Intermittent breeding in the Herring Gull Larus argentatus and the Lesser Black-backed Gull Larus fuscus

Thirty-three percent and 37% of adult Herring Gulls Larus argentatus and 34% and 40% of adult Lesser Black-backed Gulls Larus fuscus , all with some previous breeding experience, failed to breed in 1993 and 1994, respectively. About half of these intermittent breeders of both species failed to breed in both years. Common amongst the intermittent breeders were smaller individuals, birds with low survival chances and relatively unsuccessful breeders.     

Sex-biased mortality in the Lesser Black-backed Gull Larus fuscus during the nestling stage

Theory predicts that skewed progeny sex ratios should be relatively common in vertebrate populations. In most birds this has proved hard to substantiate due to the difficulties associated with identifying the sex of large samples of chicks. This study reports the success of a new molecular DNA technique in determining the sex of 601 newly-hatched Lesser Black-backed Gulls Larus fuscus
There was no evidence of any adaptive sex ratio within broods. Male chicks were found to be disproportionately large and to grow at a faster rate than females. The overall sex ratio changed significantly from 0.484 (male/male + female) at hatching to 0.399 by fledging, probably due to male susceptibility to starvation. Mortality also increased significantly with hatching order, an effect often observed in species like the Lesser Black-backed Gull where hatching is asynchronous. I discuss the possibility that hatching asynchrony may in fact be a strategy employed to prevent excessive skews developing in progeny sex ratio whenever variable differential mortality is likely.
The results appear to vindicate Fisher's (1930) hypothesis which predicts the overproduction of the 'cheaper' sex. However, as the skewed sex ratio may be determined more by unpredictable environmental factors, such as food supply and weather conditions rather than parental strategy, this interpretation should be treated with caution.     

Nest predation and nest site selection among Eiders Somateria mollissima: the influence of gulls

Hooded Crows Corvus cornix, Great Black-backed Gulls Larus marinus and Herring Gulls L. argentatus were the main nest predators in an Eider population in southwest Sweden. The clutch sizes of Eider nests within gull colonies did not differ from those outside gull colonies. The proportion of Eider nests destroyed by predators was significantly lower within than outside gull colonies, especially on islands with Lesser Black-backed Gulls L. fuscus. Although the difference was not significant, the survival time of simulated Eider nests was higher within than outside gull colonies. On Eider islands with gull colonies, foraying crows spent more time within the colony area than expected by chance. However, crows apparently avoided an area around each gull nest, and we suggest that the colonies, to some extent, protected Eider nests against predation. The density of Eider nests was higher on gull islands than on gull- free islands, and higher within than outside the gull colonies. However, the association with gulls was weak compared to that displayed by some other waterfowl.     

Timing of primary moult in adult Herring Gulls and Lesser Black-backed Gulls

Summary Timing of primary moult in relation to the breeding cycle is presented for 113 actively moulting adult Herring Gulls and 79 Lesser Black-backed Gulls from Walney Island, England. Moult in both species occurred about when the eggs hatched in mid-May. The entire Herring Gull population began to moult the primaries within a period of 50 days. Lesser Black-backed Gulls started to moult 10 days later than Herring Gulls but many birds were not yet moulting as late as 5. August when I left the study area. It is suggested that the population of Lesser Black-backed Gulls consists of residents and migrants and that the former begin to moult earlier than the latter.

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Der zeitliche Ablauf der Handschwingenmauser von Silber- und Heringsmöwe

Zusammenfassung Bei 133 ad. Silber- und 79 ad. Heringsmöwen von Walney Island, England, wurde der Beginn der Handschwingenmauser in Beziehung zum Brutzyklus untersucht. Bei beiden Arten setzte die Mauser ungefähr Mitte Mai zum Zeitpunkt des Schlüpfens der Jungen ein. Alte Silbermöwen begannen innerhalb von 50 Tagen mit der Handschwingenmauser. Die Heringsmöwe fing etwa 10 Tage später an, doch hatten manche Individuen noch bis Anfang August nicht mit der Mauser begonnen. Die Heringsmöwen-Population auf Walney besteht vermutlich aus Stand- und Zugvögeln; erstere scheinen früher zu mausern.

     

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.     

