THE SOCIAL BEHAVIOUR OF THE WHITE WAGTAIL MOTACILLA ALBA ALBA WINTERING IN ISRAEL

White Wagtails Motacilla alba wintering in Israel are partly territorial, mostly around human habitations, and partly live in flocks around temporary food sources. Individual birds may spend part of the season (or the day) in the territory and the other part with a flock. Experiments with artificial distribution of food, in a natural habitat, brought about a change from flocking to territorial behaviour. Preliminary observations suggest that in the natural situation the pattern of food distribution may be the proximate factor which regulates the birds' behaviour, by determining whether they have to fight for their food. Pairs are formed on many territories, and may last for long or short periods. Pair formation is initiated by females, who when seeking food appease the territorial males and are able to stay with them on their territories. Females also manifest territorial behaviour. Although pairing in winter territories is similar, in the behaviour involved, to sexual pairing, it is very unlikely that winter pairing continues into, or influences, pairing for breeding. It is suggested that the function of winter pair formation is that it allows two birds to exploit one territory, and that the main advantage is to the female which is the subordinate bird of the pair. This kind of pair-formation may be analogous to non-breeding group territories reported in some other birds.     

Winter territoriality of Mute Swans Cygnus olor

There was considerable variation in winter territoriality among Mute Swan pairs around Oxford, though most pairs maintained their territories for at least half the winter. Variation was associated with temperature and with territory quality pairs left their territories most commonly during the coldest months of the winter, and pairs on good territories, with abundant vegetation and accessible pasture. stayed longer on territory than those on poor territories with sparse vegetation and little pasture. As in other species, exclusive territorial defence was maintained at intermediate levels of resource availability, and flocks were able to settle on a few particularly good territories. Circumstantial evidence suggested that winter territories may be maintained not only to provide an adequate food supply for the owners but also to ensure access to a brerding site by preventing other individuals from taking over the area.     

The wintering of migrant Dunnocks Prunella modularis in two Mediterranean habitats after fire

Migrant Dunnocks Prunella modularis are common winter visitors to the Mediterranean area. In a burnt forest, Dunnocks were present from October to April. They occupied definite individual home ranges of 0.18 ha on average (n = 83), at a density of 30 birds/10 ha in the fifth winter and 18 birds/10 ha in the sixth winter following a fire. Birds used visible posts and uttered ‘tseep’ calls often. Counter-calling was particularly frequent in the winter with the highest population density. In a burnt maquis, Dunnocks showed return rates (15%) and median recapture distances (62.5 m) not significantly different from those of territorial wintering Robins Erithacus rubecula. We suggest that Dunnock ranges can be regarded as ‘feeding territories’, promoted by an abundant, predictible and renewable food supply of rock-rose Cistus spp. seeds, characteristic of early Mediterranean successions. Ownership might be proclaimed by ‘tseep’ calls, recalling the behaviour of female Dunnocks during prebreeding territorial conflicts. Further research using colour-ringed birds may confirm Dunnock winter territoriality, exceptional for a migrant and mostly granivorous passerine.     

FOOD RESOURCES, TERRITORY DENSITY AND REPRODUCTIVE SUCCESS OF AN ISLAND SILVEREYE POPULATION ZOSTEROPS LATERALIS

We examined the relationships between food resources, territory density and some breeding parameters (individual reproductive success and parental age) of a Capricorn Silvereye population during two years' detailed study on Heron Island, Great Barrier Reef. The territorial behaviour of the Silvereyes included dawn song. aggression and distinctive vocalizations. During both years, about 85% of pairs occupied territories although the population sizes differed. The dispersion pattern of territories was not regular and was correlated with the density of figs and human food scraps. Nestlings were fed more figs as they grew, and the parents foraged in fig trees outside their territories. However, only about 60%, of territorial pairs fledged young. Pairs which fledged most young were older, fed more insects to their nestlings, and nested in areas containing more fruiting fig trees per territory. We conclude that food resources were in short supply, and that access to fig trees provided breeding birds with a quick energy source while they searched for insect protein for the young. The data are consistent with the predictions of the ‘sufficient resource‘ hypothesis of the adaptive significance of territoriality but not with its assumption that the principal benefit is the food within the territory.     

