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.     

Intercolony variations in movement patterns and foraging behaviors among herring gulls (Larus argentatus) breeding in the eastern Wadden Sea

Herring gulls (Larus argentatus) are opportunistic predators that prefer to forage in the intertidal zone, but an increasing degree of terrestrial foraging has recently been observed. We therefore aimed to analyze the factors influencing foraging behavior and diet composition in the German Wadden Sea. Gulls from three breeding colonies on islands at different distances from the mainland were equipped with GPS data loggers during the incubation seasons in 2012–2015. Logger data were analyzed for 37 individuals, including 1,115 foraging trips. Herring gulls breeding on the island furthest from the mainland had shorter trips (mean total distance = 12.3 km; mean maximum distance = 4.2 km) and preferred to feed on the tidal flats close to the colony, mainly feeding on common cockles (Cerastoderma edule) and shore crabs (Carcinus maenas). In contrast, herring gulls breeding close to the mainland carried out trips with a mean total distance of 26.7 km (mean maximum distance = 9.2 km). These gulls fed on the neobiotic razor clams (Ensis leei) in the intertidal zone, and a larger proportion of time was spent in distant terrestrial habitats on the mainland, feeding on earthworms. δ¹³C and δ¹⁵N values were higher at the colony furthest from the mainland and confirmed a geographical gradient in foraging strategy. Analyses of logger data, pellets, and stable isotopes revealed that herring gulls preferred to forage in intertidal habitats close to the breeding colony, but shifted to terrestrial habitats on the mainland as the tide rose and during the daytime. Reduced prey availability in the vicinity of the breeding colony might force herring gulls to switch to feed on razor clams in the intertidal zone or to use distant terrestrial habitats. Herring gulls may thus act as an indicator for the state of the intertidal system close to their breeding colony.     

Wie richtet eine Flußseeschwalbenkolonie (Sterna hirundo) ihr Abwehrverhalten auf den Feinddruck durch Silbermöwen (Larus argentatus) ein?

How a Common Tern (Sterna hirundo) Colony Defends itself against Herring Gulls (Larus argentatus) The subject of this study is the anti-predator behaviour of a small common tern colony near a large herring gull colony on the island of Mellum, West Germany (Fig. 1). In 1980 the number of gulls crossing this tern colony increased during the terns' chick-stage (Fig. 4). Observed predation of tern chicks was independent of tide and time of day (3., Fig. 5). The frequency of tern reactions corresponded to the number of herring gull crossings (Fig. 5, Table 1). The terns' responses increased between morning and evening (Fig. 8). Tern up-flights and attacks increased absolutely and as a percentage, with the advance of the breeding season (Fig. 3, 4). They were positively correlated with the observed chick predation and the number of pairs with chicks, most markedly with chicks older than 5 days (Figs. 3, 4; Table 1). This increased defence was maintained by fewer pairs as, by then, many had lost their own broods (Fig. 4). As the breeding season progressed, herring gulls increasingly became the main cause of tern up-flights and the object of the attacks (Figs. 9–11). The up-flights of the whole colony, which occurred frequently and spontaneously during incubation, were observed only rarely after hatching and were almost exclusively a response to herring gulls (Figs. 10, 12). The lower herring gulls flew over the colony, the more frequently common terns flew up or attacked and the more individuals were involved in these responses (Figs. 6, 13, 14). During the breeding period, communal up-flights and attacks by terns increased as a percentage (Figs. 12, 13, 15–17). Group-attacks effected changes in the gulls' flying-routes more often than did individual attacks (Fig. 18). Despite the defence behaviour and its adaptation to the predation pressure, herring gulls often succeeded in robbing chicks. This is why the breeding success of the common tern was poor (< 0.4 chicks/nest). Possible reasons for this are discussed.     

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.     

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.     

MIGRATION AND DISPERSAL OF AUDOUIN'S GULL LARUS AUDOUINII FROM THE EBRO DELTA COLONY

Oro, D. & Martinez, A. 1994. Migration and dispersal of Audouin's Gull Larus audouinii from the Ebro Delta colony. Ostrich 65:225-230.

Migratory paths and dispersal patterns of Audouin's Gulls (Larus audouinii) born in the Ebro Delta have been studied, as well as their degree of philopatry. The gulls always disperse southwards from the colony, and only the juveniles seem to disperse in other directions. Dispersal patterns of 2y, 3y and older gulls are similar, wintering mainly on Iberian Mediterranean and Moroccan Atlantic coasts. Juveniles migrate before the rest of the age classes and they winter south of older gulls, in the Senegambia region. However, conclusions on Audouin's Gulls movements are difficult to draw, owing to the large differences in resighting and recovery efforts. Data presented also suggest that Audouin's Gulls do not appear to breed before the age of four years.     

