The claim of the ruff to a place in the American list is a much stronger one than that of most of the Palaearctic species which figure in it, as it has occurred three times in Greenland, once in Canada, on many occasions in the eastern States, in Barbados, Guadeloupe, and South America.

Spring.—Apparently the ruff migrates chiefly by night and in this connection it is noticeable that the period of its greatest activity in the breeding season is during the twilight of the early morning hours. Naumann (1887) states that the first birds to arrive are the young of the preceding year, and that the adult males are the next to follow while they in turn are succeeded by the old females. Owing to the enormous development of the feathered ruff from which the species derives its names, the males are readily distinguished during the breeding season even in flight and on the ground can be recognized at almost any distance.

Courtship.—The breeding habits of the ruff are so remarkable that it is necessary to treat of them in considerable detail. When the males reach the breeding ground they are in full breeding plumage, the bare skin of the face being covered with bright yellow warts, while a disk of feathers protects the neck and two tufts project from each side of the head. The extraordinary variation in the coloring of these feather adornments renders it possible to identify individual birds, as it is rare to see two with even approximately similar coloring, and this enabled Mr. Selous (1906) to make the valuable observations which are referred to below.

Where ruffs are common, as in North Holland, one finds from time to time bare areas of ground where the grass has been worn away in patches. They are the playing grounds of the ruffs, and were known in England technically as “hills.” During the daytime they are resorted to from time to time by the male birds, which may often be seen sparring with one another, but the significance of these meetings was little understood until Edmund Selous (1906) spent a fortnight in the spring of 1906 in Holland, during which he concealed himself in a hide which commanded a good view of the “hill” at close quarters, and was often on the watch before daylight. The diary of his observations was first published in the Zoologist for 1906 and 1907, and is too lengthy and discursive to be reproduced here, but in the following condensed account most of the essential facts have been preserved. The “hill” under observation was roughly about 10 paces by 6, with 11 distinct patches where the grass had been worn bare, besides a couple of others less plainly marked. There were other similar “hills” in the neighborhood, all much alike, placed on dry ground, in the neighborhood of marshes. They are resorted to year after year, and bear traces of excrement from previous seasons, while the grass is much worn away owing to the presence of the birds. Some of the Dutch “hills” are quite close to or actually in the way of paths, and the birds when disturbed by passers-by will return to the ground within a few minutes. Like many other Dutch birds they show much indifference to the presence of man, owing to the fact that no shooting is permitted during the breeding season. On April 14 a single male was disturbed by Selous from the “hill” at 7 a. m. and did not return, but on the next day a flock of eight or nine birds arrived and at once began to fight, but not on the “hill.” It soon became evident that in order to be in place before the arrival of the birds it was necessary to get into hiding before dawn. By April 16 the “hill” was in regular use, each male resorting to the same stand and sitting there, from time to time rising and making a slight demonstration with erected tufts and ruff, sometimes followed by a slight sparring match with one of the other males, but when not engaged in fighting spending the time quite amicably side by side, in some cases almost touching. In the afternoon a similar state of things prevailed, but the proceedings were quieter and more subdued. When sparring the birds “would rush and leap high into the air against each other, rushing away and not leaping again as they came down,” and this activity was generally more pronounced when a reeve appeared on the ground. The other birds on occasions of this kind also rushed about in wild excitement, suddenly pulling themselves up with the head extended forward and remaining in this position motionless, perhaps then rushing in another direction and then gradually sinking down quickly. When a reeve arrived at the “hill,” where perhaps a dozen males were present, after the first wave of excitement had passed she would thread her way among the ruffs, which as she approached sank down to the ground by her side with bowed head. Her preference for one particular male was shown by halting at his side and gently nibbling with her bill the feathers at the back of his neck. If copulation followed, there was no interference on the part of the other males present, except perhaps occasionally by accident in the course of excited running to and fro.

After persistent watching it became evident that the attentions of the reeves were very unevenly distributed among the regular habitues of the “hill.” There were two conspicuously handsome males—one a bird with a golden-brown ruff and the other with a blue-black one—and these two seemed to be selected far more frequently than the other ruffs. The few attempts made by males to force the compliance of the female were quite unsuccessful. Perhaps it is due to this that attempts to pair between two male birds frequently took place and, strange to relate, even between two reeves also. Selection on the part of the hen birds apparently bore no relation to the fighting powers of the male, although one serious fight, which lasted some three or four minutes, took place, during which the two birds concerned fought like demons, but finally ceased by mutual consent. The two selected birds were, however, certainly among the finest and handsomest birds on the ground. While the stations were usually occupied by the same birds day after day, there were times when over 20 birds were present, so it is probable that males from other “hills” call in at times. While it was clear that one ruff paired with several reeves, it was much more difficult to prove that the reeves paired with more than one male, owing to their general similarity of appearance, though it is probable that this was the case. The following extract from Selous (1906) gives a good description of the characteristic sparring:

