First records of lesser black-backed gulls Larus fuscus crossing the Sahara non-stop

Lesser black-backed gulls Larus fuscus migrate regularly along coastlines or the Nile to their wintering grounds in sub-Saharan Africa. They usually avoid the interior of the Sahara, but occur occasionally far inland when moving upstream along rivers. We discovered several flocks of lesser black-backed gulls crossing the western Sahara about 500 km east of the Atlantic coast. Their concentrated flight directions along a northeast-southwest axis indicate that the gulls were on an intended migratory flight and not incidentally drifted inland. Ground speed of 8 flocks of gulls averaged 20 m/s and was considerably larger than the mean air speed (12 m/s) demonstrating that the gulls were strongly assisted by tailwinds. Highest ground speed was 26 m/s. Warmest temperature encountered when crossing the study site was 13° C due to relative high flight altitudes of 3,500 m above sea level. Thus, lesser black-backed gulls migrated under favourable conditions with respect to energy and water consumption. Our observations strongly suggest that the gulls performed long non-stop flights between the eastern Atlantic coast of sub-Saharan Africa and the Mediterranean Sea.     

Urban and coastal breeding lesser black-backed gulls (Larus fuscus) segregate by foraging habitat

Despite urbanization's general erosion of biodiversity, towns and cities provide novel opportunities for some species. During the 20th century, gulls (Laridae) colonized urban areas around the world where they flourished. At the same time, some coastal populations declined. The reasons for this difference are not fully understood, partly because little is known about any ecological differences between urban and non-urban gulls, such as their foraging ecology. Here we compare the movement ecology and habitat selection of Lesser Black-backed Gulls Larus fuscus graellsii breeding at two neighbouring colonies – one urban and one coastal – in north-west England. We used bird-borne GPS loggers to first compare colony-level movement behaviour and habitat selection and then investigated individual-level habitat use. We observed clear colony-level habitat segregation: urban breeders preferentially foraged in urban areas whereas coastal breeders foraged primarily in coastal habitats and avoided urban areas. Coastal breeders also had larger core and home-ranges than urban breeders, possibly due to differences in colony size. However, we also found inter-individual differences in habitat use, which may have important management implications. These findings suggest a link between nesting and foraging ecology, and thus management or environmental change altering food availability will impact gulls at the coastal and urban sites differently.     

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.     

Interspecific differences in foraging preferences, breeding performance and demography in herring (Larus argentatus) and lesser black-backed gulls (Larus fuscus) at a mixed colony

Herring gulls Larus argentatus and lesser black-backed gulls Larus fuscus breeding at Walney Island, Cumbria, the largest breeding colony of the two species in the UK, have recently shown very different population trends. The former has declined sharply, whereas numbers of the latter have been maintained for several years. Here we compare aspects of the feeding and breeding ecology of the two species in order to examine whether or not this suggests explanations for their different population trends. Comparison of the ratio of the two species in flight lines leading to different feeding sites and their diet composition showed that the lesser black-backed gulls fed more at sea and the herring gulls fed more in the intertidal zone. Urban resources were used by both these species. These differences have been consistent over the last three decades. Susceptibility to death from botulism at the breeding colony was the same for the two species. The availability of the intertidal zone for foraging appears to have declined in recent years, and this may have had a more negative impact on the herring gull. However, the breeding success of the two species remains relatively high. This study suggests that differences in foraging behaviour and food availability during the breeding season are unlikely to be responsible for the marked differences in demographic trends in the two species. Changes in local food availability during the winter would be expected to have more effect on the resident herring gull. This work highlights the need for more detailed studies of the ecology of both species during the breeding season and in winter in regions showing differing patterns of population change.     

The effect of dehydration on salt and water balance in gulls (Larus argentatus and L. fuscus).

The effects of a variety of dehydratory stresses, hypersalinity, heat and water deprivation on renal and extrarenal handling of a standard salt load have been studied. Concommitant measurements of plasma and tissue electrolytes and of pituitary prolactin levels have also been made and the possible correlation between prolactin levels and the degree of dehydration are discussed along with the possible ecological significance of the reported findings.     

