Agonistic display and spacing in the black-headed gull, Larus ridibundus

Black-headed gulls nesting on islands in a marsh habitat had a higher breeding success than those nesting on the periphery of the colony. Island birds maintained close inter-nest distances whereas peripheral nests were spaced further apart as predation of eggs increased. These changes in spacing corresponded with changes in the response distances of agonistic displays. There was no change during incubation for island birds but peripheral birds displayed to intruders at much greater distances than during pair formation. Island birds were more likely to attack and display more frequently to intruders than peripheral birds. However, in one season the peripheral birds were more aggressive during pair formation because of an increase in intrusions by unpaired females.     

Multiple maternity in black-headed gull Larus ridibundus clutches as revealed by protein fingerprinting

Social monogamy with biparental care is the norm in gulls Laridae , but egg colour variation suggests that some nests may contain mixed clutches laid by more than one female. Here we use protein fingerprinting of egg albumen to assess the occurrence of mixed maternity clutches in three colonies of black-headed gulls. Among 160 analysed clutches with >1 egg, 34% contained eggs from more than one female, and 15% of the eggs in clutches >1 came from other females than the major female (laying most eggs in nest). Among clutches with 2–3 eggs 28% were mixed, and among clutches with 4 or more eggs 89% contained eggs from two or more females. There were significantly fewer eggs from the major female in mixed nests (mean=2.06±0.63 SD) than in non-mixed nests (mean=2.82±0.43 SD). In nests without evidence of female conflict, hatching success of minority eggs was similar to that of eggs from the major female (12.5 and 8.4%, respectively). In 21% of mixed maternity nests, one or more minority eggs was buried or punctured, and 25% of eggs from major females were also found evicted, suggesting conflict between females and rejection of eggs. Intra-specific nest parasitism seems the most likely cause of mixed clutches, but there are also other possible causes.     

Maternal androgens in black-headed gull (Larus ridibundus) eggs: consequences for chick development

We tested the hypothesis that mother birds counterbalance the negative effects of hatching asynchrony for later-hatched chicks by increasing the yolk androgen concentrations in consecutive eggs of their clutch. In doing so, they may adaptively tune each offspring's competitive ability and, thus, growth and survival. However, evidence in support of this hypothesis is contradictory. The yolk concentrations of maternal androgens in the eggs of black-headed gulls increase significantly with the laying order of the eggs in a clutch. We experimentally tested the functional consequences of this increase on chick development under natural conditions by injecting eggs with either an oil or androgen solution. We created experimental clutches in which androgen levels either stayed constant or increased with laying order while controlling for differences in egg quality by using only first-laid eggs. We then compared development, growth and survival between these broods. Androgen treatment enhanced embryonic development because androgen-treated eggs hatched half a day earlier than controls, while their size at hatching was similar to oil-treated controls. Androgen treatment did not increase chick survival, but it enhanced growth. Androgen-treated, third-hatched chicks had a higher body mass and longer legs than third-hatched chicks that hatched from oil-treated eggs. At the same time, growth of first chicks (which were all oil treated) was reduced by the presence of two androgen-treated siblings, suggesting that yolk androgens enhance the competitive ability of later-hatched chicks. Our results support the hypothesis that transfer of different amounts of androgens to the eggs of a clutch is a mechanism by which mothers maximize their reproductive output.     

Lymphoid distribution in the migratory gull Larus ridibundus

We studied the distribution of lymphocytes in the main lymphoid tissues (blood, spleen and thymus) of the gull Larus ridibundus, searching for variations that might depend on the migratory cycle. We also looked for sex- and age-associated differences in lymphoid redistribution. In L. ridibundus, lymphocytes are the most commonly observed leukocyte subpopulation in blood. Moreover, changes in the distribution of lymphocytes in the lymphoid tissue occur, depending on the migratory period. The proportion of these cells in spleen is greater in the post-migratory and pre-migratory periods compared to the non-migratory period. The percentages of circulating lymphocytes are high in the pre-migratory period, but depletion occurs in the post-migratory period. In contrast, the age or the sex of the animals did not confer any major differences on the lymphoid distribution.     

