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ABSTRACT: Conservation objectives for non-breeding coastal birds (shorebirds and wildfowl) are determined from their population size at coastal sites. To advise coastal managers, models must predict quantitatively the effects of environmental change on population size or the demographic rates (mortality and reproduction) that determine it. As habitat association models and depletion models are not able to do this, we developed an approach that has produced such predictions thereby enabling policy makers to make evidence-based decisions. Our conceptual framework is individual-based ecology, in which populations are viewed as having properties (e.g. size) that arise from the traits (e.g. behaviour, physiology) and interactions of their constituent individuals. The link between individuals and populations is made through individual-based models (IBMs) that follow the fitness-maximising decisions of individuals and predict population-level consequences (e.g. mortality rate) from the fates of these individuals. Our first IBM was for oystercatchers Haematopus ostralegus and accurately predicted their density-dependent mortality. Subsequently, IBMs were developed for several shorebird and wildfowl species at several European sites, and were shown to predict accurately overwinter mortality, and the foraging behaviour from which predictions are derived. They have been used to predict the effect on survival in coastal birds of sea level rise, habitat loss, wind farm development, shellfishing and human disturbance. This review emphasises the wider applicability of the approach, and identifies other systems to which it could be applied. We view the IBM approach as a very useful contribution to the general problem of how to advance ecology to the point where we can routinely make meaningful predictions of how populations respond to environmental change.
Biological Reviews 11/2009; 85(3):413-34. · 9.07 Impact Factor
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ABSTRACT: Population models show that the response of migratory bird populations to a reduction in the area, or change in the quality, of their winter feeding areas depends critically on the shape (linear or non-linear) and strength (slope) of the relationship describing how the proportion dying from starvation changes with population density. Yet the parameters that define such density-dependent functions are extremely difficult to estimate from direct measurements of mortality at different population sizes. Even if they could be estimated, there would be uncertainty as to whether or not the relationship would remain the same, and thus provide reliable predictions, in the new circumstances for which forecasts are required. This paper summarizes studies of the Oystercatcher Haematopus ostralegus which aim to derive the shape and strength of the winter density-dependent mortality and emigration function for one estuary, under existing and new circumstances, from the responses of individual birds to each other and to their spatially and seasonally varying food supply. Based on these studies, an individuals-based, physiologically structured game theoretic distribution model has been built which predicts the carrying capacity and numbers of birds dying at different population sizes. The model also can be applied to Oyster-catchers occupying several estuaries distributed throughout their wintering range and can thus be extended to the entire biogeographical or global population. In addition, it can be used to identify some easily measured behavioural and ecological parameters that identify those species, from a wide range of taxa, that are most likely to be affected by habitat loss and change.
Ibis 06/2008; 137(s1):S56 - S66. · 2.43 Impact Factor
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ABSTRACT: Measurements were made of Oystercatcher corpses to find the most reliable measure of body condition in live birds. Total body weight was as accurate as muscle thickness in assessing the weight of the pectoral muscles, a major store of protein reserves. Total body weight was also a good indicator of fat reserves. Wing-length was the most reliable measure of body size. Body condition in live birds was measured as an individual's residual weight derived from a general linear model of (loge) body weight against age, month, (loge) wing-length and time since capture. On the Exe estuary in autumn, there was no significant difference in body condition between same-age Oystercatchers of different diets and feeding methods. In the winter months, however, body condition differed significantly between mussel feeders using different feeding methods, and between mussel feeders and birds that fed on worms and clams. Differences between diet/feeding methods were not the same for each age group. Birds with the highest mean body condition index in adults were mussel hammerers, in immatures mussel stabbers and in juveniles, worm/clam feeders. No significant difference in body condition was found between the sexes in any age group in autumn. In winter, when birds with the same diet/feeding method were considered, females were significantly heavier than males. However, when birds of all diets and feeding methods were combined, weight differences between the sexes were not apparent. We examine reasons why birds with certain diets and feeding methods may be heavier than others in winter, and why females are sometimes heavier than males. We conclude that, in most cases, it is due to differences between birds in their ability to achieve a common target weight. However, we conclude that adult females try to achieve a higher target weight than males, probably as a premigratory and prebreeding strategy. In the light of this, we consider the consequences for adult females, in terms of reproductive success and survival, of specializing in diets and feeding methods associated with lower body weights in winter.
