THE EFFECTS OF DIURNAL AND TIDAL PERIODICITIES IN THE NUMBERS AND ACTIVITIES OF HERRING GULLS LARUS ARGENTATUS IN A COLONY

The average numbers of Herring Gulls Larus argentatus present in a breeding colony on Walney Island, Cumbria, were found to vary with the tidal cycle but to remain effectively constant with time of day through the breeding season. An activity survey, based on 50 Herring Gulls observed at half-hourly intervals during March and April 1973, showed that sleep and rest varied inversely with each other with sleep increasing to 50 per cent at midday. After a peak in the proportion of gulls asleep four hours before low tide, sleeping progressively decreased until low tide; seemingly a result of resident gulls waking and remaining more alert as others left the colony in search of food. Preening was constant throughout the day and tide cycle. Other behaviours (mostly courtship and agonistic behaviour associated with territory defence) increased slightly during low tide and were more common early and late in the day. Night observations of the gulls' activities showed that there was a peak of sleeping between midnight and 02.00 hours. It is suggested that Herring Gulls have a bimodal diel sleep pattern.     

NESTING DENSITY AND BREEDING SUCCESS IN THE HERRING GULL LARUS ARGENTATUS

The relationship between nesting density and breeding success of Herring Gulls Larus argentatus was studied on the Isle of May, Scotland, in 1968. Herring Gulls nesting at the most common density started laying earlier in the season than those nesting at lower or higher densities. Therefore, although the overall spacing of nests was uniform, the nest density of birds laying later in the season progressively approached a random distribution. The onset of laying occurred in synchronized groups within the colony. Late-laid clutches were commonly situated on the periphery of the colony where the density of nests was lowest. When the laying period was divided into four time periods, in each period the tendancy was for birds nesting at the most common density to have the highest clutch-size, hatching and fledging success, and to rear the most chicks per pair to fledging. In addition, birds which spaced their nests most uniformly, presumably as a consequence of territorial behaviour, were the most successful parents.     

ASPECTS OF THE BREEDING BIOLOGY OF HERRING GULLS LARUS ARGENTATUS IN URBAN COLONIES

Herring Gulls nesting on rooftops in South Shields and Sunderland were observed during 1974–76. These colonies, then around 200 pairs in each town, have expanded rapidly despite culls. Ringing results identified the Forth area as an important source of recruits, and also suggested that females are more likely to breed away from their natal area than males. It is clear from the high breeding success that these towns provide favourable alternatives to more typical breeding sites. The gulls in the towns nest at low densities, other than on flat roofs. Their breeding success, averaging 1.2–1.6 chicks per pair, was higher than that recorded at more typical colonies, and on structurally isolated rooftop sites was almost double that on flat roofs. This high success is attributed to the absence of cannibalism in these small colonies and the lack of territorial aggression at the structurally isolated sites.     

HYBRIDIZATION OF GLAUCOUS GULLS LARUS HYPERBOREUS AND HERRING GULLS L. ARGENTATUS IN ICELAND

Until recently, the Herring Gull Larus argentatus and the Glaucous Gull L. hyperboreus were completely allopatric in western Europe, although having overlapping distributions in Canada and the Soviet Union. But beginning about 1925–1930 a large-scale invasion of argentatus from the Faroe Islands and/or Britain into Iceland occurred, resulting in extensive hybridization with hyperboreus, already present there. The hybrids are viable and apparently fully fertile. In the summers of 1964–1966 371 gulls were collected from breeding colonies in various parts of Iceland and the samples analysed by the use of hybrid indices based on the colour of the five outermost primaries. The populations of northwestern Iceland, where the greatest concentration of hyperboreus was found at the time of the argentatus invasion, were found still to consist largely of pure hyperboreus, with 18–23% of the birds showing signs of argentatus admixture. In eastern Iceland, where argentatus first appeared in numbers, and along the south coast, birds of hybrid origin predominate, 3–43% of the birds being pure argentatus while pure hyperboreus are absent except for one colony in southeastern Iceland where they made up 23% of the birds. Significant differences in composition are found among several of the colonies of eastern and southern Iceland, and possible reasons for this are discussed. Among 53 mated pairs no deviation from random matings with respect to hybrid indices was indicated. The high proportion of pure argentatus found in some colonies in spite of this indicates that pure argentatus are still immigrating from abroad. Two colonies in eastern Iceland sampled both in 1965 and 1966 showed a significant change in composition towards argentatus, further evidence for continued immigration of argentatus. A few other colonies have changed in the same direction over longer periods. Although there must be some differences between the two forms in ecological requirements as shown by their distributions, their ecologies appear similar in Iceland. Both forms take a large portion of their food from the intertidal zone of the sea-shore, and although a slight difference in the selection of foods from the shore was seen in winter in one locality, this probably did not have a genetic basis. However, argentatus may feed to a larger extent on refuse than hyperboreus. This may enable argentatus-like birds to occupy the relatively barren (with respect to intertidal foods) east coast of Iceland in numbers. An examination of several hundred museum skins of argentatus and hyperboreus from other areas shows that occasional hybridization occurred in western Europe long before the extensive contact was established in Iceland, but gene flow between the two was always limited as their ranges were widely separated. The two gulls are effectively reproductively isolated from each other where synipatric in Canada, while the situations in Alaska and in parts of the Soviet Union are unclear. Since the area of overlap without hybridization is much larger than the area of hybridization, the two forms are best regarded as separate species. Future studies may show in which direction the hybrid situation in Iceland will develop, but meanwhile it is suggested that the hybrid populations in which birds of hybrid origin amount to 50% or more be referred to as Larus argentatus/hyperboreus.     

