1. Top of page
  2. Abstract
  5. Acknowledgments

The internationally important populations of waterbirds that winter in the United Kingdom can face intense pressure from human disturbance as a result of the high urbanization found around many protected coastal or inland wetland sites. Here, I describe and evaluate an approach that has been used to investigate the spatial effects of human disturbance on waterbirds. Rather than directly investigating behavioural responses to individual disturbance events, the presence of features in the landscape associated with disturbance is instead used as a surrogate, with the essential aim being to demonstrate that bird numbers or densities are depressed or their behaviour altered in proximity to areas used by humans. This paper first describes case studies that demonstrate the limitations of the basic inference (i.e. that disturbance influences patterns of waterbird distribution or behaviour) and then how investigations might be strengthened. For conclusions to be sound, it is particularly important that other factors, such as food supply, that might also explain the spatial patterns observed are considered or other corroborative evidence presented. The approach is thus least applicable in the most heterogeneous environments where many factors, perhaps spatially autocorrelated, may explain variation in distribution or behaviour. However, greatest confidence in the validity of conclusions may be gained where studies are able to show (ideally by experimental manipulation) that species’ distributions or behaviour vary temporally in line with the levels of human use of the features examined. Although its aim and scope are thus limited, the use of a landscape approach, provided that it takes into account other factors affecting spatial variation in bird abundance or behaviour, can provide a preliminary assessment of species avoidance of key sources of disturbance that may offer a framework for more detailed investigation.

The coastal and freshwater wetlands of Europe support internationally important numbers of waterbirds in winter (Kershaw & Cranswick 2003, Rehfisch et al. 2003, Collier et al. 2005) with many sites designated as Special Protection Areas owing to their significance for these species (Stroud et al. 2001). However, many important sites are highly urbanized, with habitat having been lost to industry, housing, harbour developments and barrage schemes (Davidson et al. 1991, Burton et al. 2006, Burton in press). With these developments, there has been increased disturbance from, for example, construction work itself, road and rail traffic, and also increased and varied recreational use of wetland sites. Recreational use may include water-based activities such as sailing, water-skiing, jet-skiing and windsurfing, as well as walking, dog-walking, cycling and trail-biking (Davidson & Rothwell 1993).

The effects of these activities on waterbirds have been the subject of considerable study, with research often focused on measuring the behavioural responses shown by individuals to different disturbance stimuli (e.g. Burger 1981, Roberts & Evans 1993, Smit & Visser 1993, Fitzpatrick & Bouchez 1998, Triplet et al. 1998, 2001, Mathers et al. 2000, Lafferty 2001a, 2001b, Thomas et al. 2003). Such studies may provide information on, for example, the proportions of different activities causing disturbance, the distances at which different species take flight, or the time taken for birds to return to a site or resume foraging, though, as noted recently, the level of response shown may depend on the stress that individuals are under (Triplet & Gembarski 2001, McGowan et al. 2002, Stillman & Goss-Custard 2002, Beale & Monaghan 2004). Nevertheless, such information, particularly in relation to the frequency of disturbance, may help to indicate whether individual fitness (e.g. body condition, survival) is potentially impacted by disturbance and in this way help in the development of modelling studies to predict impacts on local populations (West et al. 2002, Goss-Custard et al. 2006, Stillman et al. 2006). Alongside such modelling studies, there is still a need for empirical evidence that disturbance may actually impact bird populations (Hill et al. 1997).

Often, information about the potential impact of human presence is required from a site perspective, for instance by those monitoring the status of sites designated for their waterbird interest. At this level, information may initially just be needed on whether the distributions of birds within a site are affected by particular human activities.

Here, I describe and evaluate an approach that has been used to investigate the spatial effects of human disturbance, whereby the presence of features in the landscape associated with disturbance is used as a surrogate. The premise of these studies is that the abundance, presence/absence or behaviour of birds will vary spatially in relation to certain landscape features because they act as sources of human disturbance. This paper first describes case studies that demonstrate the limitations of the basic inference (i.e. that disturbance influences patterns of waterbird distribution or behaviour) and then how investigations might be strengthened through consideration of other factors that might also explain spatial variation in species abundance or behaviour and temporal changes in species’ responses. The case studies focus on waterbirds, though the approach has been used for other taxa. The limitations of the aim and scope of the approach, as well as its potential role as a preliminary step for further more detailed analyses are discussed.