Winter territoriality and its implications for the breeding ecology of White-throated Dippers Cinclus cinclus

Capsule: Pairs of White-throated Dippers Cinclus cinclus which defended winter territories bred earlier than non-territorial individuals, but there was no difference in reproductive success.

Aims: The effect of winter territoriality on breeding ecology has rarely been studied in resident birds. We carried out a preliminary investigation of whether winter territorial behaviour and territory size affect the timing of reproduction, breeding territory size and reproductive success in a riverine bird, the White-throated Dipper.

Methods: We monitored an individually marked population of White-throated Dippers in the UK. Wintering individuals were classified as either territorial or ‘floaters’ according to their patterns of occurrence and behaviour, and their nesting attempts were closely monitored in the subsequent months. Winter and breeding territory sizes were measured by gently ‘pushing’ birds along the river and recording the point at which they turned back.

Results: All birds defending winter territories did so in pairs, but some individuals changed partners before breeding. Territorial pairs that were together throughout the study laid eggs significantly earlier than pairs containing floaters and those comprising territorial birds that changed partners. However, there were no significant differences in clutch size, nestling mass or the number of chicks fledged. There was no relationship between winter territory length and lay date or any measure of reproductive success, although sample sizes were small. Winter territories were found to be significantly shorter than breeding territories.

Conclusion: Winter territoriality may be advantageous because breeding earlier increases the likelihood that pairs will raise a second brood, but further study is needed. Territories are shorter in winter as altitudinal migrants from upland streams increase population density on rivers, but this may also reflect seasonal changes in nutritional and energetic demands.     

Temporal shifts in the pair-bond dynamics of New Zealand robins (

Winter is a challenging time for temperate insectivorous songbirds, due to colder temperatures, reduced prey activity and shorter diurnal foraging times. For species that are non-migratory, territorial and monogamous, winter conditions may result in within-pair competition. However, little is known about how monogamous pairs coexist on their winter territories. We investigated temporal patterns in male?female interactions of the New Zealand robin (Petroica australis to better understand mechanisms of coexistence during winter. Previous work has shown that male robins are physically dominant over females and maintain priority access to food year-round. We quantified female behaviour throughout the 2008 non-breeding season to better understand how females coexist with physically dominant males on winter territories. Results showed that pairs rarely forage in close proximity in autumn and winter, suggesting females avoid males at this time of year. Males and females begin to spend more time foraging together as winter turns to spring. During this winter?spring transitional period, females steal large amounts of food hoarded by males. These results indicate that male and female New Zealand robins use different behavioural mechanisms to coexist on their winter territories. While males are dominant physically, females show a seasonally variable strategy where they avoid males in autumn and winter, and then steal male-made caches from early spring until the onset of inter-pair cooperation and the breeding season.     