BREEDING SUCCESS AND POPULATION GROWTH IN A COLONY OF HERRING AND LESSER BLACK-BACKED GULLS LARUS ARGENTATUS AND L. FUSCUS

This paper describes the results of investigations into the factors affecting breeding success of the Herring and Lesser Black-backed Gulls Larus argentatus and L. fuscus, in the large colony on Walney Island, northwest Lancashire, between 1962 and 1965. These investigations were concerned with the incubation period, and the first ten days after hatching. The survival of chicks to ten days is 67% in Herring Gulls, and 56% in Lesser Black-backs. Most of these losses occur in the period just after hatching and are due to “cannibalism” by other gulls. This form of predation does not appear to be masking any effects from starvation or disease. The following factors contribute to egg or chick mortality:breeding too late (and, to a much smaller extent, too early); breeding in the open, as opposed to amongst cover; the facts that eggs in small clutches have a lower hatching success than those in large ones and that Herring Gull (but not Lesser Black-back) chicks in small broods are less likely to survive to ten days than are those in large broods. Chick mortality after the first ten days is not certainly known. About 30% of the eggs laid gave rise to fledged young— or about one fledged chick per pair. In the Herring Gull, the average clutch size (2.56) is lower than that of the Lesser Black-back (2.76). Both species show a seasonal decline in clutch size—this occurs earlier in the Herring Gulls than in the Lesser Black-backs. The Walney population, which stood at about 700 pairs in 1950, had reached 12,000 in 1957, and is at present about 18–19,000 pairs. It is suggested that this increase may be linked to the greater availability, or exploitation, of human garbage in the Morecambe Bay area. The population explosion between 1950 and 1957 must have been partly due to massive immigration and could not have come about through natural increase alone. The possible influences of the gulls' behaviour on the population growth are discussed. There is no evidence of any “shock disease”, although the Walney colony is very crowded. “Cannibalism” is regarded, not as evidence of a failing food supply, but as an extension of the normal hunting behaviour of these omnivorous gulls; it will be an economical means of obtaining food only in a large, dense colony, such as Walney. It may be offset by increased breeding efficiency due to social factors.     

DISEASE IN A COLONY OF BLACK-HEADED GULLS LARUS RIDIBUNDUS

An account is given of the disease factors encountered in a colony of Black-headed Gulls in the breeding season 1956.     

AGE, EGG-SIZE AND BREEDING SUCCESS IN THE HERRING GULL LARUS ARGENTATUS

The breeding biology of the Herring Gull Larus argentatus was studied in 1969, 1970 and 1972 on Skokholm, Pembrokeshire. The distribution of laying dates in three years is given; median laying date was always about 10 May. Laying dates of individual pairs were fairly constant between two years, but females up to nine years of age apparently advanced their laying date in successive years. In 1969 there was a decline in clutch-volume index during the season but this was not found in 1970 or 1972. Females laid clutches of similar volume in successive years, although the volume did vary with age, increasing up to 7 or 8 years of age and then declining. Hatching success was highest in the earliest of four periods in 1970 (69%) and lowest in the last (51%); in 1972 the corresponding figures were 71% and 63% and the seasonal trend was not significant. Chick mortality was higher in 1970 (69%) than in 1972 (59%); in 1970 there was significant seasonal variation in chick survival but in 1972 there was not. In 1970 chick survival was positively correlated with hatching weight (and hence egg-size) and chick survival was also related to growth which was itself correlated with hatching weight. There were no such correlations in 1972. Mean chick production in 1970 was 0.60 per pair and in 1972 0.70. In both years later breeders were less successful, but the seasonal range of mean success was less in 1972. In the discussion it is shown that some of these differences between years could be due to changes in the age structure of the population and it is suggested that the correlation between egg-size and chick survival in 1970 may be an artefact of the age structure.     

COLONY DESERTION AND REPRODUCTIVE SYNCHRONY OF BLACK-BILLED GULLS LARUS BULLERI

Black-billed Gulls breeding in large dense colonies on river beds in New Zealand were found to have a high rate of colony desertion early in the nesting cycle. The hypothesis is developed that individuals faced with possible desertion after they have begun to nest can reduce the likelihood of being deserted by delaying their nesting activities until a large number of others is also ready to nest. This interpretation is developed into a graphic model to show that colony desertion can thereby produce a strong selection pressure favouring highly synchronized nesting. This model successfully predicts onset synchrony, subcolony formation, occurrence of complete colony desertions mainly early in the season, occurrence of partial colony desertions, and the existence of nester non-nester conflict over whether or not to desert. Unstable food appears to be the most likely cause of colony desertion in Black-billed Gulls. Black-billed Gulls are similar, in varying degrees to Sandwich, Royal and White-fronted Terns. Convergent traits include tendencies to occupy colonies rapidly and synchronously, reduced site tenacity, and the use of open and often unstable nesting habitats. The links described here between desertion and synchrony may be relevant for some other species.     

THE FOOD OF SOME LARUS GULLS

The food of L. argentatus on Skomer and Skokholm Islands (Wales) was studied by the analysis of stomach contents, pellets and direct observation. L. argentatus is mainly a scavenger and exists for the most part on fish-waste and garbage discarded by man. At Ncwborough Warren argentatus feeds more on arable land and on the shore than do the Skokholm and Skomer birds.
Although argentatus frequently eats limpets Patella spp., there is little competition for food between them and Oystercatchers Haematopus ostralegus because the two species eat different sized limpets.
A summary of previous studies into the food of argentatus is given.
L. marinus is mainly a predator feeding on the young of other gulls, sea-birds and Rabbits but it also eats carrion, offal and fish-waste when available and during the winter. The numbers of Manx Shearwaters Procellaria puffinus killed on Skokholm and Skomer are discussed.
L. fuscus obtains most of its food on the shore and on arable land. There appears to be little competition, at least in the areas studied, between .fuscus and argentatus for food.