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.     

Egg-laying capacity is limited by carotenoid pigment availability in wild gulls Larus fuscus

In birds, experimentally increased egg production can reduce maternal condition, parenting ability and survival, and the quality of the eggs themselves. Such costs probably reflect resource limitation, but the identity of the resource(s) in question remains unclear. Carotenoids are antioxidants and immunomodulants that birds can only obtain in their diet. Trade-offs in the allocation of limiting carotenoids between somatic maintenance and egg production could therefore be an important factor underlying reproductive costs. We show that in wild lesser black-backed gulls, Larus fuscus, dietary carotenoid availability (i) constrained the capacity to re-lay following clutch removal; and (ii) affected the relationship between yolk mass and egg mass. However, whether carotenoids are limiting for egg production directly, by stimulating the synthesis or antioxidant protection of yolk precursors, or indirectly via effects on maternal health, requires further study.     

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.     

The effect of age and environment on extrarenal salt excretion in juvenile gulls (Larus argentatus and L. fuscus)

Young gulls captured from an essentially marine colony with no access to freshwater showed a significantly greater ability to excrete a hypertonic salt load than did birds of approximately the same age captured from an inland colony.
This ability was reflected by higher extrarenal secretion rates from the heavier nasal gland of the "marine" birds. That this increased ability was due to environmental exposure and not to genetic variations was shown by the fact that these differences disappeared when the two groups of birds were maintained on a freshwater diet for four weeks.
The main environmental factor stimulating the development of the nasal gland in the "marine" birds appeared to be non-availability of freshwater, as analysis of the diet of both colonies showed only minor differences.     

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.     

Maternal condition, yolk androgens and offspring performance: a supplemental feeding experiment in the lesser black-backed gull (Larus fuscus)

It has been proposed that the maternal androgens in avian egg yolk enhance offspring fitness by accelerating growth and improving competitive ability. Because egg quality is strongly influenced by maternal condition, we predicted that females in good condition would produce high-quality eggs with relatively high androgen content. We experimentally enhanced maternal condition by supplementary feeding lesser black-backed gulls (Larus fuscus) during egg formation and compared the concentrations of androstenedione (A4), 5alpha-dihydrotestosterone (DHT) and testosterone (T) in their eggs with those in eggs laid by control females. We also measured circulating levels of T in females immediately after laying. Egg androgens could affect offspring performance directly through chick development and/or indirectly through changes in the competitive ability of a chick relative to its siblings. To avoid confounding these two routes, and to separate effects operating through the egg itself with those operating through experimental changes in parental chick rearing capacity, we fostered eggs from both maternal treatment groups singly into the nests of unmanipulated parents. Contrary to expectation, mothers with experimentally enhanced body condition laid eggs with lower levels of androgens, while exhibiting higher circulating T concentrations post-laying. Despite these lower levels of egg androgen, offspring hatched from eggs laid by mothers in good condition did not show reduced growth or survival when reared in the absence of sibling competition. Our results demonstrate that yolk androgen concentrations vary with the body condition of the female at the time of egg formation and that females in good condition reduced the yolk androgen content of their eggs without altering offspring performance.     

Effects of experimentally increased egg production on female body condition and laying dates in the great skua Stercorarius skua

We investigated the effects of increased egg production on body condition as well as on measures of reproductive performance in great skuas, Stercorarius skua, over two subsequent years. We experimentally increased egg production from the normal two to six eggs. Six eggs might also be produced under natural circumstances after repeated clutch loss. After the production of the last egg we measured: (i) body mass, (ii) pectoral muscle, and (iii) haematocrit, total red blood cell count and mean corpuscular volume, as indicators of body condition. We took the same measurements of control females who had produced the normal clutch of two eggs. The measurements were repeated one year after the manipulation, and survival, laying dates, clutch sizes and hatching success were recorded for up to three consecutive years. After producing six eggs, females were lighter, had smaller pectoral muscles and lower haematological values than control females. Hatching success of eggs was significantly reduced. Even one year after the experiment there were still differences in body condition. Annual survival was not affected by the manipulations, although there was an indication that survival costs depended on whether chicks were raised after the increased egg production. While pair bonds and egg sizes were not affected in the post‐experimental year, females started breeding significantly later than in the previous year. Two years after the experiment laying dates had advanced again and were not different from those of control females. This pattern of maintaining survival and egg sizes, but delaying breeding in the post‐experimental year was found for two independent groups of females which had both been subjected to increased egg production. These results present evidence that increased egg production can have long‐term effects on female body condition and aspects of reproduction. However, although present, the costs of extra eggs appear to have been relatively small in the great skua in comparison to the two other bird species for which inter‐annual effects have been reported.     

