Contour feather moult of Ruffs Philomachus pugnax during northward migration, with notes on homology of nuptial plumages in scolopacid waders

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Abstract
Among the sandpiper family Scolopacidae, the Ruff Philomachus pugnax combines a large seasonal change in the appearance of the plumage with a very pronounced sexual plumage dimorphism. Studies on the east and south African wintering grounds of Ruffs indicate that before northward migration at least the males moult (part of) their basic (winter) plumage into a kind of alternative plumage. We studied the details of the subsequent moult into a final (supplemental) breeding plumage by quantifying the presence of three feather types—(1) winter (basic), (2) striped (alternate) and (3) breeding (supplemental)—in breast feather samples of 1441 Ruffs captured on staging areas in The Netherlands during northward migration in 1993‐97. Ruffs arriving in March show a mix of winter and striped feathers. In April, the ‘breeding feather’ type appears in both male and female Ruffs, and partially takes the place of winter feathers as well as striped feathers in males, and winter feathers only in females. The presence of three plumages in Ruffs is thus confirmed for males, but also occurs in female Ruffs and in Bar‐tailed Godwits Limosa lapponica. We suggest that striped feathers represent the ‘original’ alternative plumage feather type of the sandpiper family and that the showy feathers of the, in the European literature fortuitously appropriately named, ‘supplementary plumage’ represent an additional feather generation. Such colourful nuptial plumages could thus be derived characters that have evolved independently in several scolopacid genera, presumably under particularly strong sexual selection pressures.