Each ruff has certainly a place of its own, and the most envenomed fights appear to me to result from one bird pitching down in another’s place when he flies in. The aggrieved bird instantly rushes at the intruder and there is a fight which may last for a longer or shorter time. The birds have then a perfectly frantic appearance. They kick, strike with their wings, and especially endeavor to bite or peck each other. This biting is very noticeable, the mandibles seeming to snap with rage. They leap commonly before they close and then continue to do so as a matter of course, the object seeming to be, as with other birds, to get above the adversary and strike down upon him with wings and feet and bill. Of course, when one bird is displaced by another, others are displaced also, leading to general commotion and, moreover, the mere anticipation of any fresh arrival is matter for excitement for every ruff on the ground.

Summarizing the results of Mr. Selous’ observations, it becomes evident that the “hills” are the pairing grounds to which both males and females resort, but the period of the greatest activity is during the early morning from about 3.30 a. m. onward. Each male has a definite place and the choice of a mate rests entirely with the female, the males adopting an attitude of supplication, crouching low with partially spread wings, tail and beak pointing to the ground. The hen marks her preference by nibbling the back of the neck of the prostrate male with her bill, and soon afterwards coition takes place, the surrounding males remaining usually quiescent. One reeve was seen to pair with two ruffs in succession and it is probable that she is polyandrous as the ruff is certainly polygamous. There seemed to be no connection between the fighting power of the males and the preference exercised so strikingly by the females, but, on the other hand, the favored ruffs were apparently always handsome and strikingly colored birds.

Nesting.—The reeve makes her nest within a reasonable distance of the “hill,” but not very close to it. Naumann (1887) says that it is never less than a hundred paces distant, and most of those which I have seen were within a quarter of a mile. In the Dutch water meadows the usual nesting site is among fine, thickly growing grass, where it is difficult to see unless one’s eye is caught by a glimpse of the nest hollow. In the marshes of Lapland, where vegetation is more backward, it may be found among clumps of sedge and rushes. It is built entirely of fine grasses, and would frequently be overlooked if it were not for the sight of the bird when flushed.

Eggs.—Normally four; sets of three are quite exceptional and are probably second or third layings. They are pyriform in shape, thin shelled, and, as is frequently the case with the eggs of waders, large for the size of the bird. The ground color varies from brownish yellow to pale olive brown as a rule, but occasionally has a greenish tinge, while varieties with a clear pale blue ground are not very scarce. The markings consist of spots and blotches of brown ranging from umber to dark sepia and ashy grey shellmarks, more numerous at the large end. White eggs have been recorded. The measurements of 143 eggs, chiefly by the writer, average 43.97 by 30.7 millimeters; the eggs showing the four extremes measure 47.5 by 30.5, 43.2 by 32.1, 39.2 by 31.6, and 42.9 by 28 millimeters.

They are usually laid in central Europe during the latter half of May and early in June, occasionally in the second week of May. In northern Europe most eggs are found during the latter half of June. Only one brood is reared in the season.

Young.—Incubation is performed by the female alone. Naumann (1887) gives the incubation period as 17 to 19 days, but this is probably too low an estimate and recently Heinroth has recorded 27 days. From observations made by Mr. W. H. St. Quinten on young hatched in captivity they leave the nest very soon after hatching, apparently directly the down is dry. The reeve is a good mother, and I have seen one when flushed from eggs on the point of hatching run from the nest, flapping her wings in a helpless way as if unable to fly. Another bird was also seen trying to distract attention by running with every feather erect, looking more like some small mammal than a bird.

Plumages.—The plumages and moults are fully described in A Practical Handbook of British Birds, edited by H. F. Witherby.

Food.—The ruff is mainly insectivorous, feeding largely on beetles (Coleoptera), chiefly the smaller species. Jäckel records specimens of the following genera: Agonum, Bembidium, Parnus, Cyclonotum, Sitones, Pisodes, Rhinoncus, Philhydrus. Weevils are also mentioned by Florence. Among the Orthoptera remains of grasshoppers were found by Slater. Neuroptera met with include caddis worms and cases (Phryganeidae) and larvae of Ephemera, Trichoptera (Limnophilus by Jäckel) who also records Naucoris (Hemiptera). It will be noted that most of the above are aquatic forms. Diptera are a staple food in the north, including larvae of Tipulidae and many small species. Of Mollusca, Slater records small fresh water bivalves, Collett worms (Lumbrici). In winter the food is more largely vegetable in character, though fresh water algae and seeds of Rumex have been recorded in summer. Rice is frequently found in birds killed in India and Mellor also found in seven cases nothing but durra grains (Sorghum) from 19 to 228 in number.

