As field determinations take much effort, it would be useful to be able to predict easily the coe... more As field determinations take much effort, it would be useful to be able to predict easily the coefficients describing the functional response of free-living predators, the function relating food intake rate to the abundance of food organisms in the environment. As a means easily to parameterise an individual-based model of shorebird Charadriiformes populations, we attempted this for shorebirds eating macro-invertebrates. Intake rate is measured as the ash-free dry mass (AFDM) per second of active foraging ; i.e. excluding time spent on digestive pauses and other activities, such as preening. The present and previous studies show that the general shape of the functional response in shorebirds eating approximately the same size of prey across the full range of prey density is a decelerating rise to a plateau, thus approximating the Holling type II (' disc equation ') formulation. But field studies confirmed that the asymptote was not set by handling time, as assumed by the disc equation, because only about half the foraging time was spent in successfully or unsuccessfully attacking and handling prey, the rest being devoted to searching. A review of 30 functional responses showed that intake rate in free-living shorebirds varied independently of prey density over a wide range, with the asymptote being reached at very low prey densities (<150/m x2). Accordingly, most of the many studies of shorebird intake rate have probably been conducted at or near the asymptote of the functional response, suggesting that equations that predict intake rate should also predict the asymptote. A multivariate analysis of 468 ' spot ' estimates of intake rates from 26 shorebirds identified ten variables, representing prey and shorebird characteristics, that accounted for 81% of the variance in logarithm-transformed intake rate. But four-variables accounted for almost as much (77.3%), these being bird size, prey size, whether the bird was an oystercatcher Haematopus ostralegus eating mussels Mytilus edulis, or breeding. The four variable equation under-predicted, on average, the observed 30 estimates of the asymptote by 11.6 %, but this discrepancy was reduced to 0.2% when two suspect estimates from one early study in the 1960s were removed. The equation therefore predicted the observed asymptote very successfully in 93% of cases. We conclude that the asymptote can be reliably predicted from just four easily measured variables. Indeed, if the birds are not breeding and are not oystercatchers eating mussels, reliable predictions can be obtained using just two variables, bird and prey sizes. A multivariate analysis of 23 estimates of the half-asymptote constant suggested they were smaller when prey were small but greater when the birds were large, especially in oystercatchers. The resulting equation could be used to predict the half-asymptote constant, but its predictive power has yet to be tested. As well as predicting the asymptote of the functional response, the equations will enable research workers engaged in many areas of shorebird ecology and behaviour to estimate intake rate without the need for conventional time-consuming field studies, including species for which it has not yet proved possible to measure intake rate in the field.
A drop in body temperature allows significant energy savings in endotherms, but facultative heter... more A drop in body temperature allows significant energy savings in endotherms, but facultative heterothermy is usually restricted to small animals. Here we report that king penguin chicks (Aptenodytes patagonicus), which are able to fast for up to 5 months in winter, undergo marked seasonal heterothermy during this period of general food scarcity and slowdown of growth. They also experience short-term heterothermy below 20 °C in the lower abdomen during the intense (re)feeding period in spring, induced by cold meals and adverse weather. The heterothermic response involves reductions in peripheral temperature, reductions in thermal core volume and temporal abandonment of high core temperature. Among climate variables, air temperature and wind speed show the strongest effect on body temperature, but their effect size depends on physiological state. The observed heterothermy is remarkable for such a large bird (10 kg before fasting), which may account for its unrivalled fasting capacity among birds.
