The Ruff

Picture this: a fine, dandy-looking Renaissance gentleman in fancy, pompous clothes and a resplendent  ruff around his neck.

Sir Henry Unton (1586) unknown British artist. Source: tate.org.uk

Now picture this: a bunch of these dandy-looking gentlemen posing, strutting and fiercely scuffling in a muddy field to attract the attention of the ladies. In fancy clothes, ruff and all. That’s right.

Although I am an occasional thespian and a theatre nerd, that won’t be the focus of today’s post. This post is about a very different kind of Ruff: the living kind.

The Ruff (Calidris pugnax) is a medium-sized migratory wading bird, breeding in marshlands and grassy fields across northern Eurasia, especially in Scandinavia and Siberia, but also in the more temperate British Isles. As winter comes, they shed their breeding plumage and migrate southwards in vast flocks, fleeing the bitter cold. They spend the winter in the tropics, mainly in Africa where thousands may congregate in suitable habitats. A minority winter in areas including the Mediterranean, the Indian Subcontinent, Southeast Asia and even southern Australia. 

In Sri Lanka, it is a scarce but regular winter visitor, found in salt pans, marshes and mudflats of the dry zone coastline. I first saw it in a saltern in Bundala National Park in 2010, when the safari guide identified it for us. Ten years later, I spotted one in the same area - and this time, I was able to identify it on my own.

Ruff in Bundala National Park, January 2020

A better view of the very same Ruff, alongside a Black-winged Stilt

In Sri Lanka, Ruffs are found alone or in small groups, owing to their scarcity. They are often found in mixed-species wading flocks, which makes them easier to identify by comparison.

When summer approaches - in March-April - Ruffs all over their wintering grounds feel the need to return home to breed. They begin by putting on the beginnings of their courtship trappings: their feathers become more “scaly” in appearance, and they become more mottled.

They stock up intensively for the long journey north, but leave before they’re anywhere close to full breeding plumage, which they assume after returning home.

Females, known as Reeves, have it simple. They, like all males initially, assume a dark, scaly-stripey appearance but their small, stocky silhouettes remain mostly unchanged - and that is how they remain throughout the breeding season.

When it comes to the significantly larger males, however, things get weird. Very weird. Over their evolution, genetic mutations in their chromosomes have given rise to not one, not two, but three “male” forms, which I shall explain later once they have fully “transitioned” into their breeding plumage in the following trio of photos. (Note: I don’t own the following three photos. I have used them here under the Creative Commons license, so that even if I haven’t personally witnessed the spectacle of Ruff courtship, I’d still be able to share it here.)

The first one looks fairly normal and expected, similar to the transformation of other male waders.

Transitioning male Ruff, in Poland. Photo credits: Jerzy Strzelecki
(https://creativecommons.org/licenses/by/2.0/deed.en)

This second male looks very fancy with a lovely auburn “bob”. But he’s not there yet!

Male Ruff in transition, Lappland. Photo credits: Åsa Berndtsson

(https://creativecommons.org/licenses/by/2.0/deed.en)

Here they are in all their glory: with flamboyant ruffs around their necks and plumes on their heads, complete with attractively patterned mantles.

Two fully-plumed male Ruffs in Netherlands. Photo credits: Arjan Haverkamp

(https://creativecommons.org/licenses/by/2.0/deed.en)

Ruffs gather on open grassy fields to display, compete and mate in gatherings called “leks”. A lek is an orgy of sorts, and females often mate with multiple males (polyandry) sometimes of different forms - and even homosexual matings are known to occur.

The most typical male ruff, named the “territorial male”, usually has a black or bright chestnut ruff, and is stoutly built. These males are the most numerous, and overtly competitive, defending territories and lek spaces which they return to every year. They perform elaborate courtship displays to attract females actively (Ruffs can get very feisty and pugnacious - hence their specific name, pugnax).

The second type is the “satellite male”, which is more lightly built and has a white or mottled ruff. They do not occupy territory, instead dropping in on leks whenever displays are underway, occasionally managing to sneak an opportunity to mate. Territorial males seem to mostly tolerate them because their presence adds to the numbers and attracts more females to the lek.

There is a third, sneakier type - the rare “faeder”. Faeders never moult past the feminine breeding plumage, instead permanently mimicking females. Ruffs are the only birds in which this phenomenon has been observed. Faeders, in the guise of females, sneak into leks to mate. What’s more interesting is that they engage in homosexual mating, which the birds themselves are believed to be aware of: faeders can be mounted by typical males, and vice versa.

After all the pageantry of breeding, mated females go their separate ways to hatch and raise their young - which often may have multiple sires - on their own. As summer wanes, Ruffs shed their breeding plumage and begin to feed and gain weight, storing energy to travel south once more.

Here’s a plate from Birds of Sri Lanka, by Warakagoda & Inskipp, that features the Ruff, with focus on the breeding plumage. I don’t know why they included it - there’s no chance that a birder could see the Ruff’s true splendour in Sri Lanka - but I’m glad they did.

The Ruff. Plate 31, Helm Field Guides’ Birds of Sri Lanka (Deepal Warakagoda, Carol Inskipp, Tim Inskipp & Richard Grimmett)

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This article is here thanks to these two links. Initially I was only planning to write about the plumage transformation - but these uncovered a whole new side to the story. Check them out for a more in-depth look into the Ruff’s breeding systems:

https://www.newscientist.com/article/dn28493-ruff-bird-orgies-have-four-sexes-thanks-to-a-supergene-flip/

https://www.rug.nl/research/portal/files/6700647/2006BiolLettJukema.pdf