  1. Top of page
  2. Abstract
  5. Acknowledgments

Case study 1

At its most basic, a landscape approach aims simply to show that bird numbers are reduced in proximity to areas used by humans but with no consideration of other factors that might influence species’ distributions. Such an approach is limited to the simplest of systems where the influence of other habitat characteristics is likely to be minimal. Keller (1991), for example, showed that Pink-footed Geese Anser brachyrhynchus and Greylag Geese A. anser flocks in Scotland were distributed further from roads when feeding in agricultural fields than would be expected at random. Although food resources might be expected to be fairly uniformly distributed across the available habitat (and not be related to distance from roads) it is possible that goose numbers were reduced close to roads for other coincident reasons – for example, because the risk of predation might be greater at field edges. However, the study did demonstrate that geese close to roads took flight in response to passing cars, thus strengthening the argument that disturbance was the cause of the distributional patterns observed.

Case study 2

In more heterogeneous habitats, it is clearly important that other factors that might affect spatial variation in the aspect being studied (i.e. the abundance, presence/absence or behaviour of birds) are also taken into account. Burton et al. (2002a) used data from six estuaries in southern England taken from the Wetland Bird Survey (WeBS) Low Tide Count Scheme to investigate whether waterbird numbers were related to the presence of nearby footpaths, roads, railways and towns. Generalized linear models (GLMs) assessed whether species’ numbers varied according to the proportion of each intertidal count section within a specified distance of each landscape feature (that which maximized model fit), estuary, month, count section area, whether count sections bordered the low water mark and the proximity of sections to the nearest footpath access point (car park, town or road). Numbers of six of nine species – Common Shelduck Tadorna tadorna, Red Knot Calidris canutus, Dunlin C. alpina, Black-tailed Godwit Limosa limosa, Eurasian Curlew Numenius arquata and Common Redshank Tringa totanus– were significantly lower if a footpath was close to a count section, whereas those of Dark-bellied Brent Geese Branta bernicla bernicla were significantly greater. Numbers of three species were also reduced close to roads, four close to railways and one close to towns. In this case, analysis was limited as it is possible that such results might have been due to variations in factors such as substrate type, water depth and food supply, rather than disturbance. However, although it was not directly assessed, food availability was partially accounted for by classifying whether count sections bordered the low water mark. (As counts were undertaken at low tide, those sections that bordered the low water mark might have been expected to hold more birds as prey can be much more available in wetter sediments: Kelsey & Hassall 1989.) There is also likely to have been considerable spatial autocorrelation between the effects of the different landscape features considered; avoidance of footpaths might also have been coincident with an avoidance of the upper intertidal zone due to the risk of predation (Whitfield 2003). Nevertheless, the study found a significant temporal effect that strengthened the inference that disturbance (from footpaths at least) was a major cause of the distributional patterns observed. Numbers of four species – Dark-bellied Brent Goose, Common Shelduck, Dunlin and Common Redshank – decreased with increased proximity to a footpath access point at weekends, when the use of footpaths is likely to have been at its greatest, whereas this effect was only significant for one species, Brent Goose, on weekdays.

Case study 3

Burton et al. (2002b) used a similar methodology to investigate the effects of construction work disturbance – from the building of a barrage (Burton in press), new housing, a hotel, a road bridge and other land-claim – on waterbirds using the former estuarine mudflats of Cardiff Bay, south Wales. GLMs tested whether waterbird numbers and feeding behaviour were related to the year, month, state of tide, the count section and whether or not the section bordered active construction work. Numbers of five of seven species – Eurasian Teal Anas crecca, Eurasian Oystercatcher Haematopus ostralegus, Dunlin, Eurasian Curlew and Common Redshank – and the feeding activity of all four waders were all significantly lower on intertidal count sections adjacent to construction work. By specifying count section as a variable, models took into account not just the effect of area in explaining bird numbers, but also, indirectly, variation in resource availability. Critically, as ‘disturbed’ sections were only classified as such for the duration of active construction work, disturbance is unlikely to have been coincidentally correlated with food supply.

