Divergent selection on feather pecking behaviour in laying hens (Gallus gallus domesticus)

A selection experiment was initiated in 1996 in which selection for (HP line) and against (LP line) feather pecking was performed. The foundation stock was a White Leghorn layer strain established in 1970 and maintained since then as a random bred control line at the Institute. Six hatches were produced over three generations. At the age of 68 weeks (generation 0, 1996), 35 weeks (generation 1, 1997), 30 weeks (generation 2, 1998), and 27 weeks (generation 3, 1999) female birds were transferred to observation pens and their feather pecking behaviour was recorded. In each generation, 30 females and 8 males were selected from approximately 200 females and 60 males. The selection criterion was breeding value estimated by animal model on the trait 'number of bouts of feather pecking per bird per hour'.Feather pecking behaviour in adult hens was significantly higher in HP than in LP. In generation 2 the following was recorded: 3.10 versus 1.37 bouts per bird per hour (P<0.01), 7.04 versus 3.58 pecks per bird per hour (P<0.05) and the proportion of hens recorded feather pecking in the 180min observation period was 67 versus 56% (P<0.05). In generation 3 the following was recorded: 4.56 versus 0.63 bouts per bird per hour (P<0.001), 13.9 versus 2.51 pecks per bird per hour (P<0.001) and the proportion of hens recorded feather pecking in the 180min observation period was 75 versus 49% (P<0.001).In generation 3, plumage condition was better in LP on neck, breast, back, wings and tail, as well as overall (P<0.001). Body weight did not differ between lines in generation 2, but in generation 3, HP hens were on average heavier than LP hens at the age of 27 weeks (1435g versus 1371g, P<0.001).     

Effect of manipulating feathers of laying hens on the incidence of feather pecking and cannibalism

Feather pecking is a problem in commercial laying hens, particularly in loose-housing systems, where many hens can be affected by only a few feather peckers. In addition, feather pecking can become an even larger problem if it spreads throughout the flock. There are several possible ways that feather pecking may spread. The simplest way is that one hen may damage the feathers of a hen, and another hen may find the damaged feathers an attractive pecking target. The aim of this experiment was to determine if damaged feathers were feather-pecked more than undamaged feathers on the same body area, and to determine whether some types of feather-body area manipulations were preferred over others as pecking stimuli. Manipulations involved damaging the feathers on the rump, tail or belly of different hens, with two or three levels of severity of manipulation at each body area. Sixteen groups of 11 Lohmann Brown hens between 26 and 28 weeks were observed with the recipient, the feather pecker and the body area that was pecked all being recorded. The feather pecks were classified separately as either gentle or severe. Damaged feathers received significantly more severe feather pecks than undamaged feathers. There were also more gentle feather pecks to damaged feathers, although this did not reach statistical significance. The feather-body area manipulations that received the greatest number of severe feather pecks were the tail feathers when they were cut very short, the rump feathers when they were trimmed, and the rump when feathers were removed. These results support the suggestion that feather pecking does indeed spread through flocks by damaged feathers becoming an attractive target for feather-pecking behaviour. An unexpected result of performing the feather manipulations was an outbreak of cannibalism in half of the experimental groups. Even though there was no visible damage to the skin of the hens after having the feathers manipulated, 13 of the 16 attacked hens were wounded on the part of the body where the feathers had been damaged in some way.     

Reaction to frustration in high and low feather pecking lines of laying hens from commercial or semi-natural rearing conditions

The effect of rearing conditions on feather pecking and reaction to frustration was studied in two lines of laying hens. From commercial rearing conditions (large group, no mother hen), seven birds from a high feather pecking line (HC birds) and eight birds from a low feather pecking line (LC birds) were used. From semi-natural rearing conditions (small group, mother hen present) seven birds from the high feather pecking line (HN birds) were used. Feather pecking behaviour of HC, LC, and HN groups was recorded for 30 min. After that, each bird was food deprived and trained to peck a key for a food reward in a Skinnerbox. After training, each bird was subjected to a frustration session in a Skinnerbox, where the feeder was covered with Perspex. Three HC birds showed severe feather pecking, compared with one HN bird and zero LC birds. Differences in reaction to frustration were found between birds from different lines, but not in birds from different rearing conditions. LC birds tended to put their head in the feeder more frequently than HC birds over all sessions. Although limited, this study indicates that rearing conditions influence feather pecking, but not reaction to frustration.     

