Marcel Klaassen

Hampered Foraging and Migratory Performance in Swans Infected with Low-Pathogenic Avian Influenza A Virus

It is increasingly acknowledged that migratory birds, notably waterfowl, play a critical role in the maintenance and spread of influenza A viruses. In order to elucidate the epidemiology of influenza A viruses in their natural hosts, a better understanding of the pathological effects in these hosts is required. Here we report on the feeding and migratory performance of wild migratory Bewicks swans (Cygnus columbianus bewickii Yarrell) naturally infected with low-pathogenic avian influenza (LPAI) A viruses of subtypes H6N2 and H6N8. Using information on geolocation data collected from Global Positioning Systems fitted to neck-collars, we show that infected swans experienced delayed migration, leaving their wintering site more than a month after uninfected animals. This was correlated with infected birds travelling shorter distances and fuelling and feeding at reduced rates. The data suggest that LPAI virus infections in wild migratory birds may have higher clinical and ecological impacts than previously recognised.

Long-distance endozoochorous dispersal of submerged macrophyte seeds by migratory waterbirds in northern Europe—a critical review of possibilities and limitations

We review whether migratory Anatidae, i.e., swans, geese and ducks, could be acting as vectors for dispersal of Zostera, Ruppia and Potamogeton propagules by endozoochory (carrying seeds in their guts). We list six prerequisites that must all be fulfilled, if successful dispersal should occur. Several Anatidae species feed on these macrophytes, and undertake rapid long-distance movements, making dispersal possible. We identify four problems, which in combination leads us to conclude that long-distance dispersal events are likely to be rare. (i) Most long-distance movements are out of phase with the reproductive efforts of the plants, and if birds arrive at sites when plants still bear seeds, they are likely to depart well after seed stocks have been depleted. (ii) Seed transport by birds will usually be uni-directional, from north to south on autumn migrations. (iii) Most of the gut contents of migratory birds are likely to have been discarded within 300 km of departure. (iv) In many cases, birds will arrive in habitats seriously different from those they departed, i.e., any seeds carried along will have low chances of surviving in their new site. We suggest that northbound dispersal by endozoochory can only occur during spring if waterbirds feed on seeds that have not been depleted and remained frozen down or buried in sediments, or during moult- or post-moult migrations. Moult migration takes place in summer in phase with the reproductive efforts of the plants. Also epizoochorous dispersal (external attachment) is subject to restrictions i, ii and iv.

Moult and basal metabolic costs in males of two subspecies of stonechats: the European Saxicola torquata rubicula and the East African S. t. axillaris

The circannual patterns in resting metabolic rate (RMR) of males of two subspecies of stonechats, the European Saxicola torquata rubicula and the East African S. t. axillaris, are compared. As the birds from the two subspecies were raised and kept under comparable laboratory conditions, differences in metabolic rate between the two subspecies had to be genetically determined. RMR peaked during moult in both subspecies. During the rest of the year RMR was fairly constant in both subspecies and assumed to reflect basal metabolic rate (BMR). African stonechats had a 22% lower mass specific BMR than European stonechats, which is thought to reflect a genetical physiological adaptation to the differences in environmental circumstances they experience in the field. A low BMR makes an animal more susceptible to cold. Hence, the relatively high plumage mass in the African compared to the European stonechat may be functionally linked to its relatively low BMR. Moult costs, calculated from the plumage masses and the differences in RMR inside and outside the moult period, tended to be higher in the European compared to the African stonechats. These data and an interspecific comparison of moult costs over various species of birds support the earlier notion by Lindström et al. (1993) that moult costs are more closely linked with BMR than with body mass or rate of moult. The relation between moult costs and BMR and the fact that the efficiency of moult is extremely low (3.8 and 6.4% for European and African stonechats, respectively) suggest that the maintenance of specific tissues necessary for moult is a large cost factor. Alternatively, impaired insulation during moult may necessitate an increased metabolic capacity which may be associated with an increased RMR.

