Feeding ecology and diet

The principal food of all species is adult and larval insects of a very wide variety and a great size range, from tiny midges and chironomid larvae to locusts and dragonflies. Insects taken include Diptera (especially Scatophagidae, Tipulidae, and Chironomidae), beetles (including cockroaches), grasshoppers and locusts, crickets, Hemiptera (including aphids), mantids, ants, termites, and wasps. Virtually the full range of insect prey is taken by the members of each genus, and some of the most popular food items are beetles (including weevils) and grasshoppers, plus dragonfly larvae, adult and larval Diptera, and Lepidoptera larvae, while termites are usually taken whenever available. Wagtail species take a variety of larval aquatic insects, such as dragonfly and mayfly nymphs, and caddisfly and stonefly larvae.

Other invertebrates eaten include spiders, crustaceans such as Isopoda, Amphipoda and crabs, annelid worms, Myriapoda, and small terrestrial, freshwater and marine mollusks. Vertebrate prey includes small fish, small frogs, tadpoles and small chameleons. Plant material is sometimes eaten, especially in the winter, and includes grass seeds, weed seeds, berries, grass blades, pine tree seeds, and tree buds; even young vegetables are reportedly eaten by the Australasian pipit. The olive-

backed pipit feeds chiefly on insects in the summer and seeds in the winter. Some species occasionally take more unusual foods, including carrion, and the cape wagtail will forage around human habitation, eating raw meat, fat, cheese, maize meal, bread, and cake.

The birds forage on open ground, in grass and herbaceous vegetation, among domestic stock, at water margins, in shallow water, on floating vegetation, and in trees and bushes; some species even follow the plough. The red-throated pipit also forages in seaweed on beaches, while the rock pipit wades in seawater, following retreating waves on beaches. Foraging methods vary with species, and include the following main techniques: (1) picking from the ground or the water surface while walking; (2) run-picking by making darting runs to catch prey; (3) immersion, by plunging the head into water; (4) fly-catching and aerial pursuit, by making short to long flights in pursuit of prey; (5) hovering to catch airborne prey or prey on the water surface; (6) probing in ground vegetation, crevices or leaf-litter. The long tail of wagtails helps the birds' balance when run-picking and flycatching, and assists in the control of aerial maneuvers when pursuing insects in flight.

In the nonbreeding season, some wagtail and pipit species feed in flocks, exploiting large patches of food. Some wagtails maintain individual winter feeding territories to defend dependable but localized food supplies, especially adjacent to water. Territory boundaries are vigorously defended with displays involving head-bobbing and short jumps into the air. Territoriality may vary with food abundance, and individuals may switch between defending patchy resources and feeding communally at widespread patches. Winter pairs may occupy territories, or an adult may share the territory with an immature bird.

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