Studies of less familiar ijo Wryneck R. E. F. Peal Photographs by R. G. Carlson, Eric Hosking, A. Klaver, A. jV. H. Peach and Walter Tilgner Plates 9-16 The Wryneck Jynx torquilla is unusual among the birds featured in this series in being a British breeding species, though now a very rare one. In the last century it was widely distributed, nesting in all English counties except Cornwall and Northumberland, as well as in eight counties in Wales, and country people were familiar with its call. It has been decreasing for at least 150 years for reasons not clearly understood (Monk 1963). It no longer breeds in Wales; in England nesting is now restricted virtually to the south-east, but even there records are few and well scattered. In the Scottish High­ lands, however, after some eleven sightings in late May or June in 1951-68 (Harthan et al. 1964, Macmillan 1969), there were three proved cases of breeding in 1969, all on Speyside (Burton et al. 1970). It is probable that these birds were connected with the Fenno- Scandian population (see below) rather than the English. The Wryneck was classed by Peters (1948) with the closely related Red-breasted Wryneck J. ruficollis of tropical and southern Africa as the only species in the subfamily Jynginae. Its upperparts are grey, brown and buff, motded and variegated and with streaks, especially on the back; below it is paler and, in particular on the chin, breast, flanks and undertail-coverts, more barred. It is about 16-17 cm l°ng and can easily be mistaken in the field at a distance for a passerine. The fineness of detail in the plumage is well shown on plates 9-16. The upperparts can blend very closely into the bark of a tree, as is shown by the lower on plate 12a. There are nine long primaries in the wing and a tenth (the outermost) which is shorter (plate 16); the number of secondaries is normally ten. The barring on the outer webs of the primaries is clearly visible on plates 9, 13 and 16b. In juvenile plumage the outermost primary measures about 50 mm from the carpal joint, being slightly more than half the length of the longest primary. This feather is lost in the post-juvenile moult, which affects all the plumage except the secondaries (Stresemann and Stresemann 1966), and in subsequent plumages it is much shorter, as can be seen in plate 16a. This provides a means of distinguishing juveniles in the hand. In those studied in Switzerland 66 PLATE 9. Wryneck Jynx torquilla perched near nest, Suffolk, June 1945 {photo: Eric Hashing). The plumage is a mixture of grey, brown, buff and off-white, liberally mottled, streaked and barred; the large, strong toes are positioned two forward and two back, the inner toes being much shorter than the outer ones (pages 66-72) PLATES IO and II. Wrynecks Jynx torquilla at nest-holes, Norway, June 1964, and Suffolk, June 1945 (photos: Eric Hosking). Right, young being fed in glass-backed nestbox, Netherlands, July 1963 (photo: A. Klaver): the runt seemed four to five days old, most of the other nestlings being twelve; nine young eventually fledged, including the runt which left the nest six days after most of the brood (page 70)

PLATE 12 {left). Wrynecks Jynx torquilla, Austria, June 1972, showing the camou• flage caused by the variegated plumage (photos: A. jV. H. Peach). The birds nest in both natural and excavated holes, though they very rarely make their own (page 68)

PLATE 13. Wryneck at nest with bill full of , mainly pupae, Norway, June 1964 [photo: Eric Hosking). In Europe chicks are fed almost entirely on ants (page 69) PLATE 14. Wryneck Jynx torquilla trapped while on spring migration at Azraq, Jordan, 18th April 1965; after being retrapped there on 23rd, it was found dead on 9th May at Kozelets, U.S.S.R., about 2,100 km NNW. Below, another shot taken at the Suffolk nest featured on plates 9 and 10b, June 1945 {photos: Eric Hosking) PLATE 15. Wryneck Jynx torquilla calling by nest-hole, Suffolk, June 1934 (photo: Eric Hoiking). Both sexes call frequently before and during the egg-laying period, and also when second clutches are being laid (page 70). Below, taking food to young in the Austrian nest shown on plate 12, June 1972 (photo: R. G. Carlson) PLATE 16. WrynecksJ^Tu torquilla landing at and leaving nest-hole {photos: Walter Tilgner): note the barring on the outer webs of the primaries, and the very short first primary which is much longer in juveniles (pages 66-67) Wryneck studies 67 by Sutter (1941), the outermost primary was shed when the birds were 66-67 days old and the post-juvenile moult was completed at an age of 13 weeks. This moult and the complete adult moult, which occurs at about the same time of year, appear to be concluded in the European population before the autumn migration is begun, but farther east birds may commence migration while still in moult (see Dementiev and Gladkov 1966-68). The outermost tail-feathers are much shorter than the others and lie below the pair immediately inside, as is just apparent in plate 16. This is in contrast to the (), which have the outer tail-feathers above the pair inside. The toes are usually directed two forwards and two backwards, the inner toes being shorter than the outer. The