ON THE DEFINITIONS AND FUNCTIONS OF DOMINANCE AND TERRITORIALITY

1. Dominance/subordinance is a relationship between two individuals in which one defers to the other in contest situations. Each such relationship represents an adaptive compromise for each individual in which the benefits and costs of giving in or not giving in are compared. Familiar associates in groups or neighbours on nearby territories may develop relatively stable dominant-subordinate relationships based on individual recognition. Although the aggressive aspects of dominance are usually emphasized, the less conspicuous actions of the subordinate individual are actually more important in maintaining a stable relationship. 2. In evolutionary terms, dominance essentially equals priority of access to resources in short supply. Usually the subordinate, who would probably lose in combat anyway, is better off to bide its time until better able to compete at another time or another place. Both individuals save time, energy, and the risk of injury by recognizing and abiding by an established dominant-subordinate relationship. 3. Dominance can be either absolute or predictably reversible in different locations or at different times. Of the various forms of dominance behaviour, rank hierarchies and territoriality represent the two extremes of absolute and relative dominance, respectively. A dominance hierarchy is the sum total of the adaptive compromises made between individuals in an aggregation or organized group. Many animals seem to be capable of both absolute and relative dominance, and within species-specific limits the balance may shift toward one or the other. High density, or a decrease in available resources, favours a shift from relative to absolute dominance. Some species may exhibit both simultaneously. Social mammals may have intra-group hierarchies and reciprocal territoriality between groups, while the males of lek species may exhibit ‘polarized territoriality’ by defending small individual territories, with the most dominant males holding the central territories where most of the mating takes place. 4. Territoriality is a form of space-related dominance. Most biologists agree that its most important function is to provide the territory holder with an assured supply of critical resources. Territoriality is selected for only when the individual's genetic fitness is increased because its increased access to resources outweighs the time, energy, and injury costs of territorial behaviour. 5. Territoriality was first defined narrowly as an area from which conspecifics are excluded by overt defence or advertisement. The definition has been variously expanded to include all more or less exclusive areas without regard to possible defence, and finally to include all areas in which the owner is dominant. I define territory as a fixed portion of an individual's or group's range in which it has priority of access to one or more critical resources over others who have priority elsewhere or at another time. This priority of access must be achieved through social interaction. 6. My definition excludes dominance over individual space and moving resources, and includes areas of exclusive use maintained by mutual avoidance. It differs from most other definitions in its explicit recognition of time as a territorial parameter and its rejection of exclusivity and overt defence as necessary components of territorial behaviour. There is an indivisible continuum of degrees of trespass onto territories, and functionally it is priority of access to resources that is important rather than exclusive occupancy. 7. There is a similarly indivisible continuum in the intensity of behaviour needed to achieve priority of access to resources. Deciding whether or not an exclusive area is defended leads to the pointless exercise of trying to decide which cues indicating the owner's presence are conspicuous enough to merit being called defence. Concentrating on overt defence emphasizes the aggressive aspects of territorial behaviour rather than the equally or more important submissive aspects such as passive avoidance.     

Population density,resource patterning,and territoriality in the Everglades pygmy sunfish

The means by which pygmy sunfish compete for food are influenced by the density of the population and the dispersion of the prey as well as by the sex and dominance status of the individual. At all densities when the prey were predictably located in a central clump, males established territories. When prey were dispersed randomly in both the high and low density population, males abandoned territorial behaviour and, like females, swam freely about the aquaria. Only in the intermediate density populations did the males maintain territories and continue to defend resources. Development of a simple cost-benefit model shows that male territorial behaviour is governed to a large extent by economic considerations. Despite these overall patterns, differences in competitive strategies were observed within populations. Dominant individuals tended to possess territories nearest the central patch of prey, and use the most intense and physiologically exhausting displays. Only under the most stressful conditions did they acquire significantly more food than subordinates, and even then these benefits were not translated into increased growth, largely because dominant fish engaged in disproportionately more energy-consuming contests.     

Territorial behaviour in the Western Atlantic Grey seal (Halichoerus grypus)

Virtually all published accounts of territorial behaviour in the Grey seal apply to breeding males only. Long-range field studies reveal, however, that lactating cows also maintain territories for a period of several weeks. During the summer months, adults and sub-adults have specific resting places to which they return each day at low water and which they defend against other individuals of their own species.     

