Rudolf H. Drent

Subordinates explore but dominants profit: resource competition in high Arctic barnacle goose flocks

Social dominance plays an important role in assessing and obtaining access to patchy or scarce food sources in group-foraging herbivores. We investigated the foraging strategies of individuals with respect to their social position in the group in a flock of nonbreeding, moulting barnacle geese, Branta leucopsis, on high Arctic Spitsbergen. We first determined the dominance rank of individually marked birds. The dominance of an individual was best described by its age and its sex-specific body mass. Mating status explained the large variation in dominance among younger birds, as unpaired yearlings ranked lowest. In an artificially created, competitive situation, subordinate individuals occupied explorative front positions in the flock and were the first to find sites with experimentally enriched vegetation. Nevertheless, they were displaced quickly from these favourable sites by more dominant geese which were able to monopolize them. The enhanced sites were subsequently visited preferentially by individuals that succeeded in feeding there when the exclosures were first opened. Data on walking speed of foraging individuals and nearest-neighbour distances in the group suggest that subordinates try to compensate for a lower energy intake by exploring and by lengthening the foraging bout. Observations of our focal birds during the following breeding season revealed that females that returned to the study area were significantly more dominant in the previous year than those not seen in the area again. Copyright 2001 The Association for the Study of Animal Behaviour.

Travelling to breed

Traditionally, investigation of the dynamics of avian migration has been heavily biased towards the autumn return trip to the wintering quarters. Since the migratory prelude to breeding has direct fitness consequences, the European Science Foundation recently redressed the balance and sponsored a workshop on spring travels. We here survey the findings elucidating the complications arising during migration directed towards the breeding quarters. The evidence that early nesting confers advantage is overwhelming, hence demands of reproduction pose a constraint on both time and energy resources during spring. Both during migration and upon arrival there must therefore be strong selection in favour of saving time. Experimental results (e.g. using supplementary feeding) show that the date of laying is generally proximately constrained by the inability of the female to find sufficient nutrients. A key question thus concerns the implementation of the ‘capital’ strategy for breeding, the female accumulating nutrient stores along the way to bridge periods of shortage upon arrival on the breeding grounds. Eight studies on waterfowl (geese and eiders) and shorebirds (turnstones and knots) nesting in the arctic combine tracking of individuals (satellite telemetry) with direct observation (marked birds) and reconstruction of the origin of nutrients laid down in the eggs or in the form of body stores of the female parent (stable isotope signatures of tissues compared to potential food sources). The consensus emerges that in most cases a mixed strategy prevails, with nutrients garnered locally supplementing ‘imported’ body stores. The species concerned face a shortage of feeding time during incubation and suffer an energy deficit. Successful breeding thus depends on adequate fat depots that form part of the ‘capital’ the parents bring with them. Some headway has been made in predicting the extent of dependence on body stores for breeding in relation to body mass and length of migration from rates of fat deposition during stopover and fat consumption during flight for waterfowl. This work poses a challenge to refine field studies accordingly. The pressure to save time en route highlights the need to effectively exploit rich food resources. Several case studies underline the crucial significance of a very limited set of stopover sites, each with a narrow time window of optimal harvest. The influence of man at such sites often verges on the disastrous, and ongoing climate change may unhinge the finely tuned timing long-distance migrants depend on. There is a real need to extend this work to provide the scientific basis to underpin adequate conservation actions.

An Exploration of the Energetics of Leap-Frog Migration in Arctic Breeding Waders

Of the many problems posed by the phenomenon of bird migration, the interpretation of current migratory patterns in an evolutionary perspective remains a very thorny one. A widespread feature is for populations of the same species occupying distinctly disjunct breeding ranges to show differences in their wintering range as well. In some cases the more northerly breeding forms pass over the other populations to winter beyond the more southerly breeding forms. This migratory pattern, termed leap-frog migration, was first recognized more than 50 years ago and has been extensively discussed by Salomonsen (1955a,b), who in particular extended the list to include a number of waders. All authors who have grappled with this problem are in agreement that the displacement of the populations in winter must be attributed to the avoidance of competition (for review see Greenberg 1986). We accept this essential ingredient, and from this basic premise we will explore the implications of the spectacular differences in wintering area for the energy budgets of populations so segregated. In essence, we believe that pinpointing the essential differences in the energetic balance of the leap-frogging component may help to identify the selective advantages associated with the evolution of this migratory pattern.

AN ANALYSIS OF HATCHLING RESTING METABOLISM - IN SEARCH OF ECOLOGICAL CORRELATES THAT EXPLAIN DEVIATIONS FROM ALLOMETRIC RELATIONS

From data in the literature, an allometric equation is compiled for hatchling resting metabolic rate and an attempt is made to explain residual variation in terms of hatchling type, yolk and water content, embryonic and postnatal growth rate, and environmental circumstances (latitudinal distribution). The body mass exponent for resting metabolism in hatchlings was 0.86 and, thus, substantially different from the values compiled for adult birds (0.67-0.75). Relatively high hatchling metabolic rates were found for birds exhibiting high embryonic and postnatal growth rates, as well as for those species that hatched at high latitudes. A functional explanation is postulated for the correlations between hatchling metabolism and these three variables.

