Thesis (Ph.D.) - University of St Andrews, February 2010.

Redshank selected low tide feeding sites on the Ythan Estuary in a manner consistent with the "Ideal Free" model (Fretwell and Lucas 1970). Predictions derived from the "Despotic" (Fretwell 1972) model were contravened. Redshank feeding rates were independent of the density in the mud of their major prey species, Corophium volutator, the size of prey available however, was crucial in determining food intake. High densities of Corophium too small to be taken had an apparent confusing effect, resulting in increased search costs. The shift in the Redshank distribution towards the mouth of the estuary in late autumn, previously associated with decreasing daylength (Joffe 1978), was consistent with a change in the cue, from energy intake per unit time to energy intake per unit feeding cost, used by Redshank to select their lowtide feeding sites. Increase in the proportion of Redshank feeding at low tide, the switch from roosting to feeding in agricultural land at high tide, increase in the degree of fidelity to low tide sites and decrease in the amount of movement about the estuary at low tide all indicate an increase in the pressure on Redshank to feed efficiently. Although no significant turnover in the winter population was detected, a PASCAL simulation revealed that turnover of 3 0% of the population size, or higher, would have gone undetected. Significant turnover was noted during the autumn period. Lower quality Redshank, e.g. juveniles and light birds, were predominant among the autumn emigrants. Severe weather conditions in 1981/82 resulted in a 22% reduction in the winter population, almost entirely through increased mortality. Juveniles and, despite the "Ideal Free" distribution, Redshank on the lower section of the estuary, the least preferred in autumn, were apparently more vulnerable to severe weather conditions.

In order to meet their energy budget animals must often increase their risk of predation, either through their choice of foraging location or by decreasing anti-predation behaviours, which are incompatible with foraging. I investigated the starvation-predation risk trade-off in redshanks overwintering in the area of the Firth of Forth in Scotland over different spatial scales. On a small spatial scale, where redshanks foraged in an area where risk of attack was high I investigated the role of competition for food and decreased individual vigilance within groups and how this related to predation risk, additionally I looked at the relative roles of individual variation in time spent exposed to risk and variation in anti-predation behaviours on individual survival time. On larger spatial scales of 100s of meters and over several kilometres, I considered how choice of overwintering site was affected by predation risk, profitability and population density. Time available to feed increased with increased group size, allowing redshanks to compensate for increased competition and allowing large groups to form, thus decreasing individual predation risk. Individuals that spent less time exposed to attacking predators survived for longer, however individuals constrained by cold weather to spend long periods exposed to risk could increase their survival through increased intake rates and vigilance. On an intermediate spatial scale redshanks selected overwintering sites based on profitability rather than risk, and only used less profitable site when population density was high. On a large spatial scale redshanks increased their use of less profitable sites in warmer weather, but did this without increasing their risk of predation. Overall this suggests that across most spatial scales redshanks can minimise their predation risk by their choice of foraging location, but when forced by weather conditions or competition to be exposed to attack, capture reducing behaviours also reduce predation risk.

Disturbance from engineering works is an increasing problem in terrestrial and marine ecosystems throughout the world. Many reported declines in population size, breeding success and body condition have been diagnosed as the result of anthropogenic disturbance, however little is known about the effect of long-term disturbance from large-scale engineering works. Understanding the mechanisms by which animals respond to anthropogenic activities is fundamental to explaining interactions, and resolving potential conflicts between humans and wildlife. This thesis focuses on the factors affecting the habitat use and foraging decisions in wintering shorebirds and wildfowl. The first half of this thesis considers the direct and indirect impacts on waterbirds of a major engineering project in central Scotland; construction of the new Clackmannanshire Bridge at Kincardine-on-Forth. For individual bird species in close proximity to the bridge site, round-the-clock construction work had consequences ranging from neutral to considerably negative. Cormorant Phalacrocorax carbo declined in the area, probably as a result of the disturbance of an important low tide roost. Redshank Tringa totanus, previously abundant in the prey-rich areas adjacent to the construction site, were displaced into poorer areas for most of the construction period; where they may also have suffered from increased interference competition and elevated risk from raptorial predators. Some positive effects of industrial development were also revealed; radio-transmitters combined with tilt-switch posture sensors indicate that Redshank were able to capitalise on the improved nocturnal visibility in areas around Grangemouth docks to assist with foraging and predator detection. Evidence is presented that birds switched foraging strategy (from sight to touch feeding) depending on ambient light levels; whereby artificial light was used in a similar manner to moonlight to assist with prey detection. Redshank also avoided riverine areas at night that were used frequently by day, probably in response to an elevated threat from nocturnal predators. As the predator landscape changes from day into night, birds adopt different strategies to minimise the risk from nocturnal predators. It is clearly important, therefore, that information on nocturnal distributions is available to inform decisions on site management, especially where anthropogenic activity continues throughout the diel cycle. Behavioural decisions were shown to vary widely within a species depending on individual state, metabolic demands and previous exposure to human disturbance. Prey resources were shown to change dramatically over the course of a winter. In response to this decline, the home range of Redshank contracted over a winter season. Similarly, animals responded less and took greater risks in response to experimental disturbance events later in the winter than earlier in the winter, and on days when the temperature was lower. This effect was strongest for individuals occupying heavily disturbed areas, which were possibly already compensating for lost feeding time and a negative energy balance. The results were consistent with the hypothesis that those individuals that respond most obviously to human disturbance were those least likely to suffer fitness consequences. This is the opposite from what is commonly assumed when behaviour is used as an index of disturbance impacts, most notably in the use of flush distance in the design of wildlife buffer zones. In conclusion, this study demonstrated various negative impacts of disturbance, including local displacement, due to construction activity on overwintering waterbirds. It also revealed two key, but poorly understood, phenomena relating to mechanisms for coping with anthropogenic disturbance: routine utilisation of artificial light to extend night-time feeding opportunities amongst Redshank and an adaptive flexibility in escape responses across a range of species under varying conditions of risk.