Surveillance of wild birds for avian influenza virus

TOC Summary: A targeted, hypothesis-based approach and local surveys over broad geographic areas are needed.

Ecophysiology of avian migration in the face of current global hazards

Long-distance migratory birds are often considered extreme athletes, possessing a range of traits that approach the physiological limits of vertebrate design. In addition, their movements must be carefully timed to ensure that they obtain resources of sufficient quantity and quality to satisfy their high-energy needs. Migratory birds may therefore be particularly vulnerable to global change processes that are projected to alter the quality and quantity of resource availability. Because long-distance flight requires high and sustained aerobic capacity, even minor decreases in vitality can have large negative consequences for migrants. In the light of this, we assess how current global change processes may affect the ability of birds to meet the physiological demands of migration, and suggest areas where avian physiologists may help to identify potential hazards. Predicting the consequences of global change scenarios on migrant species requires (i) reconciliation of empirical and theoretical studies of avian flight physiology; (ii) an understanding of the effects of food quality, toxicants and disease on migrant performance; and (iii) mechanistic models that integrate abiotic and biotic factors to predict migratory behaviour. Critically, a multi-dimensional concept of vitality would greatly facilitate evaluation of the impact of various global change processes on the population dynamics of migratory birds.

Effects of snow cover on the timing and success of reproduction in high-Arctic pink-footed geese Anser brachyrhynchus

During four breeding seasons, 2003–2006, we studied the relationship between snow cover and nesting performance in pink-footed geese (Anser brachyrhynchus) in a key breeding site on Svalbard. Snow cover in late May, i.e., at the time of egg laying of geese, was derived from MODIS satellite images. Snow cover had a profound cascading effect on reproductive output via the number of nesting pairs and timing of nesting, which affected nest success, while there was only a tendency for a negative effect on clutch size. Hence, we estimated a five-fold difference in the number of young produced (to post-hatching) between years with little snow and years with high snow cover. The results from the study area correlated with whole-population productivity estimates recorded in autumn. Thus, snow cover derived from MODIS satellite images appears to provide a useful indicator of the breeding conditions in the Arctic.

Cues and the optimal timing of activities under environmental changes

Organisms time activities by using environmental cues to forecast the future availability of important resources. Presently, there is limited understanding of the relationships between cues and optimal timing, and especially about how this relationship will be affected by environmental changes. We develop a general model to explore the relation between a cue and the optimal timing of an important life history activity. The model quantifies the fitness loss for organisms failing to time behaviours optimally. We decompose the immediate change in fitness resulting from environmental changes into a component that is due to changes in the predictive power of the cue and a component that derives from the mismatch of the old response to the cue to the new environmental conditions. Our results show that consequences may range from negative, neutral to positive and are highly dependent on how cue and optimal timing and their relation are specifically affected by environmental changes.

The consequences of climate‐driven stop‐over sites changes on migration schedules and fitness of Arctic geese

1. How climatic changes affect migratory birds remains difficult to predict because birds use multiple sites in a highly interdependent manner. A better understanding of how conditions along the flyway affect migration and ultimately fitness is of paramount interest. 2. Therefore, we developed a stochastic dynamic model to generate spatially and temporally explicit predictions of stop-over site use. For each site, we varied energy expenditure, onset of spring, intake rate and day-to-day stochasticity independently. We parameterized the model for the migration of pink-footed goose Anser brachyrhynchus from its wintering grounds in Western Europe to its breeding grounds on Arctic Svalbard. 3. Model results suggested that the birds follow a risk-averse strategy by avoiding sites with comparatively high energy expenditure or stochasticity levels in favour of sites with highly predictable food supply and low expenditure. Furthermore, the onset of spring on the stop-over sites had the most pronounced effect on staging times while intake rates had surprisingly little effect. 4. Subsequently, using empirical data, we tested whether observed changes in the onset of spring along the flyway explain the observed changes in migration schedules of pink-footed geese from 1990 to 2004. Model predictions generally agreed well with empirically observed migration patterns, with geese leaving the wintering grounds earlier while considerably extending their staging times in Norway.