Skipping the Baltic : the emergence of a dichotomy of alternative spring migration strategies in Russian barnacle geese

1. Since the early 1990s, an increasing proportion of barnacle geese, Branta leucopsis, bound for breeding sites in the Russian Arctic delay their departure from the wintering quarters in the Wadden Sea by 4 weeks. These late-migrating geese skip spring stopover sites in the Baltic traditionally used by the entire population. 2. Individual geese from an arctic colony tracked by satellite or light-level geolocators during spring migration in 2004 and 2005 predominantly followed the new strategy, but a minority still maintained the traditional pattern. Despite a spread of more than 50 days in departure date from the Wadden Sea, both early and late departing females laid their eggs within the short time-window conferring breeding success. 3. The spread of these new migration routines coincided with a strong increase of overall numbers and the exploitation of new spring staging resources in the Wadden Sea. Counts from Estonia demonstrate that numbers have levelled off recently at the Baltic staging sites, suggesting that the capacity of these staging sites in spring has been reached. Although onset of spring affects migratory timing in barnacle geese, it cannot explain the observed delay in departure from the wintering grounds. 4. We hypothesize that the new migratory strategy has evolved in response to increased competition for food at spring staging sites in the Baltic. According to an analytical model of optimal migration, the geese should skip the Baltic whenever the energy deposition rate falls below 88% of the Wadden Sea value.

Energetics of Avian Growth: The Causal Link with BMR and Metabolic Scope

Our point of departure is Lack’s (1968) viewpoint that avian growth rate is one of the parameters adjusted in the course of evolution to help match the needs of the brood to the foraging ability of the parents in nidicolous species. A second selective pressure, valid especially for nidifugous species, is to minimize the period of heightened predation risk when the chicks are small and cannot yet fly. Furthermore, Lack reasoned that the seasonal timing of growth required synchrony, between the period of maximal demand of the growing young and the period of greatest food abundance. A further selective pressure for rapid growth rate might be expected in strongly seasonal environments where it is imperative to complete development before the onset of unfavourable conditions. These considerations are based on the premise that changes in growth rate bring about large changes in the daily ration required to raise the chick, i. e. the energetic consequences of alteration of growth rate loom large in the daily energy budget.

Short-term and long-term facilitation of goose grazing by livestock in the Dutch Wadden Sea area

We studied the impact of livestock grazing on the distribution ofBranta bernicla bernicla (Dark-bellied Brent goose) in the Dutch Wadden Sea during spring. It was hypothesized that livestock facilitate short-term (within-season) grazing for geese as well as long-term (over years). Therefore we measured grazing pressure by geese in salt marsh and polder areas that were either grazed (spring-grazed) or ungrazed during spring (summer-grazed). Additionally, we carried out a preference experiment with captive geese to test the preference between spring-grazed and summer-grazed polder swards. We furthermore compared patterns of use by geese between long-term ungrazed and grazed salt marshes.In May, there is a difference in grazing pressure by geese between polder pastures that are grazed or ungrazed during spring. In this month, the ungrazed polder pastures are abandoned and the geese shift to either the grazed polder pastures or to the salt marsh. Vegetation in the polder that had been spring-grazed had a lower canopy height and a higher tiller density than summer-grazed vegetation. The captive geese in the preference experiment showed a clear preference for vegetation that had been spring-grazed by sheep over ungrazed vegetation. Goose grazing pressure was negatively correlated to canopy height, both on the polder and on the salt marsh. Within the plant communities dominated byFestuca rubra andPuccinellia maritima, marshes that were intensively grazed by livestock generally had higher grazing pressure by geese than long-term ungrazed or lightly grazed salt marshes.

Fueling Incubation: Differential Use of Body Stores in Arctic and Temperate-breeding Barnacle Geese (Branta leucopsis)

ABSTRACT. We compared the use of body stores in breeding Barnacle Geese (Branta leucopsis) in traditional Arctic colonies in the Barents Sea with that in recently established temperate-zone breeding colonies in the Baltic Sea and North Sea by studying female body-mass loss and use of fat and protein stores during incubation. Average daily body-mass loss was almost identical in the 2 temperatebreeding populations (17.0 g and 16.5 g in Baltic Sea and North Sea, respectively), whereas Arctic-breeding females lost significantly less (10.6 g day-1). Temperate-breeding females initiated incubation with body mass 125 g higher than that of Arctic breeders, but at the end of incubation, body mass was similar among the 3 populations, averaging 1,458 g. Body-mass loss during incubation amounted to 23% (North Sea), 22% (Baltic Sea), and 15% (Barents Sea). Fat mass, as measured by isotope dilution in a subsample of females, was consistently higher in North Sea than in Barents Sea birds, but both populations showed similar rates of fat-mass loss (9.4 g day-1, on average). By contrast, loss of fat-free mass (assumed to represent wet protein) amounted to 9.3 g day-1 in North Sea birds but only 1.5 g day-1 in Barents Sea birds. Energy content of 1 g utilized body mass was 21.1 kJ (North Sea) and 34.9 kJ (Barents Sea), which equates to 376 kJ day-1 and 415 kJ day-1 drawn from stored energy, respectively. We suggest that differences in nest-attendance and postincubation demands are responsible for the differential use of body stores in temperate- and Arctic-breeding Barnacle Geese.