breeding range of the Wryneck stretches from Britain and Portugal in the west in a band across Eurasia to northern Japan in the east; it extends north of the Arctic Circle in Fenno-Scandia, while in the south it includes the larger Italian islands and takes in parts of northern Algeria and Tunisia. A decrease, considerable in some parts, has been observed in a number of countries on the continent of Europe (see Peal 1968), and breeding has almost certainly ceased in north-west France (L. J. Yeatman in litt.). It is migratory over almost all its range, wintering mainly in tropical Africa north of the equator, from India across to south-east China, and in southern Japan. In the Mediterranean area there is some leap-frogging, since the subspecies which breed in Italy (tschnsii) and North Africa (mauretanica) are mostly resident, while birds of the nominate race breeding farther north winter farther south, with very few exceptions (Vaurie 1959, 1965). There are occasional records of Wrynecks wintering north of the Mediter­ ranean in areas where they are not known to breed, such as the Camargue in southern France (Penot 1962, A. R. Johnson in litt.), Cornwall (Booth et al. 1968) and Pembrokeshire (Donovan 1968). Migration from European breeding grounds is directed approxi­ mately south-west or SSW in autumn and the reverse on return in spring. In Britain and Ireland most migrants are recorded in autumn, after weather conditions conducive to falls of Continental migrants, whereas in spring numbers are smaller but have tended to fluctuate much less from year to year. Thus in the years 1954-58 and 1964-65 the mean annual number of migrants in autumn was 113, with 394 in the peak year, whereas in spring the mean was 23, with a maxi­ mum of 30 (Monk 1963, Peal 1968). In autumn most are found on the east coast, particularly in Norfolk and Suffolk but also regularly in Yorkshire, Northumberland, Shetland, Kent and Essex; in spring this species is most commonly reported in Kent and Shetland. Spring migration is at its height in Egypt in the last week of 68 Wryneck studies March (Meinertzhagen 1930) and in Spain from 1st to 20th April (Bernis 1970). In Switzerland the main arrivals are between about 5th and 22nd April (Dr E. Sutter in Glutz von Blotzheim 1962), and in England breeding birds used to appear mainly from the first week of April until mid-May (Witherby et al. 1938-41). Farther north they come later. More than 100 birds killed at Danish lights were all between 10th April and 21st May (Hansen 1954) and Danish breeding birds arrive at the end of April or early in May (Salomonsen 1967). The mean dates of first arrivals in south and central Sweden are 6th and 8th May respectively (Rendahl 1964). About 240 km ESE of Moscow they arrive in the second half of April (Stephan 1961) and in the Leningrad area mostly in the first half of May (Velichko 1963). The bird shown on plate 14a was caught at the oasis of Azraq, Jordan, on 18th April 1965, retrapped there five days later, and found dead on 9th May at Kozelets, mid-way between Chernigov and Kiev in the U.S.S.R., about 2,100 km NNW of Azraq (Ferguson-Lees 1966). The commonest breeding habitats are orchards, parkland and other places with scattered trees; the birds will breed at the edge of woodland but seldom in dense woods, except by a ride or other clearing. They are hole-nesters but very rarely excavate their own nest-holes, although Abro (1962) twice saw Wrynecks greatly extending holes previously used by other species. The holes in which they nest may be natural or excavated, and they use nestboxes readily; much less commonly they will occupy crevices in walls or buildings, and even holes in the ground. The search for a nest-site begins soon after arrival when most hole-nesting species have already started to breed. There is often a shortage of suitable holes and Wrynecks will take possession of the nests of other hole-breeders, destroying their eggs or young and often ejecting nest material. Species displaced have included Great Spotted Dendrocopos major, Great Tit Parus major (the commonest victim), Blue Tit P. caeruleus, Coal Tit P. ater, Marsh Tit P. palustris, Willow Tit P. montanus, Nuthatch Sitta europaea, Redstart Phoenicurus phoenicurus, Pied Flycatcher Ficedula hypoleuca, Collared Flycatcher F. albicollis, Starling Sturnus vulgaris, House Sparrow Passer domesticus and Tree Sparrow P. montanus (Lohrl 1940, Banner- man 1955, Abro 1962, Busse and Gotzman 1962, Creutz 1962, Dekhuijzen-Maasland et al. 1962, Menzel 1968, Ruge 1971). Menzel saw one Wryneck remove the eggs of another. Oppenoorth (1938) reported an attack by a Wryneck on a nestbox inhabited by tits, in which both Wryneck and tit were killed. The Spotted Fly­ catcher Muscicapa striata is among the species which, according to Bannerman (1955), Boutillier (1914) stated were often evicted, but in the original reference the species of flycatcher was not given. Wryneck studies 69 Wrynecks may disturb a number of nests, and Lohrl (1940) stated that a single bird had destroyed 14 clutches in one year in an orchard at Stuttgart; he took six Wrynecks, of which four had been seen disturbing other nests, and five of these were