The behaviour of seabirds foraging at fishing boats around Shetland

Among the different types of fishing vessels around Shetland, whitefish trawlers attract the largest numbers of scavenging seabirds and provide the most food. Offal was almost all consumed by seabirds, predominantly by Fulmars Fulmarus glacialis , which excluded other species by their aggression. Fulmars generally ignored discarded whole fish, which were mainly taken by Great Black-backed Gulls Larus marinus , Gannets Sula bassana and Great Skuas Catharacta skua . Although flatfish were usually ignored because seabirds found them difficult to swallow and they sank faster, most discarded roundfish were consumed. Herring Gulls L. argentatus , Lesser Black-backed Gulls L. fuscus and Kittiwakes Rissa tridactyla were rarely able to obtain offal or discards. Herring Gulls and Lesser Black-backed Gulls spent much time on the periphery of feeding flocks while Kittiwakes rarely attempted even to join these. Most of the birds at trawlers were in adult plumage, and it is suggested that the low proportion of immature birds present was a further reflection of the highly competitive feeding conditions at trawlers. We suggest that likely changes in fishing practice and seabird population sizes in the immediate future may result in Herring Gulls, Lesser Black-backed Gulls and Great Skuas finding feeding on waste around trawlers increasingly difficult, so they may be further displaced by Fulmars, Gannets and Great Black-backed Gulls.     

SPECIES ISOLATION BETWEEN THE HERRING GULL LARUS ARGENTATUS AND LESSER BUCK-BACKED GULL L. FUSCUS

There is a large, mixed colony of the two “ring” species, the Herring Gull Larus argentatus and Lesser Black-backed Gull L. fuscus, on Walney Island, northwest Lancashire. These birds are nesting at the very high density of one nest/40 square yards, or more, but although they defend their territories against both species indiscriminately, there is effectively no hybridization. This paper discusses the nature of the species isolation mechanism, and its function. Since the two species can produce fertile hybrids, the mechanism must be of an ecological/behavioural nature, rather than morphological incompatibility. It is shown that there are slight differences in breeding season and habitat, but these do not seem to be great enough to account for the high degree of isolation. It is likely that species isolation depends primarily on the female's choice of a mate. It is suggested that, as specific cues, she uses the differences in call-note tones, and the colour of the back (and perhaps also of the eye-ring), or both. Herring Gulls and Lesser Black-backs are adapted to slightly different niches. The overlap is so great, however, that any hybrid is unlikely to be at a disadvantage; but the overlap can only have arisen very recently, as a result of both species taking advantage of the increased availability of human refuse. It is possible that the isolation mechanism was evolved to cope with earlier conditions, when food was more limited, and the species' niches more sharply defined.     

Genetic Affinities Within the Herring Gull Larus argentatus Assemblage Revealed by AFLP Genotyping

To date, the taxonomic status of circumpolar breeding populations of the Herring Gull Larus argentatus, the Lesser Black-backed Gull Larus fuscus, and the closely related Yellow-legged Gull Larus cachinnans has been based on differences or similarities in phenotype, morphology, and feeding and premating behavior. To shed some new light on the many taxonomic uncertainties surrounding these taxa, we describe the results of a large DNA study based on comparing the distribution of 209 biallelic markers among 109 gulls, representing 11 gull taxa of the Herring Gull assemblage and the Common Gull Larus canus. A detailed phylogenetic analysis failed to show clustering of individuals into groups representing either geographic origin or phenotype. Alternatively, birds were grouped into taxa defined on the basis of phenotype and geographic origin or phenotype alone. Genetic analyses revealed significantly different genetic distances between all pairs of taxa. However, based on these genetic distances, again no consistent phylogenetic tree could be constructed. Analysis of molecular variance indicated that about 77% of the total genetic variability among these gulls could be explained by within-taxon differences. Only 23% of the total genetic variability was due to genetic differences between taxa, irrespective of their species or subspecies status. Although this seems to challenge the current taxonomic treatment of the herring gull assemblage, our results are too premature and too incomplete to recommend a drastic change. Received: 27 January 2000 / Accepted: 18 September 2000     

Disturbance-dependent Yellow-legged Gull (Larus michahellis) predation on Larid chicks decreases with chick age