Composition of Fuel Stores and Digestive Limitations to Fuel Deposition Rate in the Long-Distance Migratory Thrush Nightingale, Luscinia luscinia

During their autumn migratory phase, thrush nightingales (Luscinia luscinia) previously starved for 2 d were allowed to refuel under three different ambient temperature conditions (-7°, 7°, and 22°C). During the refueling period, as well as during the preceding control and starvation periods, food intake, body mass, and feces production were monitored. In addition, daily energy expenditure was measured during the refueling period. The compilation of the energy balance during the refueling period revealed an energy density of the deposited tissue of 33.6 kJ g⁻¹. Assuming that the deposited tissue consists of fat and protein exclusively, with energy densities of 39.6 and 5.5 kJ g⁻¹ wet mass, respectively, we estimated the deposited tissue to consist of 82% fat and 18% wet protein (6% dry protein and 12% water). Nitrogen balances during control, starvation, and refueling phases and during a period of prolonged and complete starvation indicated that 5% of the nutrient stores consisted of dry protein. Our results support recent findings that nutrient stores for migration often contain protein in addition to fat and consequently are 15%-25% less energy rich than pure fat stores. These proteins might be stored as muscle or other functional tissue and may be required to support the extra mass of the stores and/or reflect an incapacity of the metabolic machinery to catabolize fat exclusively. Fuel deposition rate was positively related with ambient temperature, whereas food intake rate was unaffected by temperature. These results indicate that the rate of fuel deposition is limited by a ceiling in food intake rate; when this ceiling is reached, fuel deposition rate is negatively affected by daily energy expenditure rate. To a certain extent, the ceiling in food intake rate varies depending on feeding conditions over the previous days. These variations in food intake capacity probably reflect the building and breakdown of gut tissues and/or gut enzyme systems and might be insensible and not evolutionary adaptive. Significant energetic costs, however, are probably associated with the maintenance of gut tissues. It is therefore feasible that changes in digestive capacity are regulated and are directed at energy economization.

Flocking and feeding in the fiddler crab (UCA tangeri): prey availability as risk-taking behaviour

For a full understanding of prey availability, it is necessary to study risk-taking behaviour of the prey. Fiddler crabs are ideally suited for such a study, as they have to leave their safe burrow to feed on the surface of the intertidal flats during low tide, thereby exposing themselves to avian predators. A study in an intertidal area along the coast of Mauritania showed that small crabs always stayed in the vicinity of their burrow, but large crabs wandered in large flocks (also referred to as droves) to feed on sea-grass beds downshore. Transplanting downshore feeding substrate to the burrowing zone of the small crabs proved that they too preferred to feed on it. Since small crabs can be preyed upon by more species of birds, this suggests that the decision not to leave the burrowing zone might be related to the risk of being fed upon by birds. We calculated predation risk from measurements on the density and feeding activity of the crabs, as well as the feeding density, the intake rate and the size selection of the avian predators. Per hour on the surface, crabs in a flock were more at risk than crabs feeding near their burrow. Thus, though flocking crabs may have benefited from ‘swamping the predator’ by emerging in maximum numbers during some tides only, this did not reduce their risk of predation below that of non-flocking crabs. Furthermore we found that irrespective of activity, large crabs suffered a higher mortality per tide from avian predators than small crabs. This suggests that large crabs could not sufficiently reduce their foraging time to compensate for the increased risk while foraging in a flock, even though they probably experienced better feeding conditions than small crabs staying near their burrow. The greater energy demands of large crabs were reflected in a greater surface area grazed. Thus, with increasing size a fiddler crab has to feed further away from its burrow and so may derive less protection from staying near to it. It seems that growing big does not reduce the risk of predation for fiddler crabs, as it does in many other species with indeterminate growth. As in such species, the most probable advantage of growing big is increased mating success. Ultimately, therefore, prey availability must be understood from the life-history decisions of the prey species.