Ibis 06/2008; 143(1):107 - 119. · 2.43 Impact Factor
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Ibis 06/2008; 142(1):132 - 138. · 2.43 Impact Factor
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ABSTRACT: Conservation managers need to be able to assess and prioritize issues that may affect their target habitats and species. In the Baie de Somme, France, conservation issues affecting overwintering shorebirds include hunting pressure, cockle fishing, recreational disturbance, Spartina encroachment, and changing sediment levels. We used an individual-based model to predict the effect of these issues on the survival of three shorebird species: dunlin Calidris alpina, oystercatcher Haematopus ostralegus and curlew Numenius arquata. In the model, removing hunting from the mudflats in the eastern part of the estuary had the greatest positive effect on shorebird survival. Oystercatcher survival decreased markedly when stocks of large cockles were reduced to <250 m-2 or numbers of fishermen per day were doubled. Short-term disturbance events, such as walkers, had more effect on shorebird survival than long-term events, such as fishermen. Dunlin, as a protected species, were able to feed outside the Réserve Naturelle and were unaffected by disturbance within the Réserve. Oystercatcher survival decreased when the number of disturbance events within the Réserve exceeded one h-1, and curlew survival when disturbance events exceeded six h-1. Spartina encroachment caused dunlin survival to decline steadily as feeding habitat was lost. Dunlin were also the species most affected by changes in sediment levels, likely to occur through either sedimentation or sea level rise.
Oryx 03/2008; 42(02):265 - 277. · 1.83 Impact Factor
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ABSTRACT: Abstract Individual-based models (IBMs) predict how animal populations will be affected by changes in their environment by modeling the responses of fitness-maximizing individuals to environmental change and by calculating how their aggregate responses change the average fitness of individuals and thus the demographic rates, and therefore size of the population. This paper describes how the need to develop a new approach to make such predictions was identified in the mid-1970s following work done to predict the effect of building a freshwater reservoir on part of the intertidal feeding areas of the shorebirds Charadrii that overwinter on the Wash, a large embayment on the east coast of England. The paper describes how the approach was developed and tested over 20 years (1976–1995) on a population of European oystercatchers Haematopus ostralegus eating mussels Mytilus edulis on the Exe estuary in Devon, England.The paper goes on to describe how individual-based modeling has been applied over the last 10 years to a wide range of environmental issues and to many species of shorebirds and wildfowl in a number of European countries. Although it took 20 years to develop the approach for 1 bird species on 1 estuary, ways have been found by which it can now be applied quite rapidly to a wide range of species, at spatial scales ranging from 1 estuary to the whole continent of Europe. This can now be done within the time period typically allotted to environmental impact assessments involving coastal bird populations in Europe.The models are being used routinely to predict the impact on the fitness of coastal shorebirds and wildfowl of habitat loss from (i) development, such as building a port over intertidal flats; (ii) disturbance from people, raptors, and aircraft; (iii) harvesting shellfish; and (iv) climate change and any associated rise in sea level. The model has also been used to evaluate the probable effectiveness of mitigation measures aimed at ameliorating the impact of such environmental changes on the birds. The first steps are now being taken to extend the approach to diving sea ducks and farmland birds during the nonbreeding season.The models have been successful in predicting the observed behavior and mortality rates in winter of shorebirds on a number of European estuaries, and some of the most important of these tests are described. These successful tests of model predictions raise confidence that the model can be used to advise policy makers concerned with the management of the coast and its important bird populations.
Natural Resource Modeling 02/2008; 21(1):3 - 71. · 0.49 Impact Factor
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ABSTRACT: Introductions of non-native species are seen as major threats to ecosystem function and biodiversity. However, invasions of aquatic habitats by non-native species are known to benefit generalist consumers that exhibit dietary switches and prey upon the exotic species in addition to or in preference to native ones. There is, however, little knowledge concerning the population-level implications of such dietary changes. Here, we show that the introduction of the Manila clam Tapes philippinarum into European coastal waters has presented the Eurasian oystercatcher Haematopus ostralegus ostralegus with a new food resource and resulted in a previously unknown predator-prey interaction between these species. We demonstrate, with an individuals-based simulation model, that the presence of this non-native shellfish, even at the current low density, has reduced the predicted over-winter mortality of oystercatchers at one recently invaded site. Further increases in clam population density are predicted to have even more pronounced effects on the density dependence of oystercatcher over-winter mortality. These results suggest that if the Manila clam were to spread around European coastal waters, a process which is likely to be facilitated by global warming, this could have considerable benefits for many shellfish-eating shorebird populations.