Voice.—The ruff is an extremely silent bird. One may live among them in the Dutch polders for weeks together in the spring, and never hear a sound from them, except a very low guttural or quacking note from a reeve when disturbed from a family of young. Naumann (1887) however describes a note kack, kack, kick, kack, which is probably the same as that which Slater writes as wick, repeated rapidly several times. Unless much persecuted it is confiding in its habits and pays little attention to the presence of men.

Enemies.—In England its extermination was primarily due to the reckless way in which the breeding stock was netted not only in autumn but also on arrival in spring on the “hilling” grounds, in order to be fattened for the table. Montagu’s account of the state of things in his day has been quoted at length in many books on British birds, so that it is not necessary to repeat it here. As Pennant spoke of 40 or 50 dozen birds being taken by a single fowler in a season it is not surprising that the stock was rapidly reduced to so low a level that in spite of belated efforts to protect the birds and their eggs, it has now practically disappeared. On the continent where it is still locally common it has to contend against other avian enemies, more especially the harriers (Circus), but also the goshawk, peregrine, and to some extent gulls and crows.

Fall.—The distinctive plumes of the male are shed in June and in July the autumn plumage is assumed even in the extreme north. The males begin to disappear from their breeding haunts in central Europe in late July and early August, while the females are said to leave in September.

Winter.—In their winter quarters ruffs are generally sociable and are to be met with in small flocks, which show a preference to fresh water marshes rather than the sea coast. In India and the Sudan the flooded fields are a great attraction and the staple food consists of grain.

DISTRIBUTION

Breeding range.—Formerly at many places in England from Northumberland southward, lingering until recently in Norfolk. Belgium, locally in North France, Holland, Denmark, North Germany, and Bavaria (Jäckel). Hungary, Czechoslovakia, the Baltic Republics, Norway, Sweden, Finland, Russia north to Kolguev, Waigatz, and Dolgoi and south to Bessarabia and Orenburg. In Asia, Siberia north to 72° on the Yenisei and east to the Lena and Kolyma valleys, south to Dauria, Turgai and Uralsk.

Winter range.—From Europe through Africa south to Cape Province, but scarce on passage in the Atlantic Islands. Asia, south to India, Ceylon (scarce), Burma, also recorded from China and Japan, Borneo (Labuan), etc.

Spring migration.—At the Straits of Gibraltar the northward movement begins in January and does not cease till May, but it is at its height in April. Some birds winter in the Mediterranean region and these have usually gone by the middle of April. In the eastern Mediterranean the passage begins in the Ionian Isles about March 15 and in Corfu in March.

Fall migration.—The return passage through southern Spain takes place in August and September, while in Greece, where it is less noticeable, it appears to take place toward the end of September.

Casual records.—The species has been rather frequently detected on the Atlantic coast of North America and (rarely) in the interior eastern States. Among these occurrences are: Ontario, Toronto Island, spring of 1882; Nova Scotia, near Halifax, May 27, 1892; New Brunswick, Grand Manan; Maine, Scarboro, April 10, 1870, Upton, September 8, 1874, and Camden, September 14, 1900; New Hampshire, Seabrook, September 23, 1907; Massachusetts, Newburyport, May 20, 1871, Chatham, September 11, 1880, and Nantucket, July, 1901; Rhode Island, Seaconnet Point, July 30, 1900, and Point Judith, August 31, 1903; New York, two on Long Island, one of which was taken May 18, 1868, and Freeport, September 27, 1914; New Jersey, Barnegat (Elliott collection); Virginia, Four-mile Run, September 3, 1894; North Carolina, Raleigh, May 6, 1892; Ohio, Licking Reservoir, November 11, 1872, and Columbus, April 23, 1878; and Indiana, English Lake, April 12, 1905.

It also has been taken in the Lesser Antilles, Barbados, previous to 1848; Colombia, probably between the Rio Negro and the Orinoco or adjacent regions; and Alaska, St. Paul Island, September 7, 1910. It is of rare occurrence in Japan (Yezo). There is one record from Iceland (Reykjavik, September, 1820).

Egg dates.—In Holland, 4 dates between May 7 and 14; 18 dates between May 15 and 31, 8 dates between June 1 and 22. In Lapland and North Russia, 16 dates between June 8 and 27.