THE ORIGINS OF ADAPTIVE INTERSPECIFIC TERRITORIALISM

1. In order to understand fully the evolution of a behavioural trait one must not only consider whether it is adaptive in its present environment but also whether it originated as an adaptation to existing selective forces or as a fortuitous consequence of selection for a different role in other environments (i.e., as a pre-adaptation) or of selection for different traits (e.g., as a pleiotropic effect). In this paper interspecific territorialism is examined in species of humming-birds, sun-birds, tropical reef fishes, stingless bees, stomatopods, crayfish, and limpets as a means of determining its adaptiveness and its origins. 2. Humming-birds form complex assemblages with species sorted out among the available resources. Dominant species establish feeding territories where flowers provide sufficient nectar. A few large, dominant species, usually uncommon, are marauders on others' territories. Subordinate species establish territories where flowers are more dispersed or produce less nectar, or they fly a circuit from nectar source to nectar source when flowers are even more dispersed, a foraging pattern called ‘traplining’, or they steal nectar from the territorial species by being inconspicuous while foraging. Two species, Amazilia saucerottei and Selasphorus sasin, subordinate in one-to-one encounters, are able to take over rich resources by establishing several small territories within a territory of a dominant and forcing it to forage elsewhere. 3. Among humming-birds, territorial individuals attacked not only subordinate competitors but marauding humming-birds and some insects, which stayed in the territory and foraged at will, and seemingly inappropriate targets, such as non-competitors. This suggests that the stimulus for aggression is ‘any flying organism near the food resources’, regardless of its appearance. The behaviour rather than the identity of the intruder is the stimulus. 4. Sun-birds resemble humming-birds to the extent that dominants establish territories on rich nectar sources and subordinates establish territories on less rich nectar sources or steal from the territories of dominants. The diversity of foraging patterns is not so great as in humming-birds, perhaps because so few species of sun-birds have been studied. However, the advantage of territorialism has been measured in the sun-bird Nectarinia reichenowi. Individuals with territories lose much less nectar to competitors than do those without territories. 5. Field work on three species of tropical reef fishes involved a single aggressive species whose individuals attacked a wide range of species intruding on their territories. The stimulus for aggression in Pomacentrus jenkinsi seemed to be an “object moving through [its] territory”. As suggested for humming-birds, the stimulus is the behaviour rather than the identity of the intruder. 6. The relationships found in stingless bees, stomatopods, crayfish, and limpets are simpler. The dominant and subordinate species divide the resources in their habitat, the dominants' aggression preventing the subordinates from using resources that were otherwise available to them. 7. A general pattern emerges. Mutual interspecific territorialism occurs between species that (i) have different geographic ranges, (ii) occupy different habitats, or (iii) use different resources within the same habitat. Examples of two species holding separate territories on the same resources within the same habitat are rare and occur when the dominant species is rare relative to the available resources. These observations are contrary to the usual view that interspecific territorialism is an adaptation that permits co-existence of potential competitors within the same habitat. 8. Interspecific territorialism is sometimes adaptive and sometimes maladaptive, depending upon the species and the situation. 9. The general pattern of occurrence of the behaviour and the general nature of the stimulus for aggression, i.e., the behaviour rather than the identity of the intruder, suggest that interspecific territoriality is a fortuitous consequence of selection for intraspecific territorialism, the latter being not only an adaptation to the presence of conspecific competitors but a pre-adaptation to the presence of competitors of other species, should they occur.     

Bill morphology reflects female independence from male parental help

The study of territorial polygyny in birds has been influential in the development of the theory of social mating systems. Alternative female mating options have been studied within the framework of the polygyny-threshold model and later as the outcome of conflicts of interest between individuals. However, little attention has been given to variations between individual females, and how this affects their mating behaviour. Here, we test the hypothesis that some females are better adapted to raise nestlings without male assistance, and thus to mate polygynously. Specifically, we investigate whether intraspecific variation in female bill morphology is related to mating behaviour. This hypothesis is derived from earlier studies showing that, in both intra- and interspecific comparisons, uniparental care by females is correlated with the catching of larger prey items than when both parents provision the young. Using the polygynous dusky warbler (Phylloscopus fuscatus) as a model species, we found that, in accordance with our prediction, females with deep bills were more likely to mate as a secondary female. Moreover, regardless of mating status, females with deep bills settled in territories with more food and they received less male assistance in feeding their offspring. We argue that females with stronger bills are better adapted to exploit the abundance of large food items in rich territories and thus to raise young on their own. Our results demonstrate the importance of studying variations between individual females, and provide evidence for an extended version of the 'constrained-female hypothesis'. As bill depth is a highly heritable trait, our study strongly suggests that variation in female mating behaviour is not only related to ecological factors and female condition (as shown elsewhere) but also to heritable morphological traits.     