Proceedings of the Royal Society B: Biological Sciences 07/2007; 274(1616):1449-55. · 5.41 Impact Factor
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John D Goss-Custard,
Niall H K Burton,
Nigel A Clark,
Peter N Ferns,
Selwyn McGrorty,
Christopher J Reading,
Mark M Rehfisch,
Richard A Stillman,
Ian Townend,
Andrew D West,
David H Worrall
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ABSTRACT: In behavior-based individual-based models (IBMs), demographic functions are emergent properties of the model and are not built into the model structure itself, as is the case with the more widely used demography-based IBMs. Our behavior-based IBM represents the physiology and behavioral decision making of individual animals and, from that, predicts how many survive the winter nonbreeding season, an important component of fitness. This paper provides the first test of such a model by predicting the change in winter mortality of a charadriid shorebird following removal of intertidal feeding habitat, the main effect of which was to increase bird density. After adjusting one calibration parameter to the level required to replicate the observed mortality rate before habitat loss, the model predicted that mortality would increase by 3.65%, which compares well with the observed increase of 3.17%. The implication that mortality was density-dependent was confirmed by predicting mortality over a range of bird densities. Further simulations showed that the density dependence was due to an increase in both interference and depletion competition as bird density increased. Other simulations suggested that an additional area of mudflat, equivalent to only 10% of the area that had been lost, would be needed by way of mitigation to return mortality to its original level. Being situated at a high shore level with the flow of water in and out impeded by inlet pipes, the mitigating mudflat would be accessible to birds when all mudflats in the estuary were covered at high tide, thus providing the birds with extra feeding time and not just a small replacement mudflat. Apart from providing the first, and confidence-raising, test of a behavior-based IBM, the results suggest (1) that the chosen calibration procedure was effective; (2) that where no new fieldwork is required, and despite being parameter rich, a behavior-based IBM can be parameterized quickly (few weeks), and thus cheaply, because so many of the parameter values can be obtained from the literature and are embedded in the model; and (3) that behavior-based IBMs can be used to explore system behavior (e.g., the role of depletion competition and interference competition in density-dependent mortality).
Ecological Applications 01/2007; 16(6):2215-22. · 5.10 Impact Factor
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John D Goss-Custard,
Andrew D West,
Michael G Yates,
Richard W G Caldow,
Richard A Stillman,
Louise Bardsley,
Juan Castilla,
Macarena Castro,
Volker Dierschke,
Sarah E A Le V Dit Durell, [......],
Alejandro Perez-Hurtado,
Danny Rogers,
Gregor Scheiffarth,
Humphrey Sitters,
William J Sutherland,
Patrick Triplet,
Dave H Worrall,
Yuri Zharikov,
Leo Zwarts,
Richard A Pettifor
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ABSTRACT: 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-2). 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.
Biological Reviews 12/2006; 81(4):501-29. · 9.07 Impact Factor
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John D. Goss-Custard
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ABSTRACT: 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.