males. Both sexes search for a nest-hole and draw the partner's attention to a possible site by calling from within or in front of it (Ruge 1971), as in plate 15a. After laying an egg, the female leaves the nest and flies on to a tree or fence and calls: the male answers, calling increases and copulation often ensues. The clutch can consist of as many as 14 eggs. Incubation takes about 12 or 13 days and the young fly from the nest at, very approximately, 21 days old. Faeces are passed after or between feeds and put by the young at the edge of the nest. At first they are not taken away and are probably consumed by the adults, but after the young are about seven days old the parents normally remove the faeces from the nest until shortly before the young leave (see plate 3 in Klaver 1964). Sometimes they are not removed, probably when the nest-hole is too crowded for the adults to reach them (see Bussmann 1941, Ruge 1971). Second broods are not rare and there is one record of three broods in one summer near Antwerp (Arnhem i960). The size of clutches and broods tends to decrease as the breeding season progresses. Incubation of a second clutch in another nest may begin even while young of the first brood are still in their nest (Szocs 1942, Arnhem i960). Wrynecks feed predominantly on ants. Detailed studies in Europe, outside Russia, of food given to nestlings have shown that it consists almost entirely of ants, and the breeding site is usually near a place where there are a number of mound nests of these . This dependence on a single type of prey makes the Wryneck vulnerable to any changes adversely affecting the number of ants or their availability. In Norway Eric Hosking (verbally and in Hancock 1965) saw adult and pupal wood ants Formica being taken to a nest- box with young, and in plate 13 a ball of pupae and some adult ants can be seen in and on the bird's beak. In studies in Belgium (R. A. G. Arnhem in litt.), the Netherlands (Klaver 1964) and Germany (Konig 1961, Dornbusch 1968, W. Keil in litt.), however, the Wrynecks have taken no wood ants to the young (except one imago of F. rufa with other ants) but instead other, generally smaller, species, even where wood ants have been common. In England it is probable that they prey predominantly on Lasius niger (Black Lawn ) and L. flavus (Yellow Ant). In Russia, although ants are the preferred food, it would appear that young are successfully reared on other insects (Pokrovskaya 1963, Strokov 1963). There is a small population of Wrynecks in the Netherlands, almost all known breeding there being in nestboxes. Broods of at least 70 Wryneck studies nine have been reared there in boxes intended for tits. One such box measured 93 X 121 X 232 mm internally, with an entrance hole 36 mm in diameter, but it is desirable to have a larger floor with a concave base if a nestbox is intended specifically for Wrynecks. The British Trust for Ornithology recommends a floor measuring 115 mm X 150 mm and an oval entrance hole 35 mm X 45 mm (Flegg and Glue 1971). Plate 11 illustrates a brood in the Hoge Veluwe near Arnhem, Netherlands, which had been moved into a glass-backed nestbox adjacent to the one in which they had hatched (Klaver 1964). The photograph was taken on 8th July 1963. On 10th June there had been two eggs (one undersized and infertile), and the adults were believed to have deserted another nest with eggs two days previously. On 27th June there had been six young about one day old and five eggs. Eggs are normally laid at daily intervals in the early morning, so clearly incubation had begun before the clutch was completed. (Incubation of first clutches usually begins when the full number of eggs has been laid, or a little before; with later clutches or with particularly large first clutches it often starts earlier.) The brood included one runt, easily recognisable in the fore­ ground, which Klaver stated then seemed four to five days old, most of the others being twelve days old. A runt is not uncommon widi hole-nesting birds and, if food is in short supply, it will not usually survive long (Lohrl 1968), but in this brood nine young fledged, including the runt which left the nest six days after most of the brood. 'Heel' pads are clearly visible on the runt in plate 11: these are found on nestlings of some piriform and other species. The 'song' of the Wryneck, a short, far-carrying, repeated 'kee- kee . . .', is given, often at frequent intervals, until the start of incubation or the completion of the clutch, and also when a sub­ sequent clutch is being laid. Unpaired birds may continue calling until about the end ofJune . In some cases, however, a pair may call little, and E. G. Philp (verbally), in whose garden in Kent Wrynecks bred in 1966 and 1967, heard calling very rarely in the second year. It may be infrequent when a pair has selected a suitable nest site and there are no other pairs in the neighbourhood.

ACKNOWLEDGEMENTS I am indebted to Mrs Cornelia Grindle, M. R. E. Ruffer and Commander R. D. M. W. Thomas-Ferrand for checking translations, and to my wife for much assistance, including typing the drafts of this paper.

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