A comparison of Herring Gull Larus argentatus and Lesser Black-backed Gull Larus fuscus nest sites: their characteristics and relationships with breeding success

Nest sites used by Herring Gulls and Lesser Black-backed Gulls in a mixed colony were compared. Correlations between features surrounding a nest and breeding success at different stages of the breeding cycle are presented. Lesser Black-backed Gulls nested on more vegetated and flatter areas than did Herring Gulls, even though the latter species had a lower hatching success at less vegetated sites. The difference in the general topography of nest sites between the species suggests that the Lesser Black-backed Gull, through an individual defence strategy, may be better adapted to use sites which are accessible to ground predators than the Herring Gull. The fledging success of Lesser Black-backed Gulls tended to increase with an increased proportion of relatively tall vegetation close to the nest. The Lesser Black-backed Gull may therefore be able to leave its young unattended and rely on their concealment for protection against predators. In contrast, the Herring Gull may rely more on parental vigilance to protect young. More frequent attendance by adult Herring Gulls at the nest site during chick rearing compared with Lesser Black-backed Gulls supports this hypothesis.     

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.     

Summer in the city: behaviour of large gulls visiting an urban area during the breeding season

Capsule: Large numbers of Herring Gulls Larus argentatus and Lesser Black-backed Gulls Larus fuscus from a traditional colony in the Netherlands visit an urban area for food in the chick rearing period, causing nuisance while doing so.

Aims: To assess the potential contribution of large gulls breeding in a traditional colony to gull–human conflicts in cities.

Methods: Colour-ringed gulls from a nearby colony were counted in the study area throughout the 2014 breeding season. The average numbers observed daily per species and sex were compared between different breeding phases.

Results: Fluctuations in numbers of both species could be explained by breeding phase. Numbers of females of both species and male Lesser Black-backed Gulls dropped significantly during laying and incubation. Numbers peaked post-hatching, coinciding with increased engagement in nuisance events and more frequent displacements within the study area.

Conclusion: Large gulls from a traditional colony frequently visited the urban study area, especially when food demand for chick provisioning was high, suggesting that city visits are motivated by accessibility of urban food. The proportion of rooftop breeding gulls in this area is low. Therefore, measures to avoid nuisance should focus on reducing food availability rather than controlling gull breeding.     

ADAPTATIONS TO NESTING HABITAT IN THE REPRODUCTIVE BEHAVIOUR OF THE BLACK-BILLED GULL LARUS BULLERI

SUMMARY The nesting habitats adopted by most colonies of Black-billed Gulls Larus bulleri are river-beds that are subject to flooding. A number of respects in which the reproductive behaviour of Black-billed Gulls differs from that of at least most other gulls, such as Black-headed Gulls, can be viewed as adaptations, or byproducts of adaptations, to such nesting habitats:—

• (a) A different breeding site from the year before is often selected
• (b) The bulk of pair formation is accomplished before the gulls occupy their breeding sites; nesting territories are set up by mated pairs
• (c) In hostile encounters during the pair formation phase, site attachments are weak or transitory
• (d) In agonistic situations generally, attack thresholds appear to be relatively high, and fleeing thresholds relatively low
• (e) High intensity forms of“Choking” appear to be missing from the agonistic display repertoire
• (f) The growth of nest groups is rapid and orderly; nesting territories are small and the concentration of nests in the groups high
• (g) The time between occupation of the gullery site and the start of laying is short, and the synchrony of laying is high; this is probably related to the close proximity of the nests
• (h) The parents and young abandon the nest very soon after the egg hatch, and no “brood” nests are made
• (i) The young may develop locomotory powers more quickly than is the case in most other species; they can swim at an early age, and take to the water in tightly packed groups during alarms

The question of why these gulls should choose such vulnerable breeding sites is discussed.     

Reproduction and survival of Glaucous Gulls breeding in an Arctic seabird colony

ABSTRACT.   Despite being widespread and easily observed, little is known about the life history of Glaucous Gulls ( Larus hyperboreus ). From 1984 to 2007, we examined their breeding biology and demography at Coats Island, Nunavut, Canada, where they nest alongside a colony of 30,000 pairs of Thick-billed Murres ( Uria lomvia ). The gulls fed mainly on murre eggs and chicks and by scavenging adult carcasses. The median age at first breeding was 5 yr, and the mean age was 4.8 ± 0.9 yr. Adult survival was estimated as 0.84 ± 0.03 (SE). The mean clutch size was 2.56 eggs and the mean number of young reared per year was 1.6 (range = 0.9–2.2). Birds reared at the colony provided 40% of recruits. Assuming that survival of locally reared chicks that emigrated was similar to that of chicks that returned to the colony, about 22% of the young gulls survived to breeding age. The timing of breeding by Glaucous Gulls appeared related to the timing of laying by murres. Although the demographic characteristics of Glaucous Gulls in our study were similar to those of populations of other large gulls, adult survival was at the lower end of the range for populations of large Larus gulls. There is some evidence that Glaucous Gulls exhibit lower survival than large gulls breeding in temperate areas, possibly because of contaminant burdens. In general, however, the demographic characteristics of large gulls show little variation and are probably a product of their common phylogeny.     

