Andrew D. West

Predicting the impacts of disturbance on shorebird mortality using a behaviour-based model

To assess the long-term effects of human disturbance on birds, ways of predicting its impacts on individual fitness and population size must be found. In this paper we use a behaviour-based model to predict the impact of human disturbance on oystercatchers (Haematopus ostralegus) on their intertidal feeding grounds in the Exe estuary in winter. The model predicted that, for the same overall area disturbed, numerous small disturbances would be more damaging than fewer, larger disturbances. When the time and energy costs arising from disturbance were included, disturbance could be more damaging than permanent habitat loss. Preventing disturbance during late winter, when feeding conditions were harder, practically eliminated its predicted population consequences. Although disturbance can cause increased mortality, it was not predicted to do so at the levels currently occurring in the Exe estuary.

Carrying capacity in overwintering migratory birds

The carrying capacity of the food supply in a non-breeding site for migrating birds can be measured either as the maximum number of bird-days that it can support or as the maximum numbers that can survive the non-breeding season. We identify the circumstances when it is appropriate to measure carrying capacity in either way, but we then show that birds may often emigrate or starve well before carrying capacity is reached, however it is defined. It is therefore unsafe to argue that, because the carrying capacity would not be reached, a change in site management would not affect birds. The key question in environmental impact studies of this kind is whether a change in site management would decrease the survival rate or the proportion of birds accumulating the body reserves required to migrate to the breeding grounds.

Intake rates and the functional response in shorebirds (Charadriiformes) eating macro-invertebrates

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.

TEST OF A BEHAVIOR-BASED INDIVIDUAL-BASED MODEL: RESPONSE OF SHOREBIRD MORTALITY TO HABITAT LOSS

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).

Benefits to shorebirds from invasion of a non-native shellfish

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.

A BEHAVIOR‐BASED MODELING APPROACH TO REDUCING SHOREBIRD–SHELLFISH CONFLICTS

Bottom cultivation of mussels on intertidal flats is practiced throughout the world. This often generates conflicts between commercial interests and competing birds such as oystercatchers. At the Menai Strait, United Kingdom, the overwinter consumption of 242 tonnes (1 metric tonne = 1000 kg) of commercially harvestable mussels (>40 mm) by oystercatchers in 1999–2000 was worth £133 000 (

The availability and quality of the mussel prey (Mytilus edulis) of oystercatchers (Haematopus ostralegus)

226 000 U.S. dollars). This represents 19% of the value of the landings. We used a behavior-based simulation model to predict the extent to which such losses can be reduced by novel commercial management practices, and to explore the consequences for the oystercatcher population. Simulations of novel lay management practices indicated that the losses of commercially harvestable mussels to oystercatchers can be considerably reduced by altering the shore level and/or extent of the commercial lays. We propose a novel management strategy for the bottom cultivation of mussels in intertidal areas. Seed mussels (15–20 mm) should...

Predicting mortality in novel environments: tests and sensitivity of a behaviour-based model

Abstract Spatial variations in the availability and quality of the mussel Mytilus edulis food supply of Oystercatchers Haematopus ostralegus on the Exe estuary, England, are described. Oystercatchers open mussels by stabbing into gaping mussels (or prising open closed ones) or by hammering a hole in either the dorsal or ventral shells. Spatial variations in the food supply are considered at four scales. In decreasing order of size, these are (i) whole mussel beds, (ii) zones within a mussel bed, (iii) different places within one zone, and (iv) different places within one clump of mussels. The first two scales are clearly related to exposure time. Both between and within the 12 main mussel beds of the estuary, most upshore mussels are up to 10% less likely than downshore mussels to be hidden under mud. However, upshore mussels of a given length contain less flesh, have thicker ventral shells and, except on high-level beds subject to wave erosion, have thicker dorsal shells than downshore mussels. Mussels at the top of the shore also contain the highest infestations of the helminth parasite of Oystercatchers, Psilostomum brevicolle. At a particular shore level, mussels of a given length have less flesh and thicker shells, though only on the dorsal side, in areas of high mussel density. Within one clump, mussels of a given length with thick dorsal shells have more flesh than those with thin shells. In contrast, flesh content is slightly higher in mussels that are thin on the ventral side. Flesh content and shell thickness on both sides are unaffected by whether a mussel is visible at the edge of a clump or hidden inside. Simulations with a model of foraging Oystercatchers suggested that variations in mussel availability (visibility and shell thickness) and flesh content at all four scales could sometimes have an important influence on intake rate. Most published values of intake rates of Oystercatchers eating surface-dwelling mussels may be biassed (and often considerably over-estimated) by the erroneous assumption that prey of average flesh content are taken.