Food‐Searching Behavior of Titmice in Patchy Environments

Abstract

The food—searching behavior of titmice (Paridae) was investigated in laboratory and field to determine how effective individual birds were at locating patchy sources of hidden food. A series of three experiments was performed using tame great tits, Parus major. In Experiment 1, individual birds were offered a choice among six densities of hidden food in discrete patches. The birds learned to concentrate their search strongly on the more dense patches, and five of the six birds preferred to search at the highest density. In Experiment 2, the food distribution was altered by interchanging patches of the highest and lowest density gradually. The birds initially found fewer prey per trial, but soon recovered to near their original performance by switching their search effort to the area containing the second highest prey density. Most birds failed to respond to prey appearing where they had previously been absent. Experiment 3 varied the size of prey items among four areas containing equal numbers of hidden prey. A group of six birds learned to search selectively in areas containing larger prey and, as a result, a greater total quantity of prey. In all experiments, there were marked differences in performance by individual birds. The experiments support Royama's (1970, 1971) hypothesis that great tits distribute their search effort in relation to spatial differences in the profitability of feeding sites. Field observations on a pair of great tits and a pair of blue tits, Parus caeruleus, showed that searching behavior of wild tits was also very selective in space, at least when the birds were feeding nestlings. Birds often returned to previous capture sites and were more likely to do so when they found prey there quickly. These observations suggest that the laboratory experiments have real relevance to the food searching of wild tits. The searching capacity of great tits is discussed; it is suggested that they can approach ideal responses in stable but patchy feeding environments. Some factors leading to deviations from ideal responses are discussed.