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.     

SELECTION OF COLONY SITES AND NEST SITES BY COMMON TERNS STERNA HIRUNDO IN OCEAN COUNTY, NEW JERSEY

Common Terns nested on 34 of 259 saltmarsh islands along 46 miles of Ocean County coastline, New Jersey. They nested on low islands of Spartina with less than 12%, windrow ranging in size from 0.6 to 108 acres. All islands selected by terns faced at least two miles of open water from at least one direction. Selection of islands seemed a response to vegetation, size, distance to the nearest island, distance to the nearest shore and exposure to open water. Of the 225 islands without nesting terns, only three fulfilled appropriate criteria. Eighty per cent of nests were situated on windrow. Terns laid larger clutches on windrow compated to those nesting in Spartina. Nearest neighbour distance on islands with low predation varied from 85 to 485 cm and was a function of space, vegetation, and the size of the island. Terns nested closer together on windrow compared with Spartina. Considering all colonies, the mean nearest neighbour distance negatively correlated with the number of nests. Skimmers, Oystercatchers, Laughing Gulls, and Herring Gulls nested in some of the tern colonies. All tern colonies on islands also occupied by nesting Herring Gulls suffered over 10% egg-loss by predation. Proportionally more nests were preyed upon in windrow than in Spartina. Clutch sizes in nests in windrow on islands with predation were significantly lower than those in windrow on islands without predation, even though clutch sizes in Spartina were similar on islands with and without predation. Flooding by exceptional high tides destroyed significantly more nests in Spartina than in windrow. The nests that survived in Spartina were built deeper, and their rim heights were higher than nests destroyed by tides. Choice of colony and nest sites is discussed in terms of the balance of two selection pressures: predation and tidal flooding. Tern nests on windrow are more exposed to predators, but safer from flooding, whereas those nests in Spartina are more susceptible to flooding and less vulnerable to predators. The recent invasion of Herring Gulls into Ocean County has significantly increased the rate of nest predation among affected colonies of Common Terns. It is possible that in future years the terns' behaviour may be modified in response to this new selective pressure.     

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.     

Management-Induced Reproductive Failure and Breeding Dispersal in Double-Crested Cormorants on Lake Champlain

ABSTRACT We studied breeding dispersal of double-crested cormorants (Phalacrocorax auritus) associated with management practices that suppressed their reproduction on Lake Champlain in the northeastern United States. We implemented an experiment on one colony by spraying corn oil on cormorant eggs in portions of the colony and leaving other portions untreated. Gulls (Larus spp.) consumed cormorant eggs during the oiling process, but we reduced and then eliminated predation levels after the first year of the study. We used mark-recapture techniques within the experimental framework to measure rates of breeding dispersal for cormorants from the experimental colony and an unmanaged colony in Lake Champlain. Egg oiling increased the movement rate to the unmanaged colony by 3% during the year with no egg predation by gulls. When gulls depredated cormorant eggs at high rates during egg oiling, movement to the unmanaged colony increased by 20%. When cormorants are managed to reduce population sizes, methods that limit dispersal away from the managed colony may be most effective. Such methods would mitigate effects to nontarget populations and allow for a greater portion of the metapopulation to be managed.     

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.

     

COLONY AND HABITAT SELECTION OF SIX KELP GULL LARUS DOMINICANUS COLONIES IN SOUTH AFRICA

Kelp Gulls nested in six colonies on rocky islands, sand and rock cliffs, sand dunes and on a sand, gravel island in a salt lake. The colony sites selected were different from the surrounding areas and were generally inaccessible to ground predators. Kelp Gulls nested in a wide variety of habitats. Within colonies, the gulls did not nest randomly with respect to habitats but preferred to nest on flat, stable areas with some cover (either rocks or vegetation). They generally avoided heavily vegetated areas, areas devoid of vegetation and areas with steep slopes. Their specific habitat choices relate to predation and cannibalism pressures. Intermediate cover provides adequate protection for chicks while allowing for increased visibility and open escape routes for parents. Nearest neighbour distances were similar among colonies and habitats, indicating strong social attraction.     

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.     

Earthworms in the diet of Herring Gulls Larus argentatus breeding on an off-shore island

ABSTRACT

Pellets of indigestible material regurgitated by Herring Gulls Larus argentatus breeding on Lady Isle, Firth of Clyde, Scotland, were collected in 2018 and 2019 and examined for earthworm chaetae. Nearly two-thirds (65.6%) of the 314 pellets came from gulls that had consumed one or more earthworms. Significantly fewer pellets (57.6%) contained chaetae in 2018, a relatively dry May to July, than in 2019 (72.0%) when rainfall was close to the long-term average for May to July. There were significant associations between the presence of large quantities of terrestrial vegetation in the pellet and the detection of large numbers of earthworm chaetae and/or fragments of terrestrial arthropods, suggesting that recent consumption of these food items is likely when pellets contain large quantities of vegetation.     

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.