Conservation of intertidal habitats in the UK is vital in order to continue to support nationally and internationally important populations of non-breeding waterbirds. Historic reclamation for agriculture and industry has resulted in the loss and degradation of large areas of these intertidal habitats in estuaries and they continue to be threatened by sea-level rise. Managed realignment is one method which is increasingly being used to restore intertidal habitats. As managed realignment is a relatively new restoration technique, the extent to which knowledge of the biology of estuaries is applicable to managed realignment sites is unclear. Habitat restoration is often unsuccessful or incomplete, so a detailed knowledge of both the natural system and the characteristics of restored systems will usually be necessary to recreate fully-functional estuarine habitats. This thesis focuses on Nigg Bay Managed Realignment Site (Nigg Bay MRS), the first managed realignment site in Scotland, and follows the first four years of ecological development to gain an understanding of how breached realignment can be used to restore intertidal habitats to support non-breeding waterbirds. This thesis has six major aims: (i) to describe the development of saltmarsh, (ii) to describe the development of intertidal flat, (iii) to describe the colonisation by non-breeding waterbirds (iv) to determine how tidal cycle and weather affect patterns of waterbird use, (v) to determine which factors affect the spatial distribution of waders and finally (vi) to determine the patterns of use by individual birds. Four summers after the re-establishment of tidal conditions, almost all of the saltmarsh species recorded on the nearby saltmarsh had colonised Nigg Bay MRS, although recognisable communities had yet to establish. Three winters after the re- establishment of tidal conditions in Nigg Bay MRS, the sediments had a significantly smaller particle size and higher organic matter content compared to the fine sands of the adjacent intertidal flats. The intertidal invertebrate community also differed from the adjacent intertidal flats. Nigg Bay MRS attracted large numbers of non-breeding waterbirds and supported each of the most common wader and wildfowl species present in the wider estuary. Nigg Bay MRS performs a number of important functions for non-breeding waterbirds by: (i) providing a foraging and resting habitat when the tide is absent and intertidal sediments in Nigg Bay are exposed; (ii) providing a foraging resource as the tide passes over the intertidal sediments within the site once the intertidal flats in Nigg Bay are inundated; and (iii) providing a high tide roosting site. On days with low temperatures and high wind speeds, more waterbirds use Nigg Bay MRS, suggesting that it is likely to be providing sheltering benefits. Nigg Bay MRS also provides top-up feeding habitat. The factors that often influence the spatial distributions of waders in estuaries appear to be operating within Nigg Bay MRS. Wader densities are greater on the intertidal flats when they are accessible than on the saltmarsh. Wader densities are also greatest close to creeks and drainage channels, possibly due to higher invertebrate densities, more accessible prey or sheltering benefits. Colour-ringing and radio-tracking of Common Redshank established that Nigg Bay MRS has a subset of regular users, including both adults and juveniles, and the wader assemblage at night may differ from the assemblage during the day. These findings are discussed in terms of the implications for locating, designing and managing future managed realignment projects.

Previous research into bottom-up processes on saltmarshes has mainly focused on the influence of plant succession on herbivores. This study will present original research exploring the influence of bottom-up processes in a saltmarsh ecosystem between three trophic levels: Orchestia, redshanks, and sparrowhawks. Density dependence, may be the dominant top-down effect when higher numbers of sparrowhawks and redshanks are present, and may mask top-down and bottom-up trait effects which are constant. Bottom-up effects begin to emerge when cold conditions force redshanks from muddy creeks onto the saltmarsh to forage for Orchestia, because their primary prey, Corophium become less available. Larger flocks form and feeding on Orchestia requires them to balance a need to profit from the best available feeding patches and to be vigilant to sparrowhawk attack. Redshank vulnerability is compounded, because Orchestia hide in cold temperatures, so probing in the soil with their heads down makes them more vulnerable to sparrowhawk attack. Larger flocks may be able to exploit areas closer to sparrowhawk-concealing cover at the terrestrial boundary because they feel safer in greater numbers. Warmer temperatures make Orchestia more active which attracts redshanks, which can simultaneously feed and be vigilant because they peck and catch crawling and jumping Orchestia with their heads up. Consequently, increased flock size may temporarily depress Orchestia abundance, so that redshanks become spaced, leaving isolated individuals more vulnerable to attack. Therefore, it is a temperature-dependent bottom-up process which impacts upon both Orchestia and redshank behaviour, which then may influence the hunting success of sparrowhawks. Whether the characteristics of this saltmarsh ecosystem and the trophic dynamics can be compared to other examples is questionable. Saltmarshes probably differ in their topography and the way in which environmental conditions affect them that then defines which species are present and how these species interact.