Head-bobbing and non-bobbing walking of black-headed gulls (Larus ridibundus)

Head-bobbing walking (HBW) and non-bobbing walking (NBW) of black-headed gulls were compared from kinematic and behavioral/environmental viewpoints. The birds walked with a longer stride length and lower stride frequency during the HBW than during the NBW. With respect to these two parameters, the HBW of black-headed gulls was similar to that of other head-bobbers, and the NBW was similar to that of other non-bobbers. The stride length and the amplitude of head bobbing were correlated. These results suggest that the head-bobbing and gait parameters are related. From a behavioral viewpoint, HBW was observed during seeking-type foraging by wading, and NBW was observed during waiting-type foraging on a flat substrate. The type of foraging behavior and/or substrate condition probably determines whether the birds walk with or without head bobbing.     

Craniogenesis in the laughing gull (Larus ridibundus L.). The reconstruction of the basic plan of the bird

To contribute to the knowledge of the cranium of older bird embryos, the chondrocranium and osteocranium of the Black-headed Gull (Larus ridibundus) are described. The chondrocranium of the Black-headed Gull is compared with the chondrocranium of other bird species with special consideration of functional and phylogenetic aspects. As a result a "grundplan" of the bird chondrocranium is reconstructed. Most of the chondrocranial autapomorphies of birds are connected with the enlargement of the eyes, the development of a beak and a prokinetic skull, and the reduction of the olfactory system.     

Structural adaptation of bird hemoglobins to high-altitude respiration and the primary sequences of black-headed gull (Larus ridibundus, Charadriiformes) alpha A- and beta/beta'-chains

Two hemoglobin components HbA (alpha A2 beta 2) and (alpha D2 beta 2) have been detected by analytical electrophoresis in the lysed erythrocytes of the adult Black-Headed Gull (Larus ridibundus). We report the complete primary structure of the alpha A- and beta-chains of the major hemoglobin component HbA. Following the chain separation and isolation of the tryptic peptides by RP-HPLC, the amino-acid sequence was established by automatic Edman degradation in spinning cup and gas-phase sequencers. The primary structures of alpha A- and beta-chains from the Black-Headed Gull HbA differ by 11 and by 6 amino-acid residues from the corresponding chains of Greylag Goose. These changes are randomly distributed over both alpha-helical and interhelical regions. The presence of beta/beta'-chains is indicated by the observation of Ile/Leu at position beta 78. An exchange at position beta 55 (D6)Leu-Asn which is known to be involved in the alpha 1 beta 1-interface with alpha 119(H2)Pro has been found. It is suggested that packing contacts in the alpha 1 beta 1-interface are important for high altitude respiration in birds.     

Efficacy of Nigella sativa in alleviating benzo[a]pyrene-induced immunotoxicity in broilers

The immune response of broiler chickens exposed to intra-tracheal (i.t.) administration of benzo[a]pyrene (BaP) with and without Nigella sativa (Ns) supplementation was investigated. A total of 120 day-old chicks were divided into four groups comprising 30 birds each, into a control, Ns, BaP, and BaP+Ns group. Immune responses to Newcastle disease (ND) were evaluated by haemagglutination inhibition (HI), phytohaemagglutinin (PHA) skin test and carbon clearance assay (CCA). In most instances, there was a significant increase (p<0.05) in the ND-HI antibody titers, PHA skin-swelling response and phagocytic activity in the BaP + Ns group compared to that of the BaP group. Likewise, organ weight and indices of the spleen, bursa of Fabricius and thymus of birds from the BaP + Ns group were also higher (p<0.05) than that of the BaP group from day 1 until day 21. It is concluded that exposure to BaP may exert adverse effects on the immune system of broilers which may increase their susceptibility to disease, and Ns supplementation significantly reduces these alterations.     

The relation among gonadal steroids, immunocompetence, body mass, and behavior in young black-headed gulls (Larus ridibundus)

Abstract We experimentally examined the effect of testosterone on the antibody response to a single immunization with sheep red blood cells in young black-headed gulls. This species performs a number of testosterone-mediated elaborate postural displays in social interactions and breeds in dense colonies in which there is a high likelihood of infectious diseases. In young chicks, only one-third were capable of responding to immunization. In the responding chicks, testosterone enhanced antibody titers. Even when antibody responsiveness was measured >1 mo after the termination of hormonal treatment, antibody responses were enhanced in birds treated with androgen but not estrogen. At 9 mo of age, all birds responded to immunization, but there was no effect of testosterone on antibody titers. In these juveniles, frequency of display behavior was negatively related to changes in body mass, which suggests that displaying is energetically costly. Despite decreased body mass, antibody titers were highest in birds that displayed more frequently. This suggests that displaying signals immunocompetence. The results are discussed in relation to the immunocompetence handicap hypothesis and to what is known of the influence of the bursa of Fabricius and social stress on antibody production.     