Territorial and foraging behaviour of two corallivorous butterflyfishes

The territorial and foraging behaviour of two Red Sea butterflyfish species was studied at three sites in the Gulf of Aqaba. Chaetodon austricaus , a generalist corallivore that exploited evenly distributed food resources, maintained exclusive pair territories. Individuals fed by grazing for brief periods on a coral colony before moving to the next. In contrast, C. trifascialis , a specialist corallivore that exploited patchily distributed food resources, demonstrated considerable variation in territorial behaviour that ranged from the defence of exclusive solitary territories, to the shared use of a large home range. Individuals showed highly variable feeding behaviour, from grazing analogous to C. austriacus , to continual use of a single large coral colony. The results of this study demonstrate how the use of space by individuals depends critically upon the distribution of key food resources. In addition, the results demonstrate that foraging behaviour of territorial butterflyfish can be quite plastic, and adapt to local conditions.     

Butterflyfish social behaviour,with special reference to the incidence of territoriality: a review

Synopsis Butterflyfishes (Chaetodontidae) are among the best studied of coral reef fishes. Feeding ecology and some aspects of behaviour have been firmly established. However, spacing behaviour remains controversial. Two major studies made in the 1970s concluded that the majority of species were not territorial. We suggest that these and other studies which have concluded that territories are not held have generally suffered from short observation periods, and have not mapped the ranges occupied by individuals. Further, low frequencies of agonistic behaviour have been interpreted as evidence of non-territoriality. By contrast, studies which have proven territoriality have had long observation periods and have mapped ranges. These have shown that territories are usually maintained with very little overt aggression. Spacing behaviour and feeding behaviour are clearly linked, with territoriality common among benthic-feeding species, especially obligate corallivores. Species with broad dietary flexibility tend to have flexible social systems, while plankton feeders are usually gregarious. The widespread occurrence of monogamy in butterflyfishes appears linked to territoriality, the majority of territorial species identified to date occurring predominantly as pairs. Data currently available suggest that this is because pair defence of the territory is more efficient than by individuals. However, several alternative hypotheses for the evolution of monogamy based on spawning constraints and predation risk cannot yet be ruled out.     

A comparison of the song and territorial behaviour of a long-distance migrant, the Marsh Warbler Acrocephalus palustris, in summer and winter

Colour-ringed populations of Marsh Warblers Acrocephalus palustris on breeding and wintering grounds were compared with respect to song and territorial behaviour. Song and territorial defence had a short duration on the breeding grounds, peaking during the period of mate acquisition and mate-guarding. Some males later showed polyterritorial behaviour. Wintering Marsh Warblers sang and showed defensive behaviour for a longer period which included moult. This behaviour could be described in terms of defence of a short-term resource (a fertile female) in the summer and long-term defence of a reliable food resource in the winter.     

Resource and habitat sharing by the stoplight parrotfish,Sparisoma viride, a Caribbean reef herbivore

Synopsis We compare the vertical distribution, substrate preferences, grazing behaviour and social interactions of the stoplight parrotfish,Sparisoma viride, with that of other scarids at a fringing reef off Bonaire (Netherlands Antilles). Earlier reports thatS. viride only displays territorial behaviour against conspecifics are confirmed by the non-aggressive nature of this parrotfish and the low time expenditure on interactions with other herbivores. The vertical distribution ofS. viride is largely identical to that of several other scarids, acanthurids and pomacentrids, whose ranges may completely coincide withS. viride territories. Comparison of the substrate use of the five most common scarids inside a singleS. viride territory yields no evidence of food partitioning. We suggest that the lack of interspecific territorial behaviour inS. viride is explained by the inability to economically defend a territory against all potential food competitors. Factors that may favour territory sharing between herbivores are fine-scale resource partitioning and shared defence, both of which would reduce the costs of territorial life. However, more detailed investigation of herbivore food selection is required before definite conclusions can be drawn.     