Effect of rearing density on pecking behaviour and plumage condition of laying hens in two types of aviary

The objective of this experiment was to investigate the effect of rearing density on pecking behaviour and plumage during rearing and throughout the laying period in aviaries. Chicks were reared on sand at high (H; 13 m⁻²) or low (L; 6.5 m⁻²) density, in four rearing pens of 390 chicks and eight pens of 195 chicks, respectively, each pen measuring 30 m². Proportions of chicks per pen performing various types of pecking behaviour were recorded by scan sampling during 16 observation bouts in each rearing pen at 6 weeks of age and during 24 observation bouts at 12 weeks. Individual body weights and plumage condition were recorded. Later, these pullets were housed at 17 hens m⁻² in Tiered Wire Floor (TWF; 3 H and 3 L pens of 275 hens) and Laco-Volétage (2 H and 2 L pens of 275 hens) aviaries. At 35 weeks, two samples of eight hens from each aviary pen were observed for pecking behaviour in a test pen. Throughout the laying period, additional records were collected on pecking behaviour, body weight, plumage condition, egg production, and mortality. The L birds had better plumage condition at 6 weeks of age and throughout the laying period. These birds also ground pecked more frequently than H birds during rearing and the laying period. At 12 weeks, L birds feather pecked less than H birds, but no relationship was found between rearing density and feather-pecking behaviour during the laying period. Although TWF hens feather pecked more frequently than Volétage hens, there was no interaction between rearing density and type of aviary for the various pecking behaviours.     

Reaction to frustration in high and low feather pecking laying hens

Reaction to frustration of high (HFP) and low feather pecking (LFP) laying hens was investigated. From a HFP- and a LFP-line five birds with a HFP- and five birds with a LFP-phenotype were selected. Birds from the HFP-line were expected to show more key pecking and covered feeder pecking during frustration than birds from the LFP-line. When a bunch of feathers was presented, birds with a HFP-phenotype were expected to redirect their pecks at the bunch. Birds were trained to peck a key for a food reward in an automated Skinnerbox and subjected to two sessions: a control session, where food was available, and a frustration session, where the feeder was covered with Perspex. These two sessions were repeated in the presence of a bunch of feathers. Unexpectedly, birds from the LFP-line had a stronger reaction to frustration than birds from the HFP-line, expressed in pecking behaviour. When a bunch of feathers was offered, birds with a HFP-phenotype did not show more bunch pecking during frustration than birds with a LFP-phenotype.     

Influence of prior exposure to wood shavings on feather pecking, dustbathing and foraging in adult laying hens

It has been proposed that chicks acquire substrate preferences during an early 'sensitive' period. If a suitable substrate is absent during this period birds may develop alternative preferences for pecking at feathers. The aim of this study was to examine whether early substrate exposure has durable effects on the subsequent behaviour of adult hens. The effects of duration of substrate exposure, substrate change, age at exposure and time since exposure on adult bird behaviour were examined. From days 1 to 210, 144 laying strain birds were housed in pairs in pens with wire floors. The floors were replaced with solid floors covered in wood shavings at different ages and for different durations by allocation to 1 of 12 treatments. Adult birds that had never experienced shavings performed significantly more feather pecking than birds in any other treatment group. Thus, exposure to shavings, even for the minimum exposure duration of 10 days, was protective. However, current substrate was of great importance and adult birds housed on shavings performed significantly more ground pecking and less feather pecking than birds on wire, regardless of previous experience. From day 211 all hens were given shavings or straw, presented alternately for five 24h sessions over 10 consecutive days. Birds foraged on both substrates and their foraging behaviour was not influenced by previous experience. Dustbathing occurred primarily on shavings and was significantly influenced by the age at which birds had previously been exposed to shavings. Dustbathing on shavings was fairly constant throughout the 10-day test period in all groups, suggesting that relatively stable preferences had developed. A secondary 'sensitive period' for the formation of adult dustbathing substrate preference may have superseded the early 'imprinting' process. However, adult behaviour was generally flexible and strongly influenced by current substrate.     