Case study 4

The fourth case study considered investigated the effects of disturbance on waders wintering on agricultural grassland in coastal Sussex (Milsom et al. 1998). The study used logistic regression models (and a stepwise procedure) to determine whether the frequency with which waders used particular fields was related to three ‘disturbance’ factors, though importantly also a suite of other ground habitat and topographical variables. Three measures acted as surrogates for human disturbance: the presence or absence of public footpaths running through study fields, the presence or absence of roads along field boundaries and the proximity of each field to occupied buildings. Models indicated that Eurasian Curlew were less likely to use fields that bordered roads than those that did not, but that the presence of roads had no influence on use of fields by Lapwing Vanellus vanellus or Black-tailed Godwit, and that all three species were unaffected by the presence of footpaths and buildings. Final models also revealed relationships for all three species between the frequency of field use and sward height and the degree of field enclosure.

Case study 5

Further improvements to the approach may also be made by quantifying the level of use of features such as footpaths and roads. Klein et al. (1995) looked at how disturbance affected the distribution of waterbirds in a nature reserve in Florida by identifying threshold distances at which waterbirds avoided a drive through the reserve at different visitation levels (using contingency tables and chi-squared tests) while also assessing seasonal changes. Migrant ducks (American Wigeon Anas americana, Eurasian Teal, Northern Pintail A. acuta and Northern Shoveler A. clypeata) remained more than 80 m from the drive even at low visitation levels, though more so when they had first arrived in early winter, whereas waders were displaced less far and at intermediate visitation levels, and herons, egrets, Brown Pelicans Pelecanus occidentalis and Anhingas Anhinga anhinga often remained close to the drive during high visitation levels. In contrast to the previous studies, no attempt was made to control (even indirectly) for variation in habitat quality with distance from the drive, although the variation in the strength of response of individual species to visitation levels suggested a direct link between the distributional patterns seen and disturbance.


  1. Top of page
  2. Abstract
  5. Acknowledgments

Landscape approaches, as described in the case studies above, aim to investigate the spatial effects of human disturbance by demonstrating that bird numbers or densities are depressed or their behaviour altered in proximity to areas used by humans. However, confidence that disturbance is a factor in the patterns observed in such studies may potentially be limited as behavioural responses (notably movements from sources of disturbance) are not usually directly observed. To be able to infer that disturbance is a cause of the spatial variation in bird abundance or behaviour observed it is therefore important that other factors are considered or other corroborative evidence presented.

This is particularly the case in heterogeneous habitats such as estuaries, where variation in factors such as substrate type, water depth and food supply may considerably affect waterbird distributions (Kelsey & Hassall 1989, Nehls & Tiedemann 1993, Velazquez & Navarro 1993, Yates et al. 1993). In such systems the approach would not be applicable unless some attempt were made to account for other causes of variation, and even if this is done it should be noted that some factors may be spatially autocorrelated. Spatial variation in food supply was partially, but indirectly, taken into account in models in the second case study (Burton et al. 2002a) by distinguishing between intertidal count sections that bordered the low tide mark and those that did not and, in the third case study (Burton et al. 2002b), by including the count section itself as an explanatory variable. However, it is clear that such models would be very much improved by more direct measurement of variables influencing food availability, as demonstrated in the study of grassland field use by waders by Milsom et al. (1998), or preferably the food resources themselves. A study of the effects of disturbance on habitat use in Black-tailed Godwits (Gill et al. 2001a), for example, found no effect of the presence of footpaths or marinas on the numbers of birds supported on adjacent intertidal areas once bivalve food supply had been taken into account. By contrast, McKinney et al. (2006), studying waterbirds wintering in Narragansett Bay, Rhode Island, USA, found that landscape characteristics, including the extent of surrounding residential development, explained a significant degree of the variation in species abundance and richness, having allowed for habitat characteristics such as prey density and shoreline configuration.

The use of a landscape approach may also be improved by quantifying the level of use of features such as footpaths and roads. By demonstrating that waterbird species distributed themselves further from drives when visitation levels were high, Klein et al. (1995) provided much stronger evidence that the avoidance of areas close to drives was due to disturbance rather than other coincident factors. Reijnen et al. (1996) similarly demonstrated how reductions in the densities of breeding grassland waterbirds adjacent to roads were positively related to the level of traffic (see also similar studies by van der Zande et al. 1980, Reijnen et al. 1995).