Long-term, low-dose benzo[a]pyrene-induced mutation in human lymphoblasts competent in xenobiotic metabolism

We have studied the mutagenic effects of benzo[a]pyrene (BP) administered in a long-term, low-dose fashion to metabolically competent human lymphoblastoid cells. A continuous dose as low as 0.02 microM for 20 days produced a significant increase in mutant fraction at the 6TG-resistance (HGPRT) locus. The slope of the mutant fraction over time in the 0.02 microM BP-treated culture was twice that observed in the untreated concurrent control; 0.02 microM therefore represents the doubling dose of BP for gene mutation in this cell line. For higher doses of 0.1, 0.5 or 1 microM BP, the rate (or efficiency) of induced mutation was considerably higher for the first 5 or 6 days of exposure than for the last 14-15. This did not appear to be due to a growth disadvantage against early-arising mutants. Comparison to previously published data in the same cell system (Crespi and Thilly, 1984) revealed that the long-term , low-dose protocol (0-1 microM for up to 20 days) was significantly more efficient at inducing mutations than a short-term, high-dose protocol (0-10 microM for 1 day).     

Skull deformity in a herring gull chick (Larus argentatus)

A skull deformity resulting in death of a herring gull chick (Larus argentatus) is described in detail. The bones of the skull and upper jaw were twisted, asymmetrical and of unusual size or absent. The lower jaw had an almost "spoon-like" external appearance and lacked normal articular surfaces.     

Hybridization of glaucous gull (Larus hyperboreus) and herring gull (Larus argentatus) in Iceland: mitochondrial and microsatellite data

Large white-headed gulls provide an interesting group of birds for studies of hybridization. The group is composed of 20 species of recent origin, often with weak reproductive barriers. Here we report the results from a study on the glaucous gull Larus hyperboreus, an Arctic species which has been breeding in Iceland for centuries, and the herring gull Larus argentatus which has a wide distribution in Europe but colonized Iceland in 1920s. Previous studies, based on morphological variation indicated hybridization between the two species in Iceland, have been questioned as it may just reflect variation within the species. Here we evaluate whether hybridization has occurred between the two species in Iceland by studying variation in microsatellites and mtDNA. The analysis is based on feathers taken from wings sampled in Iceland over a period of 40 years. The results are compared with samples obtained from East Greenland and published sequences of samples obtained throughout Europe. The genetic analysis reveals a distinctive grouping of the two species, although they present a shallow genealogy and an extensive sharing of the genetic variants between the two species. Several individuals show admixture for molecular markers, which may result from an incomplete lineage sorting although geographical patterns of both mtDNA haplotypes and microsatellites strongly indicate a recent hybridization in Iceland.     

The Effect of Habitat Structure and Density of Nests on Territory Size and Territorial Behaviour in the Black-headed Gull (Larus ridibundus L.)

We studied the effects of vegetation structure and nest density on territory size and aggression in the black-headed gull (Larus ridibundus). Vegetation structure influenced the types of aggressive behaviour, but not territory size. The proportion of overt aggression (attacks, fights) was highest in barren areas or those with short, sparse vegetation, and lowest in plots covered with dense, tall grass. We explain this by decreased visibility of conspecifics in habitats isolated by vegetation. In such areas, total aggression did not decrease, because the low proportion of attacks and fights was offset by an increase of more moderate (“upright”) displays. Only when nest density was high did the vegetation reduce the frequency of agonistic behaviour — but then it comprised almost exclusively attacks and fights. As expected, greater nest density was related to reduced territory size and an increased proportion of overt aggression. A high proportion of attacks and fights occurred in the plot with the lowest nest density, where at the same time there were also numerous first-year breeders. We conclude that, when analyzing the effects of habitat conditions on aggressive behaviour, it is important to consider the structure of behaviour, and not only the total frequency of all types of agonistic interactions.