Predation is one of the key factors shaping the dynamics of animal populations. In birds, nest loss due to predation can be a significant cause of low reproductive success. Ground-nesting birds are among the bird groups most susceptible to predation, mainly because their nests are easily accessible to a broad suite of potential predators. For these birds, anthropogenic disturbances can generate changes in nest predation risk by altering their antipredator behaviour and also by altering the behaviour of the predator species, i.e. the predator becoming much more aware of predation opportunities due to frequent disturbances and/or motivated to repeat predation attempts when some are successful. To date, most previous studies investigating this have focused on a single effect, either predation or disturbance, on chick survival. It remains unknown how the risk of predation with and without disturbance varies with chick age. In this study, we used behavioural observations to assess how the interaction between predators and disturbance affects predation risk in chicks and how this interacts with chick age. Specifically, we investigated the effect of disturbance caused by humans and stray dogs on the predation of Slender-billed Gull Chroicocephalus genei chicks by Yellow-legged Gulls Larus michahellis, and whether this depended on the age of the chicks. Our results revealed that disturbance had a significant positive effect on predation measures of Slender-billed Gull chicks by Yellow-legged Gulls, but that this effect was mediated both by disturbance type and the age of chicks. Stray dogs entering the colony had a stronger disturbance effect on chicks than passing humans, increasing predation risk by Yellow-legged Gulls. Our results also showed that chick age interacts with disturbance type to determine the predation risk. This is probably mediated by chicks' capacity to escape predation by gathering in a single large crèche that runs into the water when disturbed. To preserve Slender-billed Gull colonies in one of its few remaining breeding sites in Tunisia, and as gulls tend to react even when the disturbance occurs relatively far from the colonies, it is crucial to (1) restrict human access to dikes and islets where large colonies breed and (2) construct artificial islets attractive to gulls and inaccessible to stray dogs.     

Nonbreeding and nests without eggs in the Lesser Black-backed Gull Larus fuscus

Estimates of the number of breeding Lesser Black-backed Gulls Larus fuscus at the Tarnbrook Fell gullery, Lancashire, have been made from annual counts of nests since 1981. During all of these surveys, the mean percentage of nests which did not contain eggs has been remarkably constant from year to year (54%± 1.9%). A study of specific nests at the colony in 1992 showed that the great majority of these "empty" nests did not subsequently receive eggs. in 1993, a study of a sample of territory-holding gulls was made on a 0.41-ha study plot to investigate and to quantify the occurrence of empty nests. Of the 62 pairs that defended a territory and constructed a complete nest, 27% subsequently failed to produce eggs. These birds attended their territory as a pair significantly less frequently than pairs that produced clutches and achieved significantly fewer apparently successful mountings. Sixty percent of pairs constructed more than one nest, and 58% of all nests built received no eggs. The construction of empty nests was not found to be related to the density of nesting gulls. Since 1981, estimates of the number of gulls in the colony have been based on the assumption that one nest represents one pair of gulls. This has been shown not to be the case, and a correction multiplier of 0.61 must be applied to counts of nests at the colony to obtain an estimate of the number of breeding gulls.     

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.     

Effects of cleaning a tidal river of sewage on gull numbers: a before-and-after study of the River Tyne,northeast England

For most of the 20th century, the tidal reaches of the River Tyne in northeast England were used as an open sewer for organic wastes from almost a million people. This produced anaerobic conditions in the river, but supplied an appreciable source of food for five species of gulls. Observations on the number of gulls using the river were made in 1969/70 to establish the situation before a sewage treatment scheme was implemented. By 1994, the volume of untreated sewage discharged into the river had been reduced by 97% and by 86% in the study area, and a second survey was carried out to evaluate the effect of the clean-up on gull numbers. Overall, gull numbers showed a significant decrease of 37% between the two study periods. The decrease in winter was 38% and was also significant, while the decline in summer was 33%, but not significant. Considering individual species, numbers of Common and Great Black-backed Gulls declined by 93% and 91% respectively. In both these cases this is presumed to be due to the decline in available food. Black-headed Gull numbers did not change significantly and it appears they have been able to find alternative feeding sites such as mud flats exposed at low tide and the outfall where treated effluent is discharged. Numbers of Lesser Black-backed Gulls and Kittiwakes rose dramatically as a result of increases in breeding numbers in the area. Neither species feeds along the river. Numbers of Herring Gulls did not change significantly — it is likely that the effect of the decline in food availability was masked by the increase in breeding numbers in towns along the river banks. The different response of each of the species of gulls indicates the differences in their ecology and breeding status in the area.     