Biological Reviews 10/2006; 81(04):501 - 529. · 9.07 Impact Factor
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Richard A. Stillman,
Andrew D. West, John D. Goss-Custard,
Richard W. G. Caldow,
Selwyn Mcgrorty,
Sarah E. A.Le V. Dit Durell,
Michael G. Yates,
Philip W. Atkinson,
Nigel A. Clark,
Michael C. Bell,
Peter J. Dare,
Matt Mander
Journal of Applied Ecology 12/2003; 40(6):1090 - 1101. · 5.05 Impact Factor
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ABSTRACT: In animal pairs, males are often more vigilant than females. This is generally assumed to result from mate guarding (either against predators or other males). However, when males have conspicuous secondary sexual characteristics, they could be constrained to be more vigilant because of a higher predation risk than females. We attempted to distinguish between the "male constraint hypothesis" and two variations of the mate-guarding hypothesis by studying the vigilance behavior of the sexually dimorphic wigeon during early winter, when some males are in breeding plumage and some are not and when not all males are paired. The proportion of time spent vigilant by paired males in breeding plumage was five times higher than any other category of males or females. We found no significant differences between the vigilance levels of unpaired male wigeon in cryptic and in breeding plumage and therefore rejected the male constraint hypothesis. As vigilance levels of paired and unpaired females did not differ either, we rejected the hypothesis that paired males invest in vigilance to reduce their mate's need to be vigilant to predation risks. Paired females interacted less frequently with other wigeon than unpaired ones, and it is probably to protect their female from other males that paired male wigeon increase their vigilance times. Copyright 2003.
Behavioral Ecology. 01/2003; 14(5):724-729.
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Journal of Animal Ecology 05/2002; 71(3):532 - 541. · 4.94 Impact Factor
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Journal of Avian Biology 01/2002; 33(4):358-365. · 2.28 Impact Factor
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ABSTRACT: Interference is an important component of food competition but is often difficult to detect and measure in natural animal populations. Although interference has been shown to occur between oystercatchers Haematopus ostralegus L. feeding on mussels Mytilus edulis L., four previous studies have not detected interference between oystercatchers feeding on cockles Cerastoderma edule L. In contrast, this study detected interference between cockle-feeding oystercatchers in the Baie de Somme, France. Prey stealing (kleptoparasitism), one of the main causes of interference between mussel-feeders, also occurred between oystercatchers in the Baie de Somme. The kleptoparasitism rate was related to the natural variation in the food supply, tending to be higher when cockles were rare. Feeding rate was negatively related to competitor density, so providing evidence for interference, but, as in mussel-feeders, only above a threshold density of about 50–100 birds ha−1. The strength of interference at a fixed competitor density was related to the cockle food supply, usually being greater when cockles were rare. Previous studies probably failed to detect interference between cockle-feeders because competitor densities were too low, or cockles were too abundant, or because they were not conducted during late winter when interference is most intense. The study shows that natural variation in the food supply can influence the strength of interference within an animal population and provides support for those behaviour-based interference models which predict that the strength of interference will be greatest when competitor densities are high and prey scarce.
Journal of Animal Ecology 12/2001; 68(2):254 - 265. · 4.94 Impact Factor
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ABSTRACT: Summary1. Above a threshold density of ≈ 100 birds ha-1, strong interference occurred between redshank Tringa totanus (Linnaeus) feeding by sight on the amphipod crustacean Corophium volutator (Pallas). No aggressive interactions occurred between the birds and the probable cause was prey depression.2. Redshank fed in a square metre of mud that had recently been exploited by another redshank much less often than would be expected by chance. By avoiding areas where prey would have been recently exploited, the feeding rate of redshank was up to three times faster than it would have been had they not avoided other foraging redshank.3. Bar-tailed godwit fed in a square metre of mud that had been recently exploited by another godwit much more often than would be expected by chance in randomly moving birds. They tended to flock while foraging and showed no tendency to avoid areas where prey would have been recently exploited.4. There was no evidence that interference occurred between bar-tailed godwit Limosa lapponica (Linnaeus) feeding on the polychaete lugworm Arenicola marina (Linnaeus) at densities below 300 birds ha-1, even though aggressive interactions occurred between birds.
Journal of Animal Ecology 02/2000; 69(2):314 - 322. · 4.94 Impact Factor
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ABSTRACT: We develop a model of predators foraging within a single patch, on prey that become temporarily immune to predation (depressed) after detecting a predator. Interference through prey depression occurs because the proportion of vulnerable prey (and hence intake rate) decreases as predator density increases. Predators in our model are not forced to move randomly within the patch, as is the case in other similar models, but can avoid areas of depressed prey and so preferentially forage over vulnerable prey. We compare the extent to which different avoidance rules (e.g., move more quickly over depressed prey or turn if approaching depressed prey) influence the amount of time spent foraging over depressed and vulnerable prey, and how this influences the strength of interference. Although based on a different mechanism, our model produces two similar general predictions to interference models based on direct interactions between predators: the strength of interference increases with (1) increased competitor density and (2) decreased prey encounter rate. This suggests that there are underlying similarities in the nature of interference even when it arises through different processes. Not surprisingly, avoidance of depressed prey can substantially reduce the strength of interference compared with random foraging. However, we identify the region of the model's parameter space in which this reduction is particularly large and show that the only system for which suitable data are available, redshank Tringa totanus feeding on Corophium volutator, falls within this region. The model shows that, by adjusting its search path to avoid areas of depressed prey, a predator can substantially reduce the amount of the interference it experiences and that this applies over a wide range of parameter space, including the region occupied by a real system. This suggests that behavior-based interference models should consider predator search pattern if they are to accurately predict the strength of the interference.
