The ruff (Calidris pugnax) is a medium-sized wading bird that breeds in marshes and wet meadows across northern Eurasia. This highly gregarious sandpiper is migratory and sometimes forms huge flocks in its winter grounds, which include southern and western Europe, Africa, southern Asia and Australia.

The ruff is a long-necked, pot-bellied bird. This species shows marked sexual dimorphism; the male is much larger than the female (the reeve), and has a breeding plumage that includes brightly coloured head tufts, bare orange facial skin, extensive black on the breast, and the large collar of ornamental feathers that inspired this bird's English name. The female and the non-breeding male have grey-brown upperparts and mainly white underparts. Three differently plumaged types of male, including a rare form that mimics the female, use a variety of strategies to obtain mating opportunities at a lek, and the colourful head and neck feathers are erected as part of the elaborate main courting display. The female has one brood per year and lays four eggs in a well-hidden ground nest, incubating the eggs and rearing the chicks, which are mobile soon after hatching, on her own. Predators of wader chicks and eggs include mammals such as foxes, feral cats and stoats, and birds such as large gulls, crows and skuas.

The ruff forages in wet grassland and soft mud, probing or searching by sight for edible items. It primarily feeds on insects, especially in the breeding season, but it will consume plant material, including rice and maize, on migration and in winter. Classified as "least concern" on the IUCN Red List criteria, the global conservation concerns are relatively low because of the large numbers that breed in Scandinavia and the Arctic. 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. This decline has seen it listed in the Agreement on the Conservation of African-Eurasian Migratory Waterbirds (AEWA).

Taxonomy and nomenclature

The ruff is a wader in the large family Scolopacidae, the typical shorebirds. Recent research suggests that it is in a near-basal position in the genus Calidris, the sister taxon to all of the species except for the knot group (red knot, great knot and surfbird). It has no recognised subspecies or geographical variants.

These cryptic males obtains access to mating territories together with the females, and "steals" matings when the females crouch to solicit copulation.

In 2016, two studies further pinpointed the responsible region to chromosome 11 and a 4.5-Mb covering chromosomal rearrangement. The scientists were able to show that the first genetic change happened 3.8 million years ago on the resident chromosome, when a part of it broke off and was reintroduced in the wrong direction. This inversion created the faeder allele. About 500,000 years ago another rare recombination event of faeder and resident allele in the very same inverted region led to the satellite allele. The 4.5 Mb inversion covers 90 genes, one of them is the centromere coding gene N- CENPN-, which is located exactly at one of the inversion breakpoints. The inactivation of the gene has severe deleterious effects and pedigree data of a captive ruff colony suggests that the inversion is homozygous lethal. Over the course of the past 3.8 million years, further mutations have accumulated within the inversion i.e. three deletions ranging from 3.3 to 17.6 kb. Two of these deletions remove evolutionary highly conserved elements close to two genes- HSD17B2 and SDR42E1-both holding important roles in metabolism of steroid hormones. Hormone measurements around mating time showed that whereas residents have a sharp increase of testosterone, faeders and satellites only experience higher androstenedione levels, a substance which is considered an intermediate in testosterone biosynthesis. The authors conclude that one or more of the deletions act as a cis-acting regulatory mutation which is altering the expression of one or both genes and eventually contributes to the different male phenotypes and behaviour. It appears that females produce a larger proportion of females at the egg stage when they are in poor physical condition. When females are in better condition, any bias in the sex ratio is smaller or absent.

Predators of waders breeding in wet grasslands include birds such as large gulls, common raven, carrion and hooded crows, and great and Arctic skuas; foxes occasionally take waders, and the impact of feral cats and stoats is unknown. When feeding, the ruff frequently raises its back feathers, producing a loose pointed peak on the back; this habit is shared only by the black-tailed godwit. which could later reflect an uplisting of the species.

The most important breeding populations in Europe, in Russia and Sweden, are stable, and the breeding range in Norway has expanded to the south, but populations have more than halved in Finland, Estonia, Poland, Latvia and The Netherlands. Although the small populations in these countries are of limited overall significance, the decline is a continuation of trend towards range contraction that has occurred over the last two centuries. The drop in numbers in Europe has been attributed to drainage, increased fertiliser use, the loss of formerly mown or grazed breeding sites and over-hunting.

References

Further reading

  • Ruff species text in The Atlas of Southern African Birds
  • Ruff mating strategy introduction and research
  • Ruffs on postage stamps: www.birdtheme.org and
  • Ageing and sexing (PDF; 2.0 MB) by Javier Blasco-Zumeta & Gerd-Michael Heinze