Llewelyn Lloyd [1792-1876] was—like many a naturalist, both then and now—eccentric. Born in London to an aristocratic family of bankers and financiers, Llewelyn was more interested in hunting, fishing, and natural history. To pursue those passions, he decamped to southwestern Sweden in his twenties to spend a couple of decades travelling throughout Sweden and Norway. And, apparently womanizing, leaving a spate of illegitimate offspring in addition to the 9 children that he had with four partners, 3 of who he married (presumably consecutively).
In those days the southwest of Sweden was sparsely populated, with the largest town, Göteborg (Gothenberg), having less than 13,000 inhabitants in 1800. It was an ideal place for Lloyd to hunt for lynx, wolf and bear, and fish— the river Göta Älv that flows to the sea through Göteborg from Vänern, Sweden’s largest lake, was an outstanding salmon river. During his stay in Sweden he is said to have killed at least 100 bears, and could often be seen walking the streets of Vänersborg with a live bear in tow, on a chain.
Soon after arriving in Sweden, Lloyd began to produce a series of articles and books describing rural life and natural history for an English audience back home. His 1867 book The Game Birds and Wild Fowl of Sweden and Norway is widely attributed to introducing the word ‘lek’ to biology, to describe where males of a few species of birds gather to court and copulate with females. It was, in fact, his first book Field Sports of the North of Europe, published almost 40 years earlier, where this term was first applied to the display grounds of the Capercaillie (Tjader in Swedish): The capercali does not play indiscriminately over the forest; but he has his certain stations (Tjader-lek , which may perhaps be rendered, his playing-grounds. (Lloyd 1830, Vol. II, page 276) and then a few pages later to the display grounds of the black grouse (Orre in Swedish):The black-cock …. assemble together in large numbers , when , like the capercali , they have their particular stations (orr-lek). (Lloyd 1830, Vol. II, page 291)
Clearly the term ‘lek’ was in common use in Sweden for these mating aggressions by the early 1800s, and probably long before. In Swedish, the noun ‘lek’ means play, game, sport, or spawning, depending on the context. So a lek was a place where some birds (or mammals or fishes) appeared to be playing a game—a mating game.
In that first Field Sports book, Lloyd was primarily interested in leks was because they were an easy place to find and shoot game birds, here quoting (in translation) a Mr Greiff, (Lloyd 1830, Vol II, page 291):
To shoot black game [grouse] at this time…is a most amusing sport. They commence their play rather earlier in the season than the capercali; and in the beginning of April, forty or fifty, or even more, may be seen together. Like the capercali, they meet at an early hour in the morning. Their lek is generally on large mosses , downs , meadows, &c. and even on the surface of lakes, rivers, &c. which remain frozen late in the spring. An old black-cock, who is called the playing-cock, (or several, if the pack be sufficiently large,) acts as the master, and does not allow the others to play; but the young cocks are suffered to blow and fight with each other, and to remain with the hens. The playing-cock ought never to be shot, because that may occasion the dispersion of the whole pack; but those that blow and fly about the lek may be killed, and sometimes one may get several at a shot when they are in the act of fighting.
Mr Greiff was the royal gamekeeper, Johan Ludvig Bogislaus von Greiff [1757-1828], who had written a book in 1821 about hunting and trapping in Sweden. As Lloyd had written his book within a couple of years of arriving in Sweden, his own Field Sports book relies heavily on von Greiff’s observations, translated from the Swedish. Lloyd’s second book about birds Game Birds and Wild Fowl, published 37 years later, is much richer in detail based on Lloyd’s own travels and observations. There he describes the leks of Ruff, Black Grouse, Great Snipe (his double or solitary snipe), and Capercaillie, clearly using the term ‘lek’ in the same way it is used today to describe aggregations of courting males: “a number of display courts, or territories, of males…at which individual males are visible to one another while competing for mating rights with visiting females.” (Frith 2025).
In addition to the many insights into the migrations, songs, plumages, courtship, and aggressive behaviours of birds, Game Birds and Wild Fowl is richly illustrated. It includes, for example, more than 40 coloured engravings of birds (e.g., Black Grouse, above) and mammals from paintings by the celebrated Swedish artist Magnus Körner [1808-1864] and some exquisite woodcuts (e.g. Ruffs, below) by the Prussian Joseph Wolf [1820-1899] who Sir Edwin Landsborough [1802-1873] said was “..without exception, the best all-round animal artist who ever lived”.
Whilst the main title of that book refers to birds, more than half of the pages are devoted to detailed descriptions and illustrations of otters, seals, walrus and a variety of saltwater fishes, and the cities and countryside of southern Sweden. It is also a treasure trove of the many hunting and trapping techniques used by the country folk of Sweden, and even includes an in-depth treatment of hybrids (Rackel-Fogel) between the Black Grouse and Capercaillie that had shot, previously considered to be a distinct species by some.
I have never seen reference to Lloyd or his books in any of the many books on the history of ornithology. That’s a pity as his books are rich with information about behaviour in a time when few ornithologists seemed to be interested in the behaviours of birds. Those books are well worth reading even today for early ideas about leks, for the historical information about hunting and trapping, and for the superb illustrations.
Further Reading
Frith, C.B. (2025) Use and definitions of the terms arena, lek and court in describing avian courtship sites. Ibis 167: 295-298. VIEW
Lloyd, L. (1830) Field sports of the north of Europe; comprised in a personal narrative of a residence in Sweden and Norway, in the years 1827-28. London: H. Colburn & R. Bentley. VIEW
Lloyd, L. 1854. Scandinavian adventures, during a residence of upwards of twenty years; representing sporting incidents, and subjects of natural history, and devices for entrapping wild animals. With some account of the northern fauna. R. Bentley VIEW
Lloyd, L. (1867) The game birds and wild fowl of Sweden and Norway; with an account of the seals and salt-water fishes of those countries. London: F. Warne and co. VIEW
Lloyd, L. (1870) Peasant life in Sweden. London: Tinsley Brothers. VIEW
von Greiff, J. L.B. (1821) Anteckningar angående Jagt och Djufångst i Swerige. [Notes concerning the hunt and animal capture (trapping) in Sweden] Stockholm: H. A. Nordstrom. VIEW
Image credits
Lloyd and bear from Vänersborg Museum
Black Grouse and Ruff from Lloyd (1867) in the public domain