A novel avian W chromosome DNA repeat sequence in the lesser black-backed gull (Larus fuscus)

The phenol emulsion reassociation technique was used to isolate and clone a female specific, repetitive DNA sequence fromLarus fuscus. The repeat, designated P2000-17, is restricted to the W chromosome, although related sequences occur elsewhere in the genome ofL. fuscus. Similar sequences were detected in the genome of six other bird species from outside the genus Laridae, but the sequence occurs less frequently and to a similar extent in both sexes. The 298 bp DNA sequence of P2000-17 was determined and found to have extensive sequence identity to the rabbit dihydropyridine (DHP) receptor calcium channel. P2000-17 is represented once within a larger 8.6 kb tandem repeat (LfW-1), which has a complex internal DNA sequence. LfW-1 is highly conserved between repeat motifs and may comprise 3% of the female genome. The possible evolutionary origin of LfW-1 is discussed in relation to the repeat types found on the W and Y chromosomes of other species.     

Impact of egg harvesting on breeding success of black-headed gulls,Larus ridibundus

Gull colonies world-wide have been harvested for their eggs for centuries with minimal knowledge of the impacts on breeding. Although most Laridae can replace lost eggs, they have comparatively high energetic demands for egg production. In this paper we assess the impacts of a licensed egg harvest on the breeding success of black-headed gulls Larus ridibundus, which nest colonially in an EU Special Protection Area in Hampshire, southern England. We compared egg volume, hatching and chick survival from harvested and un-harvested nests in central and fringe positions within colonies of various sizes, including colonies with no harvesting activity. Eggs from various laying stages were collected from harvested and un-harvested colonies of similar pre-harvest intrinsic quality, for comparison of their volumes, yolk-to-albumen ratios and eggshell thickness. Egg volume and the yolk-to-albumen ratio depended on laying time and location, with the largest eggs laid during the peak period by birds breeding in central positions on large colonies. Eggs produced by these peak layers also had the largest yolk-to-albumen ratios. Harvested sites were characterised by reductions in egg volume, yolk-to-albumen ratio and eggshell thickness, which translated to poorer hatching success and chick survival. Harvested sites also had a higher proportion of abnormal eggs, particularly taking the forms of small yolkless eggs and unpigmented eggs. The reduced breeding success on harvested colonies is likely to be linked to depletion of the female's endogenous reserves which can also reduce future survival and breeding propensity.     

Experimental evidence that egg color indicates female condition at laying in a songbird

The signaling hypothesis of eggshell coloration in birds isbased on the assumption that females of species with blue-greeneggs signal their phenotypic quality to their mates throughdeposition of the antioxidant biliverdin as pigment. Egg pigmentationmay be an expression of the condition of females at laying orof genetic linkages between egg color and female performancevariables. We have supplemented 16 pied flycatcher, Ficedulahypoleuca, females with mealworms before and during laying andcompared the mass and color of their eggs as measured on theday of laying to those of 16 control females with the same nestconstruction and laying dates and clutch sizes. Supplementedfemales laid significantly heavier and more intensely blue-greeneggs than control females. Egg blue-green chroma was significantlyassociated with the amount of biliverdin in eggshells. Egg color,and thus biliverdin content, is an expression of female conditionat laying.     

