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.     

EFFECT OF AGE ON THE FISHING ABILITY OF SANDWICH TERNS STERNA SANDVICENSIS

The fishing ability of first-winter and older Sandwich Terns was compared at foraging sites used by both age-groups on the coast of Sierra Leone between January and March 1970 when first-winter birds were about 7–9 months old. The methods used for measuring various parameters of fishing ability are described. It was found that a significantly larger proportion of plunge dives by the older age-group yielded prey although the difference between the two age-groups was only slight.
There were no major differences between the age-groups in diving rates or in size of fish captured. Older birds tended to dive from higher than first-winter birds.
By driving shoals of small fish to the surface, predatory fish enhanced the fishing success of both age-groups on certain days. Where man's beach seining operations yielded a superabundance of small fish near the tideline, these were captured equally well by both age-groups.
First-winter birds were not seen being fed by older birds, and although food-begging behaviour was observed once in late January it is clear that the first-winter birds were self-sufficient for food by the time this study was made.
Some implications of the observed differences in fishing success are discussed. It is suggested that, after fledging, Sandwich Terns gradually gain the judgment and skill necessary to dive profitably from increasing heights and so learn to exploit prey in a greater range of water depth as they get older.     

TIMING AND SPACING OF BROODS IN THE BLACK-HEADED GULL LARUS RIDIBUNDUS

The nests of the Black-headed Gull Larus ridibundus are closely aggregated into dense colonies and their use synchronized, these two phenomena together tending to produce a maximal clumping effect. Within such a colony however, nests were found to be spaced out to produce a non-random uniform distribution. The commonest distance between neighbouring nests was found to be about one metre, in contrast to related species. This study was concerned with two aspects of this distribution pattern; its survival value and its behavioural causation.
It was found that pairs nesting just outside the colony had a much lower breeding success than those nesting in the colony and that nests on the colony fringe had a slightly lower success than those in the centre. Pairs laying during the peak laying period had a higher breeding success than pairs laying either earlier or later in the season. Since by far the most important mortality agent was predation, it seems likely that both clustering and synchronization of nesting function as antipredator systems and arguments in favour of this are discussed.
Variations in nest-spacing within the colony were not correlated with variations in breeding success.
In the causation of the spacing between nests, territorial aggression was demonstrated to be an effective dispersion mechanism and the way in which this mechanism works was investigated in detail.
This spacing mechanism was not sufficient by itself to explain the observed densities, which were higher than one would expect from the aggression alone; there was also some tendency for birds establishing a new nest-site to cluster close to others. The interaction between this, the territorial aggression of the residents and the subsequent avoidance responses of the settling birds, can explain the nest spacing pattern and probably also the observed densities.     

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.     

PIRACY BY LAUGHING GULLS LARUS ATRICILLA: AN EXAMPLE OF THE SELFISH GROUP

In 1969, 1500 pairs of terns Sterna spp. and 50 pairs of Laughing Gulls Larus atricilla nested on Petit Manan Island, Maine, U.S.A. The gulls formed groups of up to eleven individuals to steal fish from the terns. The start of piracy coincided with hatching of the gull's eggs. Larger groups more often succeeded in stealing fish than smaller groups. Group formation cannot be ascribed to rarity of terns to chase, and is not due to shared advantage in fish seized or time spent chasing (hence energy expended). A sufficient explanation is the individual advantage to some gulls that join a small group at the expense of the initiator. Individual gulls are likely to follow different strategies. Chasing appeared to be more readily elicited by a chase in progress than by a tern carrying a fish.     