Temporal changes in response, such as found by Burton et al. (2002a) between weekdays and weekends, when the pressure from visitor numbers and recreational activities is likely to be higher, may provide clearer evidence that observed distributional patterns are the result of disturbance. Evans and Warrington (1997) and Evans and Day (2002) similarly showed that wildfowl may redistribute within or between inland waterbodies at weekends in response to recreational or hunting activities. Better evidence still may be obtained where potential new sources of disturbance are introduced to an area or existing sources of disturbance removed and corresponding changes in species’ distributions are observed. In the third case study (Burton et al. 2002b), count sections were only classified as ‘disturbed’ sections during periods of active construction work and thus the ‘disturbance’ effects reported are unlikely to have been the result of differences in food resources across the study area. The approach indeed may be best and most confidently applied in experimental, before-after-control-impact (BACI) studies (Stewart-Oaten et al. 1986) where human presence in the landscape is switched between sites temporally. Madsen (1998a, 1998b) clearly demonstrated how hunting disturbance can influence the distributions of wildfowl by establishing experimental areas where hunting was manipulated prior to the establishment of permanent refuges. Quarry goose and duck species redistributed according to the location of hunting-free areas, whereas non-quarry species did not.

Through the range of studies described, the essential aim is the same, i.e. to demonstrate spatial variation in bird abundance or behaviour in relation to landscape features associated with human activity. This aim is limited and the degree to which it can be concluded rather than merely implied that disturbance is a causal factor of the spatial patterns observed is, as discussed above, first and foremost dependent on the extent to which other explanatory variables are considered. The approach is thus least applicable in the most heterogeneous environments where many factors, some perhaps spatially autocorrelated, may explain variation. Greatest confidence in the validity of conclusions will be gained where studies are able to show (ideally where possible by experimental manipulation) that species’ distributions or behaviour vary temporally in line with the levels of human use of the features examined.

Landscape approaches do not aim to provide any evidence as to whether disturbance may impact individual fitness and thus the size of local populations. Nevertheless, provided that other factors affecting spatial variation in bird abundance or behaviour are taken into account, the use of a landscape approach can offer a framework for more detailed investigation of the mechanisms of disturbance. In a previous study, for example, Gill et al. (1996) used information concerning the reduction in Pink-footed Goose densities with decreasing distance to roads to provide the basis for estimating the numbers of birds that could have been supported by the food resources not exploited as a result of disturbance associated with roads. In particular, by revealing how bird abundance or behaviour may vary spatially in response to landscape features, landscape-based studies may highlight specific sources of disturbance where further investigation would help inform management options. The approach may also provide information on the relative tolerances of different species to the disturbance associated with human landscape features. In the second case study, the relative distances to which models suggested species were affected by footpaths proved a good match to previously reported information on the distances at which species take flight in response to walkers (Burton et al. 2002a). Similar information on the tolerances of different species or species groups to disturbance from ecotourism was obtained by Klein et al. (1995). Site management already employs the use of buffer zones around existing ‘core wildlife areas’, based on the tolerances of key species to human disturbance (Fox & Madsen 1997, Rodgers & Smith 1997, Blumstein et al. 2003). Mathers et al. (2000), for instance, studying Eurasian Wigeon A. penelope and Light-bellied Brent Geese B. b. hrota at Strangford Lough, Northern Ireland, suggested the use of a buffer of 250 m in order to exceed the tolerance distance of the more sensitive of the two species at that site.

It should be noted, however, that the extent to which disturbance does actually affect the distribution of birds within a site and thus their apparent tolerance to different factors will vary both according to the availability of alternative resources and the birds’ own state. Thus, if birds are under stress, e.g. during periods of cold winter weather, and alternative resources are restricted, they may be less easily disturbed than at other times (McGowan et al. 2002, Stillman & Goss-Custard 2002) and as a result show less avoidance of areas associated with human activity, such as construction sites or footpaths, even if the costs for fitness are high (Gill et al. 2001b). As Klein et al. (1995) noted, the information on species’ avoidance responses obtained by landscape approaches would be enhanced by study of the movements of individually marked birds, to show where birds go when disturbed and whether they are able to habituate to human activities.