EXPERIMENTS ON THE LIMITATION OF BIRD NUMBERS BY TERRITORIAL BEHAVIOUR

(1) This paper examines removal and other experimental studies on the role of territorial behaviour in the limitation of bird densities. Experimental design is discussed, as are the types of conclusions that can be drawn. (2) Experiments have been conducted on more than 40 species from a wide range of taxonomic groups. Most provided evidence for density limitation, probably mediated by territorial behaviour, and for the existence of a non-territorial sector able to take territories when territory owners were removed or when additional habitat was made available. Experiments in the breeding season indicated that some surplus birds, although younger on average then territorial birds, were sexually mature and able to breed when given the chance. In some species, replacement birds bred less well than other territorial birds, but in other species no difference was apparent between the two groups. (3) Other experiments indicated that, while density was limited by territorial behaviour in good habitats, this was not the case in poor habitats, and that some individuals would move from poor to good habitat when territory owners were removed from good habitat. (4) In some species, non-territorial birds of breeding age were present in the population despite the existence of vacant territorial sites, while in others replacements were observed on good territories but not on poor ones. The implications were that site quality influenced whether settlement, defence and breeding occurred, and that some individuals had more stringent site requirements than others. In some seabirds' sites in the centre of a colony were more attractive than sites on the edges. (5) Many land-bird species that have been studied show two peaks of territorial activity each year, in autumn and in spring, and a limitation on breeding density can occur at either or both seasons, depending on conditions. (6) The fact that densities of many bird species fluctuate greatly from year to year is not inconsistent with the limitation of density by territorial behaviour. Several mechanisms are apparent through which density might be limited by territorialism at different levels in different years, so that surplus non-territorial birds are present in years of low territorial density, as well as in years of high territorial density. (7) Experiments have shown that density is limited by the presence of territorial birds, in some species at different levels in different areas or years. The next step is to find whether density is regulated by the presence of territorial birds over a period of years in a density-dependent manner. For this, observational data are required to find whether the proportion of birds that is excluded by territorial behaviour each year varies with the total number available for settlement. Such studies can be made only on species in which non-territorial birds can be counted accurately, as well as territorial ones.     

Different wintering strategies of two Palearctic migrants in West Africa – a consequence of foraging strategies?

The wintering strategies of Pied Flycatchers Ficedula hypoleuca and Willow Warblers Phylloscopus trochilus in their West African winter quarters were compared. Pied Flycatchers arrived early in the season (September) and stayed in the study area throughout the winter. They were territorial and showed a high return rate. Intraspecific relationships were mostly expressed by territorial behaviour. Interspecific relations seemed to be unimportant. Willow Warblers arrived relatively late (November) and were absent from the area for some weeks in January and February, a behaviour which was interpreted as itinerancy. Willow Warblers were non-territorial and never returned to a site. Willow Warblers usually moved through the area in monospecific or mixed-species flocks. Habitat and microhabitat choice of these species were similar but in feeding ecology they differed by the higher diversity of feeding substrates and feeding techniques of Pied Flycatchers. The differences in the winter strategies are explained by the ability of Pied Flycatchers to defend a territory because of their diversity in foraging behaviour, whereas Willow Warblers are more specialized and are therefore forced to be more mobile to find their patchily distributed food.     

Territorial aggression does not feed back on testosterone in a multiple-brooded songbird species with breeding and non-breeding season territoriality,the European stonechat

Testosterone mediates reproductive behaviours in male vertebrates. For example, breeding season territoriality depends on testosterone in many species of birds and in some, territorial interactions feed back on testosterone concentrations. However, the degree to which territorial behaviour and testosterone are associated differs even between species with seemingly similar life histories, especially between species that also defend territories outside the breeding season. Here, we investigate the link between territorial behaviour and testosterone in European stonechats. Previous studies found that territorial aggression in stonechats depends on testosterone in a breeding, but not in a non-breeding context. We investigated whether stonechats show a rise in testosterone during simulated territorial intrusions (STI) during the breeding season. Post-capture testosterone concentrations of males caught after an STI were not higher than those of males caught in a control situation regardless of breeding stage. However, most of the males would have been able to mount a testosterone response because the same individuals that did not increase testosterone during the STI showed a substantial increase in testosterone after injections of gonadotropin-releasing hormone (GnRH). GnRH-induced and post-capture testosterone concentrations were positively correlated and both decreased with successive breeding stages. Further, territory owners with a short latency to attack the decoy expressed higher post-capture testosterone concentrations than males with a longer latency to attack the decoy. Thus, there is no evidence for behavioural feedback on testosterone concentrations during male-male interactions in stonechats. In combination with previous studies our data suggest that testosterone functions as an on/off switch of high intensity territorial aggression during the breeding season in stonechats. The among-species variation in the androgen control of territorial behaviour may be only partly a result of environmental differences. Instead, potential differences in how territoriality evolved in different species may have influenced whether and how a reproductive hormone such as testosterone was co-opted into the mechanistic control of territorial behaviour.     

