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in Britain or on the Continent and recovered from December to February in each of 10 regions of Britain and Ireland (4,267 recoveries in total) were used to produce an index of the proportions of geographical distribution in each area (Fig. 33). The percentages of Continental birds in these different regions ranged from 7 per cent to 80 per cent, based on at least 115 recoveries in each region and, in most cases, many more. The wide range of percentages obtained suggests that they are a close approximation to the actual proportions of birds of Continental origin. Ireland, Scotland and Cumbria had low proportions of Continental gulls, while the highest proportions occurred in south Wales, southern England, and eastern England from Yorkshire southwards. Overall, about 71 per cent of the wintering gulls in Britain and Ireland originated from the Continent – a high value, but consistent with the much higher overall numbers of Black-headed Gulls reported here compared with estimates in the breeding season.

      FIG 33. Estimates of the percentage of Continental Black-headed Gulls present in 10 regions of Britain and Ireland in winter based on numbers of recoveries of British and Continental ringed birds in each region between early December and the end of February.

      Sex ratio of wintering birds

      While the sex ratio of Herring Gulls and Great Black-backed Gulls (Larus marinus) captured in Britain during the winter approaches equality, that of wintering Black-headed Gulls shows a marked skew, with many more males being present. The data in Table 13 show that birds identified at breeding sites in northern England in May and June contained a minor excess of 108 males per 100 females. However, the sex ratio of wintering birds captured in England changed considerably, with a threefold excess of males in October and November, followed by the reversion of proportions to near equality between December to February. In the winter samples, most individuals were visitors from the Continent, and the skewed sex ratio in October and November suggests that either migrating males were arriving earlier than the females by an average of a month or more, or that males and females were feeding at different types of sites not sampled in those months, but were feeding at the same sites later in the winter. A total of 20 males and 11 females wing-tagged in the autumn were subsequently reported to have returned to the Continent, which gives some support for the suggestion that the bimodal pattern of the arrival time of Continental gulls (Fig. 32) represents males migrating earlier than females.

      BREEDING BIOLOGY

      Black-headed Gulls build their nest on the ground in areas with low-growing vegetation of varying density, ranging from sand dunes with much bare ground, to taller and floating vegetation around tarns and lakes. However, a few exceptions have been reported. At Seamew Crag, a small islet on Lake Windermere, Clive Hartley and Robin Sellers recorded several of the 50 pairs in the colony there nesting on top of low bushes over several years following 2009 (pers. comm.). In East Anglia in 1947, an entire colony numbering more than 300 pairs switched to nesting 2–3 m above ground level in young spruce trees, apparently in response to the flooding of their usual nest sites on the ground (Vine & Sergeant, 1948). Unlike some other gull species, there is only one old record of Black-headed Gulls nesting on a building, but in 2015 Robin Sellers found three small groups doing so, two near Perth and the other at Montrose, both in Scotland (pers. comm.).

      Philopatry and colony faithfulness

      Most Black-headed Gulls are two years old before they breed for the first time and a minority are a year older before they breed. Exceptionally, one-year-old individuals attempt to breed, although their success is very low. Many of those that have survived to maturity return to breed in the colony in which they hatched as chicks (called philopatry), but others move to other colonies, often some distance away. The high proportion returning to the natal colony indicates that the young birds retain a good memory of where they were reared. However, it is easier to find marked individuals that have returned to their original colony than those that have moved elsewhere and consequently the extent of philopatry is often exaggerated. A realistic estimate of the extent of philopatry is obtained by measuring the proportion of those ringed as chicks and have reached adult age that were recovered during the breeding season less than 20 km from their natal site. In the case of the Black-headed Gull, about 60 per cent of the young that survive to breeding age are philopatric and the remaining 40 per cent move to other colonies and in a few cases, across the North Sea, to breed on the Continent. There have also been exchanges of individuals between Ireland and Britain (Fig. 34). Once adults have bred in a colony, their attachment to it becomes high, and at least 85 per cent of those surviving to the following year return to it to breed. The few adults that move elsewhere to breed are mainly from colonies that are in the process of being deserted, in some cases as a reaction to repeated nesting failure and the presence of mammalian predators.

      FIG 34. Movements of more than 20 km of Black-headed Gulls ringed as chicks in Britain and Ireland and recovered in the breeding season when of breeding age. Many returned to breed at or near where they were reared, but a similar proportion apparently moved to other colonies. A few moved to the Continent to breed, mainly between northern France and Denmark, but one moved to Germany, another to Norway and a third to northern Sweden. Reproduced from The Migration Atlas (Wernham et al. 2002), with permission from the BTO.

      Annual reoccupation of the colony

      Black-headed Gull colonies are first visited by groups of adults in March each year. They arrive at irregular intervals during the morning, flying over the site without landing, and do not remain long at the colony or in the vicinity. Eventually, on one such visit some birds do land, but they exhibit a degree of nervousness and are easily disturbed. These visits become more frequent, but will be curtailed or prevented by cold and windy weather. As the days pass, the daily presence of the birds at the colony lasts much longer and spreads into the afternoon, but the colony is always vacated before sunset. The gulls usually leave by a synchronous ‘up’ or ‘panic’ flight, which sees them all suddenly rising high above the colony as if alarmed, yet without an obvious stimulus such as the appearance of a predator. As April progress, the amount of time the gulls spend at potential nest sites extends to the greater part of the day, but the colony is still deserted each night and reoccupied early in the morning, sometimes well before sunrise.

      Pairs are formed early and courtship displays become common (Fig. 35). In the meantime, nesting material is collected locally and brought in to the colony. Birds nesting in some coastal colonies often collect substantial and untidy quantities of brown seaweed, while those at inland sites collect dry grass locally and carry it to the selected nest site to construct the nest. More material is usually added to the nest during incubation. Colonies and nesting sites are usually close to water, and nests are often substantial structures that raise the eggs above the local water levels.

      FIG 35. Female Black-headed Gull (right) courtship-begging for food from a male. (Norman Deans van Swelm)

      The density of nests varies according to the size of the colony and the nature of the nesting site. Nests are 1–2 m or more apart where plant growth is sparse, but only 50 cm apart in dense vegetation, presumably because this tends to conceal the neighbouring pair, moderating the extent of aggression between close neighbours.

      Eggs and incubation

      Black-headed Gull eggs

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