With increasing numbers of visitors to coastal areas, there is a clear dichotomy for managers of protected areas between the need to protect birds and other wildlife from human disturbance, while still allowing public access to scenic sites and indeed the wildlife the sites support (e.g. Kenchington 1989). Although it may not reflect population-level responses, as noted by Blumstein et al. (2005), the information needed at a site level is often ‘not whether or not a species is negatively affected by human disturbance, but what the probabilities are of any given species using a particular site within a protected area under different levels of disturbance.’ Thus, studies which use landscape approaches to explore the spatial avoidance of key species to sources of disturbance may offer a valuable first step towards site management.


  1. Top of page
  2. Abstract
  5. Acknowledgments

My thanks to Alex Banks, Jenny Gill, Tony Fox and Mark Rehfisch for their useful and constructive comments, which considerably improved the paper.


  1. Top of page
  2. Abstract
  5. Acknowledgments
  • Beale, C.M. & Monaghan, P. 2004. Behavioural responses to human disturbance: a matter of choice? Anim. Behav. 68: 10651069.
  • Blumstein, D.T., Anthony, L.L., Harcourt, R.G. & Ross, G. 2003. Testing a key assumption of wildlife buffer zones: is flight initiation distance a species-specific trait? Biol. Conserv. 110: 97100.
  • Blumstein, D.T., Fernández-Juricic, E., Zollner, P.A. & Garity, S.C. 2005. Inter-specific variation in avian responses to human disturbance. J. Appl. Ecol. 42: 943953.
  • Burger, J. 1981. The effect of human activity on birds at a coastal bay. Biol. Conserv. 21: 231241.
  • Burton, N.H.K. in press. The impact of the Cardiff Bay barrage on wintering waterbirds. In Boene, G.C., Galbraith, C.A. & Stroud, D.A. (eds) Waterbirds around the World: 805. Edinburgh: The Stationery Office.
  • Burton, N.H.K., Armitage, M.J.S., Musgrove, A.J. & Rehfisch, M.M. 2002a. Impacts of man-made landscape features on numbers of estuarine waterbirds at low tide. Environ. Manage. 30: 857864.
  • Burton, N.H.K., Rehfisch, M.M. & Clark, N.A. 2002b. Impacts of disturbance from construction work on the densities and feeding behavior of waterbirds using the intertidal mudflats of Cardiff Bay, UK. Environ. Manage. 30: 865871.
  • Burton, N.H.K., Rehfisch, M.M., Clark, N.A. & Dodd, S.G. 2006. Impacts of sudden winter habitat loss on the body condition and survival of Redshank Tringa totanus. J. Appl. Ecol. 43: 464473.
  • Collier, M.P., Banks, A.N., Austin, G.E., Girling, T., Hearn, R.D. & Musgrove, A.J. 2005. The Wetland Bird Survey 2003/04: Wildfowl and Wader Counts. Thetford: BTO/WWT/RSPB/JNCC.
  • Davidson, N.C. & Rothwell, P. (eds) 1993. Disturbance to waterfowl on estuaries. Wader Study Group Bull. 68: Special Issue.
  • Davidson, N.C., Laffoley, D.d’A., Doody, J.P., Way, L.S., Key, R., Drake, C.M., Pienkowski, M.W., Mitchell, R. & Duff, K.L. 1991. Nature Conservation and Estuaries in Great Britain. Peterborough: NCC.
  • Evans, D.M. & Day, K.R. 2002. Hunting disturbance on a large shallow lake: the effectiveness of waterfowl refuges. Ibis 144: 28.
  • Evans, D.M. & Warrington, S. 1997. The effects of recreational disturbance on wintering waterbirds on a mature gravel pit lake near London. Int. J. Environ. Stud. 53: 167182.
  • Fitzpatrick, S. & Bouchez, B. 1998. Effects of recreational disturbance on the foraging behaviour of waders on a rocky beach. Bird Study 45: 157171.
