Ruff
The Ruff is a medium-sized wading bird in the sandpiper family, Scolopacidae. The original English name for this bird, dating back to at least 1465, is the ree, perhaps derived from a dialectical term meaning "frenzied". A later name reeve, which is still used for the female, is of unknown origin but may be derived from the shire-reeve, a feudal officer, likening the male's flamboyant plumage to the official's robes. The current name was first recorded in 1634 and is derived from the ruff, an exaggerated collar fashionable from the mid-16th century to the mid-17th century, since the male bird's neck ornamental feathers resemble the neck wear.
The Ruff has a distinctive appearance with a small head, medium-length bill, longish neck and pot-bellied body. It has long legs that are variable in colour but usually yellow or orange. In flight, it has a deeper, slower wing stroke than other waders of a similar size and it displays a thin, indistinct white bar on the wing and white ovals on the sides of the tail.
The Ruff shows very distinctive sexual dimorphism. Although a small percentage of males resemble females, the typical male is much larger than the female and has an elaborate breeding plumage.
The male is 11 to 13 inches in length with a 21 to 24 inches wingspan. During the May to June breeding season, the typical male's legs, bill and warty bare facial skin are orange and he has distinctive head tufts and a neck ruff. These ornaments vary on individual birds and are black, chestnut or white with the colouring solid, barred or irregular. The grey-brown back has a scale-like pattern, often with black or chestnut feathers, and the underparts are white with extensive black on the breast. The extreme variability of the main breeding plumage is thought to have developed to aid individual recognition in a species that has communal breeding displays but is usually mute. Outside the breeding season, the typical male's head and neck decorations and the bare facial skin are lost and the legs and bill become duller. The upperparts are grey-brown and the underparts are white with grey mottling on the breast and flanks.
The female, or "reeve", is 9 to 10 inches in length with a 18 to 19 inches wingspan. In the breeding plumage, the female has grey-brown upperparts with white-fringed, dark-centred feathers. The breast and flanks are variably blotched with black. In winter, her plumage is similar to that of the male but the sexes are distinguishable on size.
The plumage of the juvenile Ruff resembles the non-breeding adult but has upperparts with a neat, scale-like pattern with dark feather centres and a strong buff tinge to the underparts.
The Ruff is a migratory species, breeding in wetlands in the colder regions of northern Eurasia and wintering in the tropics, mainly in Africa. There is a limited overlap of the summer and winter ranges in the UK and parts of coastal western Europe and birds may be present throughout the year.
The Ruff breeds in Europe and Asia from Scandinavia and the UK almost to the Pacific. In Europe, it is found in cool temperate areas but over its Russian range it is an Arctic species and occurs mainly north of about 65°N. The largest numbers breed in Russia, Sweden, Finland and Norway. Although it also breeds from the UK east through the Low Countries to Poland, Germany and Denmark, the populations are much lower in these more southerly areas.
The Ruff is highly gregarious on migration and it travels in large flocks that can contain hundreds or thousands of individuals. Huge dense groups form on the wintering grounds. The males, which play no part in nesting or chick care, leave the breeding grounds in late June or early July followed later in July by the females and juveniles. Males typically make shorter flights and winter further north than 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 thereby improving their chances of successful mating. Males may also be able to tolerate colder winter conditions because they are larger than females.
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. When it is not breeding, it can be found in a wider range of shallow wetlands. Dry grassland, tidal mudflats and the seashore are less frequently used.
Male Ruffs display during the breeding season at a lek in a traditional open grassy arena. The Ruff is one of the few lekking species in which the display is primarily directed at other males rather than to the females and it is among a small percentage of birds in which the males have well-marked and inherited variations in plumage and mating behaviour.
There are 3 male forms: the typical territorial males, satellite males which have a white neck ruff and a very rare variant with female-like plumage. The behaviour and appearance for an individual male remain constant through its adult life and are determined by its genes.
The territorial males, about 84% of the total, have strongly coloured black or chestnut ruffs and stake out and occupy small mating territories in the lek. They actively court females and display a high degree of aggression towards other resident males. Between 5 and 20 territorial males each hold an area of the lek about 3 feet across and usually with bare soil in the centre. They perform an elaborate display that includes wing fluttering, jumping, standing upright, crouching with ruff erect or lunging at rivals. They are typically silent even when displaying. Territorial males are very site faithful and 90% return to the same lekking sites in subsequent seasons, the most dominant males being the most likely to reappear. Site faithful males can acquire accurate information about the competitive abilities of other males, leading to well-developed dominance relationships. Such stable relationships reduce conflict and the risk of injury and the consequent lower levels of male aggression are less likely to scare off females. Lower-ranked territorial males also benefit from site fidelity since they can remain on the leks while waiting for the top males eventually to drop out.
