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.     

THE MOULT OF REMIGES AND RECTRICES IN GREAT BLACK-BACKED GULLS LARUS MARZNUS AND GLAUCOUS GULLS L. HYPERBOREUS IN ICELAND

The moult of primaries, secondaries, and rectrices in two closely-related gulls, the Great Black-backed Gull Larus mavinus and the Glaucous Gull L. hyperboreus, was studied in Iceland. Both gulls moult their primaries in an extremely regular sequence, starting with the 1st (innermost) and ending with the 10th (oiltermost) feather. Usually two, less often one or three, primaries are growing per wing during the primary moult, which lasts for about six or seven months. Growlng primaries were estimated to lengthen on the average by 8.7 mm per day in marinus and 7.8 mm per day in hyperboreus. The secondaries, usually 24 in number, are shed in two moult waves, one starting with the innermost feather soon after the start of the primary moult and then progressing slowly outwards, the other beginning with the outermost secondary after the primary moult is about half completed and then progressing rapidly inwards. The moult is completed just before the end of the primary moult as the two moult waves meet at about the 16th secondary. There are no marked differences between the two gulls in the moult of secondaries. The moult of rectrices shows large variations in both species, some feathers being much more irregular than others in their time of shedding. In both species, indications of an obscured centrifugal pattern of replacement are seen, although the 5th (next to the outermost) rectrix is usually the last one to be shed. Significant differences were observed between the two species in the degree of regularity of shedding of some feathers and in the average position in the moulting sequence of others. The moult of rectrices starts soon after the moult of primaries is half completed. The feathers are then shed in rapid succession, and the moult is completed some time before the end of the primary moult. The need for good powers of flight at all times is undoubtedly the reason for the protracted primary moult. This in turn causes the moult to start early, in adults sotnetimes before the eggs are laid; immatures moult even earlier than this. The rectrix moult and the main part of the secondary moult do not begin in adults until the young have fledged, but then progress very rapidly. Presumably, the loss of some of these feathers would impair the flying ability to an extent sufficient to make it difficult for the gulls to care for their young, while the rapid moult is necessary in order for the replacement of these feathers to be completed by the time the primary moult is over.     

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.     

SOCIAL STIMULATION OF REPRODUCTION IN HERRING AND LESSER BLACK-BACKED GULLS

The date of the first egg laid and the position of nests were recorded for all pairs in 10 study areas within the mixed Herring and Lesser Black-backed Gull colony on the southern end of Walney Island, Lancashire. The study areas differed from one another in density of breeding birds and in proportion of each species. These data were collected in order to determine if evidence could be found for synchrony of reproduction by social stimulation (the "Fraser Darling Effect"). The gulls in different study areas showed considerable variability in onset and spread of laying; however, attempts to correlate a number of variables, such as density of breeding birds in each area with median date or spread of laying, average nearest neighbour distance with median date of egg-laying, and median laying date for one species with that of the other species in the same section of the gullery, failed to produce convincing evidence for inter- or intraspeciiic social stimulation of reproduction.     

INTRASPECIFIC PREDATION AND COLONIAL BREEDING IN LESSER BLACK-BACKED GULLS LARUS FUSCUS

The breeding biology of Lesser Black-backed Gulls was studied on Skokholm Island, Pembrokeshire, where the number of breeding pairs has been increasing at about 20% per annum since 1963. Laying was found to be synchronous within small groups. Clutch size and breeding success showed seasonal declines over the spread of breeding. The loss mainly of eggs, but to a lesser extent of chicks also, caused this overall decline in success. Hide observations indicated that the bulk of these losses arose through predation by nesting adult Lesser Black-backed Gulls on their nearest neighbours. Not infrequently the protagonist had lost its own clutch shortly before turning predator, and such a chain-sequence should lead to a steady build-up of aggressive failed breeders and so account for the observed seasonal increase in egg loss. Attentiveness to the clutch decreased with season but this was unlikely to have been important in precipitating the predation. Intermediate plant cover was associated with highest nest density and also with highest chick survival. In addition, nest density was directly correlated with chick survival. Whether plant cover and nest density separately affect chick production remains unresolved. Nevertheless, the increasing nest densities in this colony caused by the growth of the gull population are thought to be responsible for the widespread intraspecific predation, the intensity of which is probably a new feature of the gulls' breeding behaviour. The implications of this 'internal' predation for laying synchrony and aggregated nesting are discussed; these two factors of the breeding pattern probably evolved largely to combat 'external' predators. Not only are they no protection against inter-neighbour predation but appear to facilitate it. It remains to be seen whether this kind of predation will significantly affect breeding patterns with further increases in nest density.     

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.     

蛇体和蛇胆乙醇浸提液的营养成份分析

对五种蛇的蛇体及蛇胆的乙醇浸出液的营养成份进行了分析,并对两种浸出液进行了毒性试验。结果表示,蛇体乙醇浸出液含有二种氨基酸,四种维生素;蛇胆乙醇浸出液除含四种维生素外,还含有四种微量元素。毒性试验结果表明,供试动物主要脏器及血液学检查均未见任何异常变化。     

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.     

POPULATION SIZE,DISTRIBUTION AND DISPERSAL OF KELP GULLS IN THE SOUTHWESTERN CAPE,SOUTH AFRICA

Steele, W.K. & Hockey, P.A.R. 1990. Population size, distribution and dispersal of Kelp Gulls in the southwestern Cape, South Africa. Ostrich 61:97-106.

There are indications that the Kelp Gull Larus dominicanus population of the southwestern Cape Province, South Africa, is increasing. It is proposed that decreased post-fledging mortality due to supplementary food from man's activities, at sites such as fishing harbours and refuse dumps, is likely to be the main cause of this population increase. Kelp Gulls aggregate, and juveniles (first-year birds) are disproportionately abundant, at sites where supplementary food is available. Juvenile dispersal from the two largest breeding colonies, 38 km apart, is in opposite directions.     

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.