative density-dependence in the individual quality of cockles, the predicted energy intake rates of red knots declined at high prey densities (a type IV, rather than a type II functional response). Resource-selection modelling revealed that red knots indeed selected areas of intermediate cockle densities where energy intake rates were maximized given their phenotype-specific digestive constraints (as indicated by gizzard mass). Because negative density-dependence is common, we question the current consensus and suggest that predators commonly maximize their energy intake rates at intermediate prey densities. Prey density alone may thus poorly predict intake rates, carrying capacity and spatial distributions of predators.