Behavioral Ecology. 01/2000; 11(6):597-605.
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John D Goss-Custard,
Andrew D West,
Michael G. Yates,
Richard W. G. Caldow,
Richard A Stillman,
Louise Bardsley,
Juan Castilla,
Macarena Castro,
Volker Dierschke,
Sarah. E. A. Le V. dit Durell,
Goetz Eichhorn,
Bruno J. Ens,
Klaus-Michael Exo,
P. U. Udayangani-Fernando,
Peter N Ferns,
Yuri Zharikov,
et al
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ABSTRACT: 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 11 ('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(-2)). 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.
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ABSTRACT: A behaviour-based model was used to explore the effect of an extension of the port at Le Havre (Port 2000), and the effect of proposed mitigation measures, on the mortality and body condition of the three main shorebird species that overwinter in the estuary of the river Seine, France. In the model, a 20% reduction in the area of mudflats on the north side of the estuary had little effect on curlew Numenius arquata mortality and body condition but significantly increased mortality and decreased body condition in dunlin Calidris alpina and oystercatchers Haematopus ostralegus. Disturbance of feeding birds both day and night had a significant effect on the mortality and body condition of all three shorebird species, as did disturbance of roosting birds. Disturbance of feeding birds in the daytime only had a significant effect on dunlin mortality and body condition, but not that of curlew and oystercatchers. In the model, the creation of a buffer zone to reduce disturbance of feeding birds restored shorebird mortality and body condition to pre-disturbance levels. A new mudflat area was also effective in mitigating the effect of habitat loss on all three shorebirds and in mitigating the effect of roost disturbance on dunlin and curlew. However, a new mudflat area was not effective in mitigating the effect of roost disturbance on oystercatcher mortality and body condition. The effectiveness of the mitigating mudflat depended as much on its size as its quality. We believe that this is the first time that anyone has been able to forecast the efficacy of proposed mitigation measures.
Biological Conservation.
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ABSTRACT: It is increasingly important to be able to monitor and maintain the quality of estuaries for overwintering shorebirds. Bird numbers alone are not sufficient to indicate quality nor, as recent research shows, can it be assumed that site quality is being maintained simply because there is enough food to meet the birds’ physiological demands; i.e., the amount of food available per bird in autumn needs to be greater than the amount eaten. But how much greater? We used a simple individuals-based and behaviour-based model to explore the factors that affect the relationship between overwinter mortality and the amount of food available per bird in autumn (the food:bird ratio). The aims were to explore how the natural history characteristics of a shorebird species affected the shape of this relationship, in order to identify characteristics of the system that should be included in any monitoring programme of site quality. In all cases there was a minimum threshold food:bird ratio above which mortality did not vary and below which mortality increased steadily. The amount of food per bird required at the threshold varied from one to six times the amount actually consumed by each bird during winter, depending on the amount of realism included in the model. The minimum threshold ratio was most strongly influenced by the shape of the relationship between intake rate and prey density – the functional response – and, when interference competition occurred, by the distribution of the birds’ main food supply. Simulations with realistic spatial distributions of prey predicted that the minimum required amount of food per bird would be between two and five times the amount actually consumed. The threshold approach could provide a simple method of monitoring estuary quality based on food supply and bird numbers. It also provides a potentially clearly-defined and unambiguous measure of the carrying capacity of a site. If applied to a sufficient number of real estuaries, general guidelines for maintaining estuary quality based on estuary characteristics, food supply and bird species might be established.
Biological Conservation.