Understanding stopover decisions of long-distance migratory birds is crucial for conservation and... more Understanding stopover decisions of long-distance migratory birds is crucial for conservation and management of these species along their migratory flyway. Recently, an increasing number of Barnacle geese breeding in the Russian Arctic have delayed their departure from their wintering site in the Netherlands by approximately one month and have reduced their staging duration at stopover sites in the Baltic accordingly. Consequently, this extended stay increases agricultural damage in the Netherlands. Using a dynamic state variable approach we explored three hypotheses about the underlying causes of these changes in migratory behavior, possibly related to changes in (i) onset of spring, (ii) potential intake rates and (iii) predation danger at wintering and stopover sites. Our simulations showed that the observed advance in onset of spring contradicts the observed delay of departure, whereas both increased predation danger and decreased intake rates in the Baltic can explain the delay. Decreased intake rates are expected as a result of increased competition for food in the growing Barnacle goose population. However, the effect of predation danger in the model was particularly strong, and we hypothesize that Barnacle geese avoid Baltic stopover sites as a response to the rapidly increasing number of avian predators in the area. Therefore, danger should be considered as an important factor influencing Barnacle goose migratory behavior, and receive more attention in empirical studies.
2006. Spring stopover routines in Russian Barnacle Geese Branta leucopsis tracked by resightings ... more 2006. Spring stopover routines in Russian Barnacle Geese Branta leucopsis tracked by resightings and geolocation. Ardea 94(3): 667–678. By attaching 9-g loggers (recording dusk and dawn times to a memory chip) to the coded leg rings of Barnacle Geese Branta leucopsis caught in a breeding colony at Tobseda (68°35'N, 52°20'E) on the arctic coast of the Russian Federation in 2003 and by their recapture and retrieval in the 2004 season, we reconstructed the year-round movements of 19 females. We analysed spring migration movements of logged birds until they entered the zone of continuous daylight (c. 20 May), supplemented by ring reading in the colony. The technique also enabled description of incubation rhythm in these arctic breeders, allowing inferences about attempt and timing of breeding without the necessity of direct observa-tion. Although the birds did not necessarily travel together, most trav-elled on 14–15 (9 individuals) and 17–18 May (10 birds) when favourable condi...
Within three decades, the barnacle goose population wintering on the European mainland has dramat... more Within three decades, the barnacle goose population wintering on the European mainland has dramatically increased in numbers and extended its breeding range. The expansion has occurred both within the Arctic as well as by the colonization of temperate areas. Studies of ...
As field determinations take much effort, it would be useful to be able to predict easily the coe... more As field determinations take much effort, it would be useful to be able to predict easily the coefficients describing the functional response of free-living predators, the function relating food intake rate to the abundance of food organisms in the environment. As a means easily to parameterise an individual-based model of shorebird Charadriiformes populations, we attempted this for shorebirds eating macro-invertebrates. Intake rate is measured as the ash-free dry mass (AFDM) per second of active foraging ; i.e. excluding time spent on digestive pauses and other activities, such as preening. The present and previous studies show that the general shape of the functional response in shorebirds eating approximately the same size of prey across the full range of prey density is a decelerating rise to a plateau, thus approximating the Holling type II (' disc equation ') formulation. But field studies confirmed that the asymptote was not set by handling time, as assumed by the disc equation, because only about half the foraging time was spent in successfully or unsuccessfully attacking and handling prey, the rest being devoted to searching. A review of 30 functional responses showed that intake rate in free-living shorebirds varied independently of prey density over a wide range, with the asymptote being reached at very low prey densities (<150/m x2). Accordingly, most of the many studies of shorebird intake rate have probably been conducted at or near the asymptote of the functional response, suggesting that equations that predict intake rate should also predict the asymptote. A multivariate analysis of 468 ' spot ' estimates of intake rates from 26 shorebirds identified ten variables, representing prey and shorebird characteristics, that accounted for 81% of the variance in logarithm-transformed intake rate. But four-variables accounted for almost as much (77.3%), these being bird size, prey size, whether the bird was an oystercatcher Haematopus ostralegus eating mussels Mytilus edulis, or breeding. The four variable equation under-predicted, on average, the observed 30 estimates of the asymptote by 11.6 %, but this discrepancy was reduced to 0.2% when two suspect estimates from one early study in the 1960s were removed. The equation therefore predicted the observed asymptote very successfully in 93% of cases. We conclude that the asymptote can be reliably predicted from just four easily measured variables. Indeed, if the birds are not breeding and are not oystercatchers eating mussels, reliable predictions can be obtained using just two variables, bird and prey sizes. A multivariate analysis of 23 estimates of the half-asymptote constant suggested they were smaller when prey were small but greater when the birds were large, especially in oystercatchers. The resulting equation could be used to predict the half-asymptote constant, but its predictive power has yet to be tested. As well as predicting the asymptote of the functional response, the equations will enable research workers engaged in many areas of shorebird ecology and behaviour to estimate intake rate without the need for conventional time-consuming field studies, including species for which it has not yet proved possible to measure intake rate in the field.