Possible effects of local enrichment by gulls on feeding-site selection by wintering Barnacle Geese Branta leucopsis

The selection of salt-marsh feeding sites by wintering Barnacle Geese Branta leucopsis was investigated on Schiermonnikoog, The Netherlands. Use of sites by geese was estimated by counts of goose droppings. Geese spent significantly more time grazing in sites with evidence of Herring Gull Larus argentatus and Lesser Black-backed Gull Lfuscus breeding activity than in nearby non-gull sites. The nitrogen content of red fescue grass Festuca rubra was significantly greater in gull sites. We suggest that the geese preferred to feed in these locations because of the improved nutritional value of the grass. The higher nitrogen content of F. rubra in gull sites was most likely the result of nutrient inputs from gull droppings rather than goose droppings.     

Intra-population variation in the diet of an avian top predator: generalist and specialist foraging in Great Black-backed Gulls Larus marinus

ABSTRACT

Capsule: Great Black-backed Gulls Larus marinus breeding on Skokholm, UK, fed predominantly on seabirds, rabbits, refuse, and marine prey, with the majority of pairs being dietary generalists, but with some specialist pairs.

Aims: To understand the significance of Great Black-backed Gulls as top predators on a small offshore island with internationally important numbers of breeding seabirds (Skokholm, UK) by quantifying their diet and to determine how this varies within the breeding season, to test for pair-level dietary specialization and to examine the consequences of dietary differences for reproductive performance.

Methods: Regurgitated pellets were collected and analysed from 26 breeding pairs on Skokholm during 2017 and related to breeding success.

Results: Analysis of 1035 pellets revealed that, overall, Great Black-backed Gulls fed on seabirds (48% – mostly Manx Shearwaters Puffinus puffinus), mammals (38% – mostly European Rabbits Oryctolagus cuniculus), anthropogenic waste (7%), and marine prey (7%). Diet varied among pairs with 18 (73%) generalist pairs and 7 (27%) specialist pairs (of which, 5 were bird specialists and 2 were mammal specialists). Diet also varied seasonally, but pair-level dietary diversity was repeatable through the breeding season. Dietary diversity did not covary with breeding success.

Conclusion: Great Black-backed Gulls are top predators on Skokholm. Variation in diet among pairs emphasizes that not all individuals contribute equally in terms of predation. Understanding the incidence of this variation has important ecological implications, particularly where apex predators may exert a strong top-down influence.     

Habitat selection and breeding success in Yellow-legged Gulls Larus cachinnans

The habitat selection and breeding performance of Yellow-legged Gulls Larus cachinnans were studied in the Medes Islands colony, northeastern Spain, during 1995 and 1996. Of the three main habitats on the islands (shrubs, grass and bare areas), gulls first occupied those with the highest percentage of tall vegetation. Gulls tended to select nest sites with 20–75% cover despite great differences in the cover in the habitats and territories, suggesting that the presence of a suitable nest site may play a major role in the choice of breeding habitat. Nest-site tenacity did not influence the preferences of gulls at any level since the same pattern of choice was observed in an area subjected to annual culls (i.e. where most of the breeding pairs were culled annually and replaced by naive birds). In spite of great differences in the physical characteristics of the habitats, little difference was found in breeding performance of the gulls between habitats. Gulls nesting in the least preferred habitat (i.e. mainly bare) had smaller clutches than those nesting in the other two habitats, possibly as a result of their later seasonal laying. Despite the similar breeding success in different habitats, gulls did not seem to distribute according to the ideal free model reported for Herring Gulls Larus argentatus since the density in the preferred habitat (i.e. shrubs) was never higher than in the other two. We suggest that the habitat selection by Yellow-legged Gulls within the colony could follow an ideal despotic distribution.     