Shadow competition in wild juvenile sea-trout

Shadow competition occurs in a group of sit-and-wait predators when those closer to a source of mobile prey reduce the feeding success of those further from the prey source. It was examined in territorial juvenile sea trout Salmo trutta in a small stream. The fry formed groups of two to six fish with adjacent territories and a social hierarchy within each group. It was hypothesized that: (i) as group size increased, the mean number of prey eaten per fish within a group decreased and the variability in prey consumption between fish increased; (ii) prey consumption by individual fish decreased with increasing distance from the food source; (iii) group size increased as the mean water velocity immediately upstream from a group, and hence potential drifting food, increased. Five groups of fry were fed on small shrimps released upstream from each group at a rate of one every 15 s over a 10 min period, this procedure being repeated over 5 days to provide five replicates per group. Experiments were performed three times in 1967, 1969 and 1974 to provide information on 45 groups of fry. The first and third hypotheses were supported, but the second was only partially supported. In 1967 and 1969, territory size and shrimp consumption by individual fry decreased with increasing distance from the food source. This also occurred in 1974, except during a critical period for survival when fry density was exceptionally high with large numbers of sea trout lacking territories. This resulted in sea trout fry with the largest territories eating fewer shrimps than those with medium-sized territories because they spent more time defending their territories against sea trout lacking territories. This study is the first to demonstrate shadow competition in a vertebrate species, but has also shown that territorial defence may modify the consequences of shadow competition when densities are high and there is strong competition for the acquisition of a territory.     

THE WINTER FEEDING ECOLOGY OF THE REDSHANK TRINGA TOTANUS

The aim of the work was to find out how the Redshank which over-winter on the Ythan estuary, Aberdeenshire, adapt their feeding activities to the short daylength in winter. To achieve this, the feeding behaviour and daily routine in winter were compared with those in autumn and spring. Corophium volutator was the main prey on the estuary during the day. However, the temperature of the mud greatly affected the diet in the two areas where the feeding behaviour was studied in detail. At temperatures above 6°C, most of the biomass ingested consisted of Corophium. However, in one study area in one winter, Macoma balthica was taken more frequently at lower than at high temperatures. In the other area, Nereis diversicolor was taken more frequently at low than at high temperatures. These changes in diet appeared related to changes in the behaviour of the prey affecting their availability and not due to the birds changing their preference at low temperatures. The ingestion rate was not affected by the change in diet in the first area but decreased at low temperatures in the second. The sizes of Macoma and Nereis taken by the birds did not vary seasonally. The sizes of Corophium taken decreased in winter as a result of a reduction in the size of those present in the substrate. It was concluded that there was no evidence of Redshank increasing their ingestion rate in winter to compensate for the short daylength. The proportion of the time spent feeding on the estuary in daylight was greater in winter than in spring. Redshank continued feeding at high water in the surrounding fields and on the estuary at night during the winter but not in either autumn or spring. During the winter they obtained less than 50% of their daily food requirements from the estuary in daylight, mainly because of the short daylength. Consequently, they had to collect the balance at night and at high water. Seasonal changes in the numbers of Redshank were recorded. After an autumn decrease in numbers, no decrease in numbers could be detected during the winter. These findings are discussed in relation to the possible difficulties for Redshank in collecting their daily food requirements during the winter. The impact of the birds on their main estuarine prey Corophium is also discussed.