A drop in body temperature allows significant energy savings in endotherms, but facultative heter... more A drop in body temperature allows significant energy savings in endotherms, but facultative heterothermy is usually restricted to small animals. Here we report that king penguin chicks (Aptenodytes patagonicus), which are able to fast for up to 5 months in winter, undergo marked seasonal heterothermy during this period of general food scarcity and slowdown of growth. They also experience short-term heterothermy below 20 °C in the lower abdomen during the intense (re)feeding period in spring, induced by cold meals and adverse weather. The heterothermic response involves reductions in peripheral temperature, reductions in thermal core volume and temporal abandonment of high core temperature. Among climate variables, air temperature and wind speed show the strongest effect on body temperature, but their effect size depends on physiological state. The observed heterothermy is remarkable for such a large bird (10 kg before fasting), which may account for its unrivalled fasting capacity among birds.
Understanding stopover decisions of long-distance migratory birds is crucial for conservation and... more Understanding stopover decisions of long-distance migratory birds is crucial for conservation and management of these species along their migratory flyway. Recently, an increasing number of Barnacle geese breeding in the Russian Arctic have delayed their departure from their wintering site in the Netherlands by approximately one month and have reduced their staging duration at stopover sites in the Baltic accordingly. Consequently, this extended stay increases agricultural damage in the Netherlands. Using a dynamic state variable approach we explored three hypotheses about the underlying causes of these changes in migratory behavior, possibly related to changes in (i) onset of spring, (ii) potential intake rates and (iii) predation danger at wintering and stopover sites. Our simulations showed that the observed advance in onset of spring contradicts the observed delay of departure, whereas both increased predation danger and decreased intake rates in the Baltic can explain the delay. Decreased intake rates are expected as a result of increased competition for food in the growing Barnacle goose population. However, the effect of predation danger in the model was particularly strong, and we hypothesize that Barnacle geese avoid Baltic stopover sites as a response to the rapidly increasing number of avian predators in the area. Therefore, danger should be considered as an important factor influencing Barnacle goose migratory behavior, and receive more attention in empirical studies.
2006. Spring stopover routines in Russian Barnacle Geese Branta leucopsis tracked by resightings ... more 2006. Spring stopover routines in Russian Barnacle Geese Branta leucopsis tracked by resightings and geolocation. Ardea 94(3): 667–678. By attaching 9-g loggers (recording dusk and dawn times to a memory chip) to the coded leg rings of Barnacle Geese Branta leucopsis caught in a breeding colony at Tobseda (68°35'N, 52°20'E) on the arctic coast of the Russian Federation in 2003 and by their recapture and retrieval in the 2004 season, we reconstructed the year-round movements of 19 females. We analysed spring migration movements of logged birds until they entered the zone of continuous daylight (c. 20 May), supplemented by ring reading in the colony. The technique also enabled description of incubation rhythm in these arctic breeders, allowing inferences about attempt and timing of breeding without the necessity of direct observa-tion. Although the birds did not necessarily travel together, most trav-elled on 14–15 (9 individuals) and 17–18 May (10 birds) when favourable condi...
Within three decades, the barnacle goose population wintering on the European mainland has dramat... more Within three decades, the barnacle goose population wintering on the European mainland has dramatically increased in numbers and extended its breeding range. The expansion has occurred both within the Arctic as well as by the colonization of temperate areas. Studies of ...
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Papers by Götz Eichhorn