Selection method and early-life history affect behavioural development, feather pecking and cannibalism in laying hens: A review

The aim of this review is to discuss the effects of selection method and early-life history on the behavioural development of laying hens. Especially in larger groups, laying hens often develop damaging behaviours, such as feather pecking and cannibalism, leading to impaired animal welfare. We hypothesise that the propensity to develop feather pecking and cannibalism is affected by a bird's genetic background and by its early-life history. The genetic background can be influenced by genetic selection. Laying hens are traditionally selected on individual performance, which may lead to co-selection of feather pecking and cannibalism. For hens kept in small groups, it has recently been demonstrated that a novel group selection method, focusing on group performance, can help to reduce cannibalism. However, the biological background behind the success of group selection is unknown. It is also not known whether these results from small groups can be translated to larger groups of laying hens. Regarding early-life history, laying, brooding and rearing conditions have been shown to have major effects on behavioural development and on feather pecking and cannibalism. The presence of a hen during rearing has been shown to improve foraging- and social behaviour, to decrease feather pecking and to decrease fearfulness in chicks. Applying group selection and rearing laying hens in a more natural environment may be key factors in solving the problems caused by feather pecking and cannibalism, especially if the promising results of group selection from small groups in experimental settings can be translated to large-group housing systems.     

Reducing feather pecking when raising laying hen chicks in aviary systems

Aviary systems for laying hens offer several advantages over battery cages. However, pecking the feathers of conspecifics remains a serious problem that negatively affects the welfare of the birds as well as the economy of a farm. From experimental studies with small groups, it has been shown that feather pecking and foraging behaviour are related and that both behaviour are influenced by early access to litter substrate. We, therefore, hypothesised, that feather pecking in aviaries can be reduced with an adequate management in the first 2 weeks of life.Each of seven pens on six commercial poultry farms, was divided into two identical compartments (matched pair design). In one of the compartments (experimental compartment) chicks were reared for the first 2 weeks of life with access to litter (wood shavings, in one case with additional straw), while the chicks in the other compartment (control) were kept on a plastic grid. Thereafter, all chicks had unrestricted access to litter and there were no differences between the two compartments neither in housing conditions nor in management procedures.Chicks in the experimental compartments spent significantly more time foraging (week 5), showed significantly less feather pecking (weeks 5 and 14) and significantly fewer birds had damaged tail feathers (weeks 5 and 14).The study demonstrates that in aviaries, under commercial conditions, early access to litter substrate has a significant effect on the development of feather pecking. In order to reduce feather pecking and to increase foraging behaviour, it is recommended that laying hen chicks raised in aviary systems do get access to litter from day 1 on.     

Comparisons of damaging feather pecking and time budgets in male and female turkeys of a traditional breed and a genetically selected male line

Feather and skin pecking leading to feather loss and tissue damage is a welfare problem in commercial turkeys. A factorial experiment was designed to compare a line of unselected turkeys that previously did not exhibit this behaviour with a genetically selected male line with a propensity for damaging pecking. We also compared the time budgets of male and female turkeys from the two lines at 3, 6 and 9 weeks of age and at different times of the day. Damaging feather pecking occurred in 32% of male and 15% of female male line turkeys but was not observed in the traditional line. Time budgets of males and females were similar. Preening increased and resting, feeding and general pecking declined with age. Resting was higher in the afternoon than in the morning and male line turkeys were observed resting more often than traditional turkeys at 3 and 6 weeks. Traditional turkeys displayed more gentle feather pecks than male line poults at 9 weeks. Male turkeys of both lines showed more strong feather pecks and pulls at 3 weeks whereas female turkeys showed more at 9 weeks of age. The frequency of strong feather pulls recorded electronically increased with age but was not related to mortality. It is suggested that damaging feather pecking in turkeys may be the result of vigorous investigative pecking.     

Is there social transmission of feather pecking in groups of laying hen chicks?

Feather pecking is an abnormal behaviour where laying hens peck the feathers of conspecifics, damaging the plumage or even injuring the skin. If it occurs in a flock, more and more birds show it within a short period of time. A possible mechanism is social transmission. Several studies have shown that laying hen chicks, Gallus gallus domesticus, are able to modify their own behaviour when observing the behaviour of other chicks, for example, when feeding and foraging. As there is good experimental evidence that feather pecking originates from foraging behaviour, we hypothesized that feather pecking could also be socially transmitted. To test this, we reared 16 groups of 30 chicks. After week 4, the birds were regrouped into 16 groups of 20 chicks into each of which we introduced either five chicks that showed high frequencies of feather pecking or, as controls, five chicks that had not developed feather pecking. We then determined the feather-pecking rate and the frequency of foraging, dustbathing, feeding, drinking, preening, moving, standing and resting of all birds in a group. Data from the introduced birds were analysed separately and excluded from the group data. Chicks in groups with introduced feather-pecking chicks had a significantly higher feather-pecking rate than chicks in the control groups. In addition, birds in groups with introduced feather peckers showed significantly lower foraging frequencies than those in the control groups, although the housing conditions were identical and there were no differences in either the number or the quality of the stimuli relevant to foraging behaviour. The study therefore suggests that feather pecking is socially transmitted in groups of laying hen chicks. Copyright 2000 The Association for the Study of Animal Behaviour.     

