title: To graze or gorge: consistency and flexibility of individual foraging tactics in tits.
abstract: An individual's foraging behaviour and time allocated to feeding have direct consequences for its fitness. Despite much research on population-level foraging decisions, few studies have investigated individual differences in fine-scale daily foraging patterns among wild animals. Here, we explore the consistency and plasticity of feeding tactics of individual great tits (Parus major) and blue tits (Cyanistes caeruleus), using a grid of 65 automated feeding stations in a 385-ha woodland, during three winters. We use a principal component analysis to describe individual variation in six feeding parameters and examine how these differences covary with dominance-linked attributes (species, age and sex), the personality trait 'exploration behaviour', distance to territory and local competition intensity. Analysis of 933 086 feeder visits by 3134 individuals revealed that the majority of variation in the timing of feeding was explained by two principal components. PC1 ('binge-eating'), accounting for 38% of variation, captured temporal clustering of feeding, with high repeatability both within and between years (r range: 0·42-0·55). PC2 ('transience'), accounting for 27% of variance, described how much individuals used feeders and was also repeatable (r: 0·34-0·62). While exhibiting consistent individual differences, birds also showed flexibility in foraging patterns, binge-eating less and using feeders more when they experienced greater local competition. Individuals in behaviourally dominant states (great tits, males and adults) binged more than subordinate birds (blue tits, females and juveniles) when their territories were distant from feeding stations. Moreover, great tits and males used feeders more than blue tits and females respectively, while birds feeding further from their territory used feeders less than those feeding closer. 'Exploration behaviour' was unrelated to both measures of daily foraging behaviour. This study presents some of the first evidence that birds use consistent alternative foraging tactics at a fine temporal scale. Individuals are consistent in their tactics, and also adjust their foraging behaviour with changes in local competition. Hence, studies of foraging behaviour should consider the extent to which such individual-level variability in foraging behaviour is under selection.
authors.affiliation: Edward Grey Institute, Department of Zoology, University of Oxford, Oxford, OX1 3PS, UK.
authors.forename: Nicole D
authors.identifier: 0000-0001-6965-7471
authors.initials: ND
authors.lastname: Milligan
authors.affiliation: Edward Grey Institute, Department of Zoology, University of Oxford, Oxford, OX1 3PS, UK.|Department of Biology, Lund University, 22100, Lund, Sweden.
authors.forename: Reinder
authors.initials: R
authors.lastname: Radersma
authors.affiliation: Edward Grey Institute, Department of Zoology, University of Oxford, Oxford, OX1 3PS, UK.
authors.forename: Ella F
authors.initials: EF
authors.lastname: Cole
authors.affiliation: Edward Grey Institute, Department of Zoology, University of Oxford, Oxford, OX1 3PS, UK.
authors.forename: Ben C
authors.initials: BC
authors.lastname: Sheldon
mesh_terms: D000818:Animals; D004777:Environment; D005106:Exploratory Behavior; D005247:Feeding Behavior; D005260:Female; D008297:Male; D046109:Passeriformes; D012621:Seasons; D012919:Social Behavior
keywords: 
  PCA
  ; Paridae; competition; consistent differences; daily routine; diel activity; principal component analysis; repeatability