Satellite males, about 16% of the total number, have white or mottled ruffs and do not occupy territories. Instead, they enter leks and attempt to mate with the females visiting the territories occupied by the resident territorial males. Resident territorial males tolerate the satellite birds because, although they are competitors for mating with the females, the presence of both types of male on a territory attracts additional females.
A third type of male was first described in 2006. This is a permanent female mimic, the first such reported for a bird. About 1% of males are small, intermediate in size between males and females and do not grow the elaborate breeding plumage of the territorial and satellite males. This cryptic male obtains access to mating territories together with the females and "steals" matings when the females crouch to solicit copulation. It moults into the prenuptial male plumage with striped feathers but does not go on to develop the ornamental feathers of the normal male.
Not all mating takes place at the lek since only a minority of the males attend an active lek. As alternative strategies, males can also directly pursue females or wait for them as they approach good feeding sites. The level of polyandry in the Ruff is the highest known for any avian lekking species and for any shorebird. More than half of females mate with and have clutches fertilised by more than one male. In lekking species, females can choose mates without risking the loss of support from males in nesting and rearing chicks since the males take no part in raising the brood anyway.
The nest of the Ruff is a shallow ground scrape lined with grass leaves and stems and concealed in marsh plants or tall grass up to 450 yards from the lek. Nesting is solitary although several females may lay in the general vicinity of a lek. The female lays typically 4 eggs from mid-March to early June depending on latitude. Incubation is undertaken by the female alone and the time to hatching is 20 to 23 days with a further 25 to 28 days to fledging. The precocial chicks have buff and chestnut down, streaked and barred with black and frosted with white. They feed themselves on a variety of small invertebrates.
The Ruff normally feeds using a steady walk and pecking action, selecting food items by sight, but it will also wade deeply and submerge its head. During the breeding season, the Ruff’s diet consists almost exclusively of the adults and larva of terrestrial and aquatic insects such as beetles and flies. On migration and during the winter, it eats insects, crustaceans, spiders, molluscs, worms, frogs, small fish and also the seeds of rice and other cereals, sedges, grasses and aquatic plants.
The Ruff has a large range and a large population although there is some evidence of a decrease in the population in some regions and countries. However, the species as a whole is not believed to approach the thresholds for the population decline criterion of the IUCN Red List (i.e. declining more than 30 percent in 10 years or 3 generations). For these reasons, the Ruff is classified as "least concern". However, the range in much of Europe is contracting because of land drainage, increased fertiliser use, the loss of mown or grazed breeding sites and over-hunting. Therefore the Ruff is one of the species to which the Agreement on the Conservation of African-Eurasian Migratory Waterbirds (AEWA) applies.
Date: 1st October 2012
Location: Titchwell, Norfolk
The Ruff has a distinctive appearance with a small head, medium-length bill, longish neck and pot-bellied body. It has long legs that are variable in colour but usually yellow or orange. In flight, it has a deeper, slower wing stroke than other waders of a similar size and it displays a thin, indistinct white bar on the wing and white ovals on the sides of the tail.
The Ruff shows very distinctive sexual dimorphism. Although a small percentage of males resemble females, the typical male is much larger than the female and has an elaborate breeding plumage.
The male is 11 to 13 inches in length with a 21 to 24 inches wingspan. During the May to June breeding season, the typical male's legs, bill and warty bare facial skin are orange and he has distinctive head tufts and a neck ruff. These ornaments vary on individual birds and are black, chestnut or white with the colouring solid, barred or irregular. The grey-brown back has a scale-like pattern, often with black or chestnut feathers, and the underparts are white with extensive black on the breast. The extreme variability of the main breeding plumage is thought to have developed to aid individual recognition in a species that has communal breeding displays but is usually mute. Outside the breeding season, the typical male's head and neck decorations and the bare facial skin are lost and the legs and bill become duller. The upperparts are grey-brown and the underparts are white with grey mottling on the breast and flanks.
The female, or "reeve", is 9 to 10 inches in length with a 18 to 19 inches wingspan. In the breeding plumage, the female has grey-brown upperparts with white-fringed, dark-centred feathers. The breast and flanks are variably blotched with black. In winter, her plumage is similar to that of the male but the sexes are distinguishable on size.
The plumage of the juvenile Ruff resembles the non-breeding adult but has upperparts with a neat, scale-like pattern with dark feather centres and a strong buff tinge to the underparts.
The Ruff is a migratory species, breeding in wetlands in the colder regions of northern Eurasia and wintering in the tropics, mainly in Africa. There is a limited overlap of the summer and winter ranges in the UK and parts of coastal western Europe and birds may be present throughout the year.
The Ruff breeds in Europe and Asia from Scandinavia and the UK almost to the Pacific. In Europe, it is found in cool temperate areas but over its Russian range it is an Arctic species and occurs mainly north of about 65°N. The largest numbers breed in Russia, Sweden, Finland and Norway. Although it also breeds from the UK east through the Low Countries to Poland, Germany and Denmark, the populations are much lower in these more southerly areas.