Group size and perching behaviour in young domestic fowl

To test the hypothesis that young domestic fowl perform less perch-related antipredator behaviour with increasing group size, White Leghorn pullets were reared in four replicate groups of 15, 30, 60 and 120 at a constant density of 5 birds/m(2). Each pen contained perches 20, 40 and 60cm above the ground. Perch space per bird per perch level was the same for all groups. It was predicted that, with increasing group size, domestic fowl would (1) spend less time on perches (i.e. more time down on the floor); (2) be less vigilant while perching; (3) spend relatively more time preening down on the floor. As predicted, the proportion of 3- to 18-week-old birds roosting on perches during scans throughout the photoperiod decreased with increasing group size, from 41+/-1.7% in groups of 15 birds to 33+/-1.6% in groups of 120 birds. This effect was due to reduced use of the lower perches; use of the highest perches was high at all group sizes. The proportion of birds vigilant on the highest perches of those present on that perch level decreased with increasing group size. The proportion of birds engaged in the vulnerable activity of preening down on the floor increased with group size. The frequency of transitions between floor and perches was not affected by group size but birds received more disturbances from other birds when on the top perch level in the larger groups. Thus, the decline in vigilance on the top perch level with increasing group size was not due to reduced disturbance from other birds. In conclusion, despite domestication and protection from non-human predators, changes in the use of perches by young domestic fowl with increasing group size were consistent with the antipredator hypothesis.     

Use of perches and nestboxes by laying hens in relation to social status, based on examination of consistency of ranking orders and frequency of interaction

Four groups of 15-19 adult ISA Brown hens were studied in pens to assess the relationship between social status and use of perches and nestboxes. This was to test the hypothesis that subordinate hens use these resources more by day, for avoiding dominants, but that dominants use perches more at night, for roosting. The experiment consisted of a 5-week pre-treatment period, when no perches were present, and a 4-week treatment period, when each group was tested with different perch treatments (No, Low, Medium, High). All groups were observed systematically in each week, when all interactions of three types (aggressive peck, non-aggressive peck, approach/avoidance) in a group were recorded by noting the instigator and recipient (from numbered wing tags) onto a matrix. Proportions of time that each bird spent using perches and nestboxes, by day and at night, were also recorded. The results indicate that social status of individual laying hens is relatively stable across time and can be based reliably on counts of either aggressive pecks or approach/avoidances, but not non-aggressive pecks. Aggressive pecks were the most frequent type of interaction observed, and were reduced by the presence of perches. Use of nestboxes, but not perches, was greater at night than by day. There were weak tendencies for perches, and to a lesser extent nestboxes, to be used more by lower ranking birds by day, but not at night. There was some evidence of increased use of these resources by higher ranking birds at night. It is concluded that provision of perches reduces bird density on the floor (where nearly all interactions occurred), allows subordinates a means of avoiding dominants by day, reduces frequency of agonistic interactions, and should thus benefit laying hen welfare.     

The presence of extreme feather peckers in groups of laying hens

Feather pecking is a serious economic and welfare problem in laying hens. Feather damage occurs mainly through severe feather pecking (SFP). Selection experiments have proved that this behavior is heritable and lines have been divergently selected for high (HFP) and low feather pecking (LFP). The number of bouts of SFP per hen follows a Poisson distribution with a maximum nearby 0. A few studies indicate that the distribution within flocks is not homogenous but contains sub-groups of birds showing extremely high levels of feather pecking (EFP). It was the aim of the current study to re-analyze data on SFP of lines selected for HFP/LFP and their F2 cross so as to uncover hidden sub-populations of EFP birds. Data of seven selection generations of HFP and LFP selection lines as well as their F2 cross have been used. We fitted a two-component mixture of Poisson distributions in order to separate the sub-group of EFP from the remaining birds. HFP and LFP lines differed mainly in mean bouts per bird. The proportion of EFP was only marginal in the LFP as compared with the HFP and the F2 population. Selection for LFP did not result in total elimination of EFP. The presence of even small proportions of EFP may play an important role in initiating outbreaks of feather pecking in large flocks. Further studies on feather pecking should pay special attention to the occurrence of EFP sub-groups.     