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 BREEDING ECOLOGY OF ROYAL TERNS STERNA (THALASSEUS) MAXIMA MAXIMA

The breeding ecology of the New World race of the Royal Tern Sterna maxima maxima was studied at colonies in Virginia and North Carolina, U.S.A., from 1967 to 1970. Colony sites are quite varied, but isolation, good distance visibility and (especially) freedom from quadruped predators seem important if not essential prerequisites. In Virginia, most adults arrive at the colony site in the last few days of March. Courtship, displaying and copulation take place at, near, and some distance from the colony site. In this the Royal Tern differs from its near relative S. sandvicensis sandvicensis which carries out these activities away from the colony site, presumably as an anti-predator device. Courtship displays are not described, but in their essentials are similar to those of other terns. Copulations continue throughout incubation and gradually disappear when the eggs hatch. No post-copulatory displays are known. Some synchrony of egg-laying is evident, but no “mass laying” occurs, the colony increasing in size steadily over a period of weeks and months. Defaecation on the nest is normal and probably serves to strengthen the nest rim against flooding by high tides. Contrary to published reports, the normal clutch is one; the largest is two, probably often from two different females. All incubating adults examined had two brood patches. Average egg measurements are 63 × 44-5 mm, and average egg weight is 64.3 g. Egg colour varies greatly, and evaluation is difficult. Eggs are probably not cryptically coloured and individual variation, as well as nest-site, are used by returning adults to identify their own eggs. Average maximum nest density is c. 7/m². Sterna sandvicensis acuflavida nests regularly, if not obligatorily, with m. maxima; interactions between the two, and possible benefits accruing to each, are discussed. Unhatched eggs were significantly nearer other nests than were successfully hatched eggs, and possible explanations are given. Incubation lasts about 30–31 days, a week longer than in most terns; both sexes incubate. Broken eggshells are never removed by the adults. Instead, after 2–3 days, the chicks leave the nest permanently to join a creche that roams freely about the immediate vicinity of the colony. Chicks remain in the creche for about 25–30 days, leaving it at fledging, approximately 30 days after hatching. While in the creche, chicks are normally fed only by their parents, who probably recognise them both vocally and visually, using the extensive variation in voice and colour of chicks characteristic of the species. Sandvicensis acuflavida chicks also are highly variable, and join Royal creches, adults of both species attending. Variation also occurs in Royals' juvenal plumage, and seems associated with extended parental care. Feeding adults normally wander up to 40 km from their colony, and this probably facilitates the intercolonial exchange of breeding birds we recorded. They feed inshore, in shallow waters, taking small fish by dives which do not go below the surface. They regularly take small soft-shell crabs in this way, and frequently water-skim like skimmers Rynchops spp., sometimes capturing food while doing so. A relationship between crab capture and water-skimming is established for the first time, and water-skimming in non-feeding contexts is mentioned. Besides the quadruped predators which they normally avoid by fleeing the colony site, Royals have no known predators beyond the egg stage. Eggs, although not chicks, are readily taken by Laughing Gulls. Relationships between the two species are discussed, emphasising their constant association the year round.     

FACTORS AFFECTING THE EGG SIZE OF RED-BILLED GULLS LARUS NOVAEHOLLANDIAE SCOPULINUS

The factors influencing the egg size of the Red-billed Gull Larus novaehollandiae scopulinus were studied at Kaikoura, New Zealand, between 1964 and 1972. In two- and three-egg clutches there was a trend for the eggs to become smaller in the sequence of laying. Length, breadth and volume of eggs of one-, two- and three-egg clutches declined significantly as the season progressed. The size of eggs from single-egg clutches tended to be smaller than eggs from two-egg clutches laid at the same time. There were correlations between the proportions of one-egg and of three-egg clutches being laid at a given period and the mean egg volume of two-egg clutches. When the mean egg volume of two-egg clutches increased there was a corresponding increase in the proportion of two- and three-egg clutches laid. When the mean egg volume of two-egg clutches decreased there was an increase in the proportion of single-egg clutches laid. The egg size of the Red-billed Gull showed no direct correlation with the abundance or availability of food; the largest eggs were produced early in the season when food was in short supply. In spite of an increase in the food supply in the middle of the breeding season, birds laying at this time produced smaller eggs than birds which laid earlier in the season. However, early breeders which relayed at the peak in food abundance on average produced a larger replacement clutch than originals laid early in the season. It is suggested that the birds nesting early in the season are able to produce the largest eggs because they are the most efficient foragers for food, and those which nest later in the season produce smaller eggs, even at peak food abundance, because of their inefficiency or inexperience. Early breeders laying replacement clutches tended to lay larger eggs and larger clutches than birds which are producing their first clutches at the same time. Two-year-old females laid eggs which were significantly shorter than older aged birds while the breadth and volume of the egg increased with the age of the female up to the fifth year. There was a trend for females to lay larger eggs when mated with older rather than younger males. No statistical differences in egg size were detected between females changing or retaining the partner of the previous season. Female body weight and egg volume were positively correlated in females weighing less than 275 g but not for heavier females. It is suggested that the seasonal decline in egg size and clutch size results from a decrease in the availability of food and the ability of the individual to exploit the resource.     

THE PROCESS OF COLONY FORMATION AMONG HERRING GULLS LARUS ARGENTATUS NESTING IN NEW JERSEY

We examined the pattern of colony occupation and egg-laying in five colonies of Herring Gulls nesting in New Jersey, U.S.A. Colonies formed from epicentres located in sparse bushes. The number of epicentres related to the number of birds nesting in the colonies. Colonies of over 250 pairs had more than one epicentre, whereas those with under 250 pairs had only one epicentre. Epicentres were not always in the geographical centre of the colonies. New territory-hunting pairs filled in the epicentre areas, and then nested outside these areas. The egg-laying pattern followed the settling pattern, but was more synchronous than the settling pattern. There was greater synchrony of egg-laying within sub-areas of the colonies than in the colonies as a whole. Further, synchrony correlated with the number of nests in sub-areas.     

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.