The Ruff is highly gregarious on migration and it travels in large flocks that can contain hundreds or thousands of individuals. Huge dense groups form on the wintering grounds. The males, which play no part in nesting or chick care, leave the breeding grounds in late June or early July followed later in July by the females and juveniles. Males typically make shorter flights and winter further north than 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 thereby improving their chances of successful mating. Males may also be able to tolerate colder winter conditions because they are larger than females.
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. When it is not breeding, it can be found in a wider range of shallow wetlands. Dry grassland, tidal mudflats and the seashore are less frequently used.
Male Ruffs display during the breeding season at a lek in a traditional open grassy arena. The Ruff is one of the few lekking species in which the display is primarily directed at other males rather than to the females and it is among a small percentage of birds in which the males have well-marked and inherited variations in plumage and mating behaviour.
There are 3 male forms: the typical territorial males, satellite males which have a white neck ruff and a very rare variant with female-like plumage. The behaviour and appearance for an individual male remain constant through its adult life and are determined by its genes.
The territorial males, about 84% of the total, have strongly coloured black or chestnut ruffs and stake out and occupy small mating territories in the lek. They actively court females and display a high degree of aggression towards other resident males. Between 5 and 20 territorial males each hold an area of the lek about 3 feet across and usually with bare soil in the centre. They perform an elaborate display that includes wing fluttering, jumping, standing upright, crouching with ruff erect or lunging at rivals. They are typically silent even when displaying. Territorial males are very site faithful and 90% return to the same lekking sites in subsequent seasons, the most dominant males being the most likely to reappear. Site faithful males can acquire accurate information about the competitive abilities of other males, leading to well-developed dominance relationships. Such stable relationships reduce conflict and the risk of injury and the consequent lower levels of male aggression are less likely to scare off females. Lower-ranked territorial males also benefit from site fidelity since they can remain on the leks while waiting for the top males eventually to drop out.
Satellite males, about 16% of the total number, have white or mottled ruffs and do not occupy territories. Instead, they enter leks and attempt to mate with the females visiting the territories occupied by the resident territorial males. Resident territorial males tolerate the satellite birds because, although they are competitors for mating with the females, the presence of both types of male on a territory attracts additional females.
A third type of male was first described in 2006. This is a permanent female mimic, the first such reported for a bird. About 1% of males are small, intermediate in size between males and females and do not grow the elaborate breeding plumage of the territorial and satellite males. This cryptic male obtains access to mating territories together with the females and "steals" matings when the females crouch to solicit copulation. It moults into the prenuptial male plumage with striped feathers but does not go on to develop the ornamental feathers of the normal male.
Not all mating takes place at the lek since only a minority of the males attend an active lek. As alternative strategies, males can also directly pursue females or wait for them as they approach good feeding sites. The level of polyandry in the Ruff is the highest known for any avian lekking species and for any shorebird. More than half of females mate with and have clutches fertilised by more than one male. In lekking species, females can choose mates without risking the loss of support from males in nesting and rearing chicks since the males take no part in raising the brood anyway.
The nest of the Ruff is a shallow ground scrape lined with grass leaves and stems and concealed in marsh plants or tall grass up to 450 yards from the lek. Nesting is solitary although several females may lay in the general vicinity of a lek. The female lays typically 4 eggs from mid-March to early June depending on latitude. Incubation is undertaken by the female alone and the time to hatching is 20 to 23 days with a further 25 to 28 days to fledging. The precocial chicks have buff and chestnut down, streaked and barred with black and frosted with white. They feed themselves on a variety of small invertebrates.
The Ruff normally feeds using a steady walk and pecking action, selecting food items by sight, but it will also wade deeply and submerge its head. During the breeding season, the Ruff’s diet consists almost exclusively of the adults and larva of terrestrial and aquatic insects such as beetles and flies. On migration and during the winter, it eats insects, crustaceans, spiders, molluscs, worms, frogs, small fish and also the seeds of rice and other cereals, sedges, grasses and aquatic plants.
The Ruff has a large range and a large population although there is some evidence of a decrease in the population in some regions and countries. However, the species as a whole is not believed to approach the thresholds for the population decline criterion of the IUCN Red List (i.e. declining more than 30 percent in 10 years or 3 generations). For these reasons, the Ruff is classified as "least concern". However, the range in much of Europe is contracting because of land drainage, increased fertiliser use, the loss of mown or grazed breeding sites and over-hunting. Therefore the Ruff is one of the species to which the Agreement on the Conservation of African-Eurasian Migratory Waterbirds (AEWA) applies.
Date: 1st October 2012
Location: Titchwell, Norfolk
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Also in: Sandpipers
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