Heart rate variability in domestic chicken lines genetically selected on feather pecking behavior

Domestic chicken lines of the White Leghorn type differing in their level of feather pecking were developed by divergent genetic selection specifically on feather pecking behavior. We determined parameters of heart rate variability to elucidate the relative activation of the sympathetic and parasympathetic nervous systems during rest and stressful situations. A total of 48 hens were tested in 8 batches. Segments of 2 min were extracted from electrocardiograms recorded by radio-transmitter implants, before (basal undisturbed conditions) and during physical restraint and a social test. Under basal conditions mean distance between R-waves were shorter in the low and high lines compared to the control. During physical restraint, stress reactions [reduced root of the mean squares of successive differences (RMSSD), reduced high frequency (HF), high low frequency (LF/HF) and low vagal-sympathetic effect (VSE) compared to basal levels] were significant in all lines. During the physical restraint the high feather pecking (HFP) line reacted significantly stronger than control (CON) and low feather pecking (LFP) line. During social test the LFP line reacted different than the other two lines. Seemingly birds from LFP conceived the social test as less stressful than birds from the CON and HFP lines. From this it follows that (1) physical restraint generally induced higher stress reactions than the social test and (2) genetic selection for higher levels of feather pecking increased the autonomic nervous system reaction to physical restraint whereas selection against feather pecking has reduced the response to increased social contact.     

Agonistic behaviour in domestic hens: the influence of housing method and group size

The incidence of agonistic behaviour was recorded over 1 year in laying hens of two strains, housed either in battery cages or in deep-litter pens in groups of either three or six birds. Aggressive head-pecking was more frequent in pens than in cages and in groups of six than in groups of three. Threats were more frequent in pens and in groups of six light-hybrids. Other pecking at the plumage was more common in cages in groups of six and in the light-hybrid than in the medium-hybrid strain. Observations showed that hens occupy a considerable area when engaging in threat displays: this suggests that the physical restraint of crowding may play an important part in limiting this behaviour. Two reasons for the reduction of aggressive head pecking in caged birds were suggested: firstly, inhibition of the subordinate hens' behaviour because they were constantly in the sphere of influence of the dominant bird and secondly, because agonistic encounters may be triggered only by the entry of other birds into an individual's 'personal space' and not by continuous proximity.     

Feather eating and its associations with plumage damage and feathers on the floor in commercial farms of laying hens

Feather eating has been associated with feather pecking, which continues to pose economic and welfare problems in egg production. Knowledge on feather eating is limited and studies of feather eating in commercial flocks of laying hens have not been performed previously. Therefore, the main objective was to investigate feather eating and its association with plumage damage and floor feather characteristics in commercial flocks of layers in barn and organic production systems. The study was performed in 13 flocks of barn layers and 17 flocks of organic layers. Each flock was visited at around 32 and 62 weeks of age. During both visits, the plumage condition was assessed and the density of floor feathers recorded. In week 62, droppings and floor feathers were collected. Droppings were examined for presence of feather content, whereas length, downiness and pecking damage were recorded for each floor feather. In week 62, a higher prevalence of hens with poor plumage condition was found in barn (22.2%) compared with organic production systems (7.4%; P<0.001), but the prevalence of droppings with feather content did not differ between the two production systems (8.5% in barn v. 4.3% in organic; P=0.99). Our hypothesis about a positive correlation between feather eating and plumage damage was not supported as no correlation was found between the prevalence of poor plumage condition and the prevalence of droppings with feather content. However, the prevalence of pecking damaged floor feathers was positively correlated both with prevalence of droppings with feather content (P<0.05) and poor plumage condition (P<0.01), indicating a possible association between feather eating and feather pecking. In conclusion, it was confirmed that feather eating occurs on-farm, but feather eating was only found to be positively correlated to the number of floor feathers with pecking damage and not as expected to the prevalence of plumage damage. More research is needed into the sources from where feathers are selected for ingestion, that is, whether they are picked from the floor litter, plucked directly from other hens or dislodged during preening of own feathers.     