  • Fox, A.D. & Madsen, J. 1997. Behavioural and distributional effects of hunting disturbance on waterbirds in Europe: implications for refuge design. J. Appl. Ecol. 34: 113.
  • Gill, J.A., Sutherland, W.J. & Watkinson, A.R. 1996. A method to quantify the effects of human disturbance on animal populations. J. Appl. Ecol. 33: 786792.
  • Gill, J.A., Norris, K. & Sutherland, W.J. 2001a. The effects of disturbance on habitat use by black-tailed godwits Limosa limosa. J. Appl. Ecol. 38: 846856.
  • Gill, J.A., Norris, K. & Sutherland, W.J. 2001b. Why behavioural responses may not reflect the population consequences of human disturbance. Biol. Conserv. 97: 265268.
  • Goss-Custard, J.D., Triplet, P., Sueur, F. & West, A.D. 2006. Critical thresholds of disturbance by people and raptors in foraging wading birds. Biol. Conserv. 127: 8897.
  • Hill, D.A., Hockin, D., Price, D., Tucker, G., Morris, R. & Treweek, J. 1997. Bird disturbance: improving the quality of disturbance research. J. Appl. Ecol. 34: 275288.
  • Keller, V.E. 1991. The effect of disturbance from roads on the distribution of feeding sites of geese (Anser brachyrhynchus, A. anser), wintering in north-east Scotland. Ardea 79: 229232.
  • Kelsey, M.G. & Hassall, M. 1989. Patch selection by Dunlin on a heterogeneous mudflat. Ornis Scand. 20: 250254.
  • Kenchington, R.A. 1989. Tourism in the Galápagos islands: the dilemma of conservation. Environ. Conserv. 16: 227236.
  • Kershaw, M. & Cranswick, P.A. 2003. Numbers of wintering waterbirds in Great Britain, 1994/95–1998/99. I. Wildfowl and selected waterbirds. Biol. Conserv. 111: 91104.
  • Klein, M.L., Humphrey, S.R. & Percival, F. 1995. Effects of ecotourism on distribution of waterbirds in a wildlife refuge. Conserv. Biol. 9: 14541465.
  • Lafferty, K.D. 2001a. Birds at a southern California beach: seasonality, habitat use and disturbance by human activity. Biodivers. Conserv. 10: 19491962.
  • Lafferty, K.D. 2001b. Disturbance to wintering Western Snowy Plovers. Biol. Conserv. 101: 315325.
  • Madsen, J. 1998a. Experimental refuges for migratory waterfowl in Danish wetlands. I. Baseline assessment of the disturbance effects of recreational activities. J. Appl. Ecol. 35: 386397.
  • Madsen, J. 1998b. Experimental refuges for migratory waterfowl in Danish wetlands. II. Tests of hunting disturbance effects. J. Appl. Ecol. 35: 398417.
  • Mathers, R.G., Watson, S., Stone, R. & Montgomery, W.I. 2000. A study of the impact of human disturbance on Wigeon Anas penelope and Brent Geese Branta bernicla hrota on an Irish Sea Loch. Wildfowl 51: 6781.
  • McGowan, A., Cresswell, W. & Ruxton, G.D. 2002. The effects of daily weather variation on foraging and responsiveness to disturbance in overwintering Red Knot Calidris canutus. Ardea 90: 229237.
  • McKinney, R.A., McWilliams, S.R. & Charpentier, M.A. 2006. Waterfowl–habitat associations during winter in an urban North Atlantic estuary. Biol. Conserv. 132: 239249.
  • Milsom, T.P., Ennis, D.C., Haskell, D.J., Langton, S.D. & McKay, H.V. 1998. Design of grassland feeding areas for waders during winter: the relative importance of sward, landscape factors and human disturbance. Biol. Conserv. 84: 119129.
  • Nehls, G. & Tiedemann, R. 1993. What determines the densities of feeding birds on tidal flats: a case-study on Dunlin Calidris alpina, in the Wadden Sea. Neth. J. Sea Res. 31: 375384.
