Distribution and Habitat
The Ruff is a migratory species, breeding in wetlands in colder regions of northern Eurasia, and spends the northern winter in the tropics, mainly in Africa. Some Siberian breeders undertake an annual round trip of up to 30,000 km (18,500 mi) to the West African wintering grounds. There is a limited overlap of the summer and winter ranges in western Europe. The Ruff breeds in extensive lowland freshwater marshes and damp grasslands. It avoids barren tundra and areas badly affected by severe weather, preferring hummocky marshes and deltas with shallow water. The wetter areas provide a source of food, the mounds and slopes may be used for leks, and dry areas with sedge or low scrub offer nesting sites. A Hungarian study showed that moderately intensive grazing of grassland, with more than one cow per hectare (2.5 acres), was found to attract more nesting pairs. When not breeding, the birds use a wider range of shallow wetlands, such as irrigated fields, lake margins, and mining subsidence and other floodlands. Dry grassland, tidal mudflats and the seashore are less frequently used. The density can reach 129 individuals per square kilometre (334 per square mile), but is usually much lower.
The Ruff breeds in Europe and Asia from Scandinavia and Great Britain almost to the Pacific. In Europe it is found in cool temperate areas, but over its Russian range it is an Arctic species, occurring mainly north of about 65°N. The largest numbers breed in Russia (more than 1 million pairs), Sweden (61,000 pairs), Finland (39,000 pairs) and Norway (14,000 pairs). Although it also breeds from Britain east through the Low Countries to Poland, Germany and Denmark, there are fewer than 2,000 pairs in these more southerly areas.
It is highly gregarious on migration, travelling in large flocks that can contain hundreds or thousands of individuals. Huge dense groups form on the wintering grounds; one flock in Senegal contained a million birds. A minority winter further east to Burma, south China, New Guinea and scattered parts of southern Australia, or on the Atlantic and Mediterranean coasts of Europe. In Great Britain and parts of coastal western Europe, where the breeding and wintering ranges overlap, birds may be present throughout the year. Non-breeding birds may also remain year round in the tropical wintering quarters. The Ruff is an uncommon visitor to Alaska (where it has occasionally bred), Canada and the contiguous states of the US, and has wandered to Iceland, Middle America, northern South America, Madagascar and New Zealand. It has been recorded as breeding well south of its main range in northern Kazakhstan, a major migration stopover area.
The male, which plays no part in nesting or chick care, leaves the breeding grounds in late June or early July, followed later in July by the female and juveniles. Males typically make shorter flights and winter further north than females; for example, virtually all wintering Ruffs in Britain are males, whereas in Kenya most are females. Many migratory species use this differential wintering strategy, since it reduces feeding competition between the sexes and enables territorial males to reach the breeding grounds as early as possible, improving their chances of successful mating. Male Ruffs may also be able to tolerate colder winter conditions because they are larger than females.
Birds returning north in spring across the central Mediterranean appear to follow a well-defined route. Large concentrations of Ruffs form every year at particular stopover sites to feed, and individuals marked with rings or dye reappear in subsequent years. The refuelling sites are closer together than the theoretical maximum travel distance calculated from the mean body mass, and provide evidence of a migration strategy using favoured intermediate sites. The Ruff stores fat as a fuel, but unlike mammals, uses lipids as the main energy source for exercise (including migration) and, when necessary, keeps warm by shivering; however, little research has been conducted on the mechanisms by which they oxidise lipids.
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