Experimental evidence for the relationship between food supply, parental effort and chick survival in the Lesser Black-backed Gull Larus fuscus

We studied breeding success, chick growth, parental effort and chick behaviour in two groups of Lesser Black-backed Gulls Larus fuscus whose chicks were provided with additional food until 7 days after hatching or until fledging. These data were compared with those from control pairs which we studied simultaneously to test the hypotheses that food was in short supply during the chick stage at the colony site and that in such circumstances the behaviour of adults and young is mainly responsible for the low success. Pairs whose chicks were fed with additional food until fledging showed a higher fledging success than control pairs (intermediate for pairs of first experimental group). During the first week after hatching, experimental adults of both groups were present together at the territory for longer than control pairs. In adult females of experimental pairs, the length of feeding trips was shorter than in females of control pairs, whilst the rate of chick feeding was more frequent in the experimental broods. After the chicks were 7 days old, differences were significant only for the experimental pairs whose chicks were provided with additional food until fledging. Chicks fed until fledging showed a higher daily mass and wing-length increments and reached a higher fledging mass at an earlier age than both control chicks and chicks which were provided with additional food until day 7. Starvation occurred only in control chicks and in chicks of the first experimental group after we had stopped providing food. When food was in short supply, fledging success of gulls was adversely affected as a result of both starvation (because of the lower feeding rates of chicks) and a higher predation rate (arising from changes in behaviour of both adults and chicks).     

The Post-Fledging Mortality of the Blackbird in Great Britain

Some 3,000 pairs of Herring Gulls are now known to be nesting on buildings, mainly in coastal towns they are increasing most rapidly in eastern Britain, where recruitment is taking place from natural colonies. A smaller number of Lesser Black-backed Gulls has occupied rooftops in the Bristol Channel region.     

Expansion of the Mediterranean Gull Larus melanocephalus in Poland

Since 1981, when the first breeding pair of Mediterranean Gull Larus melanocephalus was recorded in Poland, the population of this gull has increased considerably. Its population size was stable until 1997, not exceeding ten pairs annually; thereafter, an increasing number of sightings were made, and during the last 5 years between 26 and 39 breeding pairs have been recorded in Poland. To date, breeding sites have been established in a total of 45 sites (maximum of 19 sites in a given year), with 27% of these (43% of all broods) found on islands located in the middle course of the Vistula River. Breeding sites have also included artificial reservoirs, such as dam reservoirs (20% of sites, 19% of broods), gravel pits (9% of sites, 13% of broods) and fishponds (24% of places, 8% of broods). Mediterranean Gulls were found to nest only within the colonies of other Laridae. Single pairs were recorded at 40% of the breeding sites, whereas a maximum of two to five pairs were recorded at 47% of the other sites. The great majority of breeding attempts were recorded in Black-headed Gull L. ridibundus colonies. In two cases, Mediterranean Gulls bred within mono-specific colonies of Common Gulls L. canus consisting of 60–150 pairs. The biggest concentrations of breeding sites were along the middle course of the Vistula river and in the southern part of Poland.     

Lesser Black-backed Gulls Larus fuscus thriving on a non-marine diet

Capsule: Lesser Black-backed Gulls Larus fuscus breeding 30?km from the coast in the Netherlands focussed entirely on terrestrial food sources and reached relatively high breeding success.

Aim: To gain insight in the foraging ecology, habitat use and breeding performance of inland-breeding Lesser Black-backed Gulls.

Methods: We received data from seven birds fitted with global positioning system (GPS) loggers. The colony was frequently visited to collect pellets and boluses and to monitor reproductive success, mortality and growth rate of chicks.

Results: The GPS data revealed that mainly terrestrial habitats were used, 98% of these GPS positions were within 25?km of the colony. Refuse dumps were the most preferred sites, but also agricultural fields and freshwater bodies were often visited. Only two of the 710 recorded trips were directed to the North Sea. The pellet and bolus analyses confirmed the GPS data: no marine food remains were found. Breeding success of birds in the enclosure was relatively high, with 90% of eggs hatched and 51% of chicks fledged (1.6 chicks/pair).

Conclusions: Relying on terrestrial food is feasible when sources are available in the vicinity of the colony. We conclude that Lesser Black-backed Gulls could theoretically shift towards inland breeding after a fishery discards ban.