  • Rehfisch, M.M., Austin, G.E., Armitage, M.J.S., Atkinson, P.W., Holloway, S.J., Musgrove, A.J. & Pollitt, M.S. 2003. Numbers of wintering waterbirds in Great Britain and the Isle of Man (1994/1995–1998/1999). II. Coastal waders (Charadrii). Biol. Conserv. 112: 329341.
  • Reijnen, R., Foppen, R., Ter Braak, C. & Thissen, J. 1995. The effects of car traffic on breeding bird populations in woodland. III. Reduction of density in relation to the proximity of main roads. J. Appl. Ecol. 32: 187202.
  • Reijnen, R., Foppen, R. & Meeuwsen, H. 1996. The effects of traffic on the density of breeding birds in Dutch agricultural grasslands. Biol. Conserv. 75: 255260.
  • Roberts, G. & Evans, P.R. 1993. Responses of foraging Sanderlings to human approaches. Behaviour 126: 2943.
  • Rodgers, J.A. & Smith, H.T. 1997. Buffer zone distances to protect foraging and loafing waterbirds from human disturbance in Florida. Wildl. Soc. Bull. 25: 139145.
  • Smit, C.J. & Visser, G.J.M. 1993. Effects of disturbance on shorebirds: a summary of existing knowledge from the Dutch Wadden Sea and Delta area. Wader Study Group Bull. 68: 619.
  • Stewart-Oaten, A., Murdoch, W.M. & Parker, K.R. 1986. Environmental impact assessment: ‘pseudoreplication’ in time? Ecology 67: 929940.
  • Stillman, R.A. & Goss-Custard, J.D. 2002. Seasonal changes in the response of Oystercatchers Haematopus ostralegus to human disturbance. J. Avian Biol. 33: 358365.
  • Stillman, R.A., West, A.D., Caldow, R.W.G. & Durell, S.E.A. le V. dit. 2007. Using individual behaviour to predict the responses of coastal bird populations to disturbance. Ibis 149 (Suppl. 1): 7381.
  • Stroud, D.A., Chambers, D., Cook, S., Buxton, N., Fraser, B., Clement, P., Lewis, P., McLean, I., Baker, H. & Whitehead, S. (eds) 2001. The UK SPA Network: its Scope and Content. Peterborough: JNCC.
  • Thomas, K., Kvitek, R.G. & Bretz, C. 2003. Effects of human activity on the foraging behavior of Sanderlings Calidris alba. Biol. Conserv. 109: 6771.
  • Triplet, P. & Gembarski, S. 2001. Évolution de la distance d’évitement d’un humain par l’Huîtrier pie Haematopus ostralegus en recherche alimentaire. Alauda 69: 543544.
  • Triplet, P., Bacquet, S., Morand, M.-E. & Lahilaire, L. 1998. La distance d’envol, un indicateur de dérangements: l’exemple de quelques espèces d’oiseaux en milieu estuarien. Alauda 66: 199206.
  • Triplet, P., Sueur, F. & Urban, M. 2001. Distance d’envol de quelques espèces d’oiseaux d’eau hivernant en Baie de Somme. Alauda 69: 457458.
  • Velazquez, C.R. & Navarro, R.A. 1993. The influence of water depth and sediment type on the foraging behaviour of Whimbrels. J. Field Ornithol. 64: 149157.
  • West, A.D., Goss-Custard, J.D., Stillman, R.A., Caldow, R.W.G., Durell, S.E.A. le V. dit & McGrorty, S. 2002. Predicting the impacts of disturbance on shorebird mortality using a behaviour-based model. Biol. Conserv. 106: 319328.
  • Whitfield, D.P. 2003. Redshank Tringa totanus flocking behaviour, distance from cover and vulnerability to Sparrowhawk Accipiter nisus predation. J. Avian Biol. 34: 163169.
  • Yates, M.G., Goss-Custard, J.D., McGrorty, S., Lakhani, K.H., Durell, S.E.A. le V. dit, Clarke, R.T., Rispin, W.E., Moy, I. & Yates, T. 1993. Sediment characteristics, invertebrate densities and shorebird densities on the inner banks of the Wash. J. Appl. Ecol. 30: 599614.
  • Van Der Zande, A.N., Ter Keurs, W.J. & Van Der Weijden, W.J. 1980. The impact of roads on the nesting densities of four bird species in an open field habitat – evidence of a long-distance effect. Biol. Conserv. 18: 299321.