Apomorphine pecking in the pigeon.

Pecking activity elicited by apomorphine was studied quantitatively in intact and thalamic pigeons. While apomorphine pecking is easily observed in the intact pigeon, it is absent in the acute thalamic animal. I reappears, however, in the chronic preparation simultaneously with the recovery of spontaneous pecking. Apomorphine pecking is described by satiation and increased by fasting. Reticular stimulation produces reversible blockade of both apomorphine and spontaneous pecking with the same parameters, without increase in heart rate. Apomorphine pecking can be observed only in the presence of visual contrast, and is always aimed at the contrast points or edges. A decrease in contrast produces a decrease in the number of pecks delivered to the contrast points and an increase in those missing the target, while the total number of pecks delivered in a unit time is unaffected. A lowering of the background luminance is followed by a decrease in the total number of pecks, which is mostly due to a reduction of those aimed at the contrast points. All these findings are discussed and it is concluded that: i) apomorphine exerts an excitatory action (direct or indirect) on the hypothalamic feeding centers; hence apomorphine pecking can be considered as a pharmacologically motivated behavior; ii) visual contrast exerts a dual action on apomorphine pecking, namely it acts both as triggering stimulus and goal object; iii) the decrease in apomorphine pecking induced by lowering the ambient light intensity is probably due to a decrease in the level of the general arousal.     

A comparison of the behaviour and production of laying hens in experimental and conventional battery cages

There is some experimental evidence that the behaviour of laying hens in conventional battery cages is restricted. The absence of litter in particular may account in part for the higher level of feather pecking in cages as opposed to pens, and for the tendency of one strain to develop stereotyped pacing, indicative of frustration, in the pre-laying period.In the present small-scale study an experimental cage was tested for its effects on the behaviour, production, and feather and foot damage of groups of six birds per cage. The experimental cages, when compared with conventional types, had an horizontal floor, were extended in height, had food and water provided at two levels, two perches and nesting boxes containing litter. Egg collection was automated; eggs and litter travelled to a collecting point and the litter was recycled to the cages.Forty-eight birds of a White Leghorn laying strain (Shaver 288) were housed at six per cage in a bank of experimental cages. Forty-eight sister birds were grouped in sixes in conventional caging for comparison. Details are given of the space allowances in each cage on various criteria, since a simple floor area comparison was not relevant.Observations lasted over a 6-month period from point-of-lay. The experimentally caged birds laid nearly all their eggs in the nest boxes, and showed more movement, reflex activity, preening and sitting but less feeding, drinking, pecking, pushing and stereotyped behaviour. The specific variables causing these behavioural differences are speculated upon, although in this type of study they cannot be identified with certainty.The experimental birds showed less feather and foot damage compared with controls in conventional cages. Egg production in the two groups was similar but samples of eggs from the experimental cages were of a higher mean weight.As yet no economic assessment of the viability of the experimental cage has been undertaken, but reference is made to comparable studies where this has been attempted.On welfare grounds there is some evidence that the experimental cage is nearer to the criteria of an “ideal” cage (Hughes, 1973) than conventional cages.     

An Experimental Analysis of Spatial Position Effects on Foraging and Vigilance in Brown-Headed Cowbird flocks

Several observational studies have found that the costs and benefits of social foraging vary as a function of spatial position in the group. However, it is difficult to make mechanistic inferences because several confounding factors, such as food deprivation levels, food availability, neighbor distance, and group size can mask or amplify spatial position effects. We attempted to address experimentally the effect of spatial position on foraging and vigilance in a group, controlling for many confounding factors. We used enclosures that restricted physical but not visual interactions between brown-headed cowbirds and manipulated spatial position, flock size, and neighbor distance. Pecking rate (number of pecks per trial duration) was not related with position, but instantaneous pecking rate (number of pecks per foraging bout duration) was higher at the edge. The proportion of time spent head-up (scanning and food-handling) was also higher at the edge. For pecking rate and proportion of time spent scanning, changes in neighbor distance influenced the behavior of edge birds to a lesser extent than central birds. These results suggest that cowbirds at the edge perceived greater predation risk and that during the limited foraging time available, edge birds tried to compensate by foraging at a faster rate.