Rapid long‐distance migration in Norwegian Lesser Black‐backed Gulls Larus fuscus fuscus along their eastern flyway

We studied the long‐distance migration of Lesser Black‐backed Gulls Larus fuscus fuscus breeding in northern Norway along their eastern flyway using geolocators in 2009 and 2010. The majority of birds wintered in lakes in East Africa and the southeast Mediterranean was the most important stopover area. Larus f. fuscus along the eastern flyway travelled at a net travel speed of 399 and 177 km/day during the autumn and spring migration, respectively, higher than published travel speeds for Dutch Larus fuscus migrating along the western flyway. The results suggest that the long‐distance migratory Norwegian L. f. fuscus seek to minimize time spent in transit, whereas lower travel speed during northerly spring migration may reflect differences in wind patterns or food conditions between spring and autumn.     

Genetic differentiation and phylogeography of gulls in the Larus cachinnans–fuscus group (Aves: Charadriiformes)

We studied mitochondrial genetic differentiation among nine taxa of large gulls of the Larus cachinnans-fuscus group, which form part of the circumpolar Herring Gull complex. Our primary interest was to see if there were unrecognized gene flow barriers, to what extent mitochondrial genetic population structure conformed to current taxonomic boundaries, and what it might reveal about possible differences in population history. Sequences (430 nucleotides) of the hypervariable control region I (HVR-I) were obtained from 580 individuals and proved highly informative within this recently diverged group of birds. Contrary to current classification, a basal split was revealed between an Atlantic-Mediterranean clade (atlantis, michahellis, armenicus) and a NW Palearctic-Central Asian clade (cachinnans, barabensis, mongolicus, fuscus-group). There was almost no mitochondrial gene flow between these two groups, although they are in geographical contact in two areas (eastern North Atlantic, Black Sea). Within each of the two major groups, there was strong phylogeographic structure with gene flow barriers between some neighbouring taxa (e.g. cachinanns vs. barabensis), but also a case of poor genetic differentiation between phenotypically distinct forms (barabensis vs. heuglini). At the subspecies level, current taxonomy corresponded well to molecular genetic structure: over 80% of the molecular genetic variance was partitioned among six (groups of) taxa. This is in sharp contrast to previous studies using allozymes and amplified fragment length polymorphism (AFLP) markers, which seemed to indicate extensive nuclear gene flow. Within-taxon haplotype phylogenies and mismatch distributions revealed contrasting demographic histories: cachinnans (Ponto-Caspian region) and atlantis (NE Atlantic) represent ancient lineages with large long-term population sizes, inland forms stem from very recent colonization events (barabensis, mongolicus) or passed through a population bottleneck (armenicus).     

Validation of water flux and body composition in glaucous gulls (Larus hyperboreus)

Water influx rates (WIR) measured with tritiated water dilution were compared with direct measures of water and energy intake in glaucous gulls (Larus hyperboreus). Total body water (TBW) measured isotopically was also compared with TBW determined by body composition analysis (BCA) of the same birds. Seventeen wild gulls were captured and studied in outdoor enclosures at Ny-Alesund, Svalbard, in July 2002. Gulls were hand-fed known quantities of Arctic cod (Boreogadus saida) or given water on the basis of one of four experimental treatments: (A) fasting, (B) fish only, (C) water only, or (D) fish and water. Water and energy content of Arctic cod was also determined. WIR of gulls (after subtracting metabolic water production) in treatments A, B, C, and D were 0, 101 +/- 5, 62 +/- 19, and 122 +/- 21 SD g d(-1), respectively. Measured water intake in each group was 0, 111 +/- 2, 64 +/- 3, and 134 +/- 15 SD g d(-1), respectively. On average, WIR underestimated measured water intake in each group. Errors were lowest but most variable for gulls fed water only (-2.2% +/- 32.8%) compared with gulls fed fish only (-9.0% +/- 5.4%) or fish and water (-9.0% +/- 7.0%). Compared with measured water intake, errors in WIR were relatively low overall (-6.9% +/- 17.4%) and comparable to previous validation studies. The difference in TBW determined by BCA versus isotopic dilution ranged between -1.02% and +8.59% of mass. On average, TBW measured isotopically (632 +/- 24 g kg(-1)) overestimated true body water by a factor of 1.033.