Sleep in the herring gull (Larus argentatus)

Sleep postures and eye state of free-ranging herring gulls (Larus argentatus) were studied during the breeding season. Three mutually exclusive behaviours were observed, namely sleep, rest-sleep and rest postures. Arousal thresholds, eye blink rates and eye closure time were obtained during these behaviours. Significant relationships existed between eye blinking, eye closure, and a raised threshold of arousal when birds were in the sleep and rest-sleep postures. During a natural disturbance, birds in the sleep posture remained in this posture but did not blink their eyes: this is called pseudo sleep. Male gulls also exhibited a lower threshold of arousal while in the sleep posture compared with females. We conclude that rhythmic eye blinking is a good indication of sleep in herring gulls.     

Manipulating daytime sleep in herring gulls (Larus argentatus)

The regular provision of a sufficient quantity of food on the territory of pairs of breeding herring gulls led to an increase in the time individuals spent on the territory during the day, relative to controls. This ‘extra’ time was not used to increase overall sleeping time, as predicted. Instead, the experimental gulls reduced their overall sleep time due to a decline in front-sleep. Back-sleep remained relatively unaffected, with both groups spending similar amounts of time in this posture. The consequences of the two postural types of sleep were examined with respect to the cost of allowing intruders to remain on an owner's territory. Intruders arrived more often on control territories and stayed longer when only one pair member was present. Intruders were seen on control territories for more consecutive scans when the solitary control gull was in a sleep posture, particularly front-sleep. It is concluded that these results imply a difference in the consequences resulting from the two sleep postures and give additional support for the multiple function hypothesis of sleep.     

Ontogeny of nesting habitat recognition and preference in neonatal herring gull chicks, Larus argentatus pontoppidan

The development of nesting habitat recognition and preferences in herring gull chicks was studied as a function of habitat and social stimuli. The test procedure consisted of relocating chicks 20 m from the nest and observing their movements. Results indicate that herring gull chicks exhibit a preference for the nest site area during the first week post-hatch, and this preference wanes after the first week until fledging. Chicks of all ages showed a preference for vegetation similar to that of the nest site area. Nest site preferences are ecotypically controlled. Vegetation characteristics, landmarks and the presence of other chicks were found to be among the cues used to recognize the nest site.     

Effects of early incubation constancy on embryonic development: An experimental study in the herring gull Larus argentatus

1.

In many birds, parental nest attendance in early incubation is variable, with eggs incubated only intermittently. The effect of this on chick hatching success is unknown.     

Colonial breeding reduces nest predation in the common gull (Larus canus)

It is often suggested that colonial breeding reduces nest predation for birds with a high defence capacity, but experimental comparison of predation at solitary and colonial nests is seldom feasible within a single species. We here report on such a test in the common gull (Larus canus). The rate of predation on experimental eggs was significantly lower near colonies than near solitary gull nests, and the eggs survived longer at the edge of a colony than farther away. Communal mobbing of nest predators is the likely reason. In both of two years, almost all nests of solitary gulls were destroyed by predators, while most clutches survived in colonies. Nest predation hence selects strongly for colonial breeding in the present population of common gulls.     

Endocrine aspects of female-female pairing in the Western gull (Larus occidentalis wymani)

Investigations of the colonies of western gulls on Santa Barbara Island, California, have revealed a surplus of females and the occurrence of female-female pairs that produce clutches with as many as six eggs. Females are able to establish and defend breeding territories, behaviours generally thought to be under the control of androgens. There are very few significant differences in circulating levels of the luteinizing hormone and androgens among breeding males, breeding females in heterosexual pairs, and breeding females in homosexual pairs. In contrast, however, only females sampled in 1877, in both homo- and heterosexual pairs, have elevated plasma levels of oestrogens, in spring, coincident with the period in which they show courtship behaviours such as food begging and solicitation of copulation. Given a sex ratio skewed in favour of females, as is apparently the case with the colonies of this species on Santa Barbara Island, and the essentially equal plasma levels of androgens in males and females, it is not difficult to rationalize the formation of female-female pairs. Our findings do not support the hypothesis that female-female pairing involves hormonal masculinization of one member of the pair.     

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.     

Development of individual call recognition in young ring-billed gulls (Larus delawarensis): An effect of feeding

Individual recognition has been documented in young gulls, but less is known of the relevant developmental mechanisms. Since parents normally emit mew calls during feeding of the young, I tested the hypothesis that feeding with individually distinctive mew calls constitutes one such mechanism. experimentally trained birds provided support for the hypothesis. Initially high approach and vocalization levels were maintained to the reinforced call, while responses to the other call declined, as did responses to both calls in untrained controls. Comparison with other species suggests that this developmental pattern is adapted to the natal social environment. In gulls, food is a potentially important mechanism influencing call ‘meaning’ (Smith 1977) between parents and offspring and between mates during courtship feeding.     

An outbreak of philophthalmosis in Larus michahellis and Larus fuscus gulls in Iberian Peninsula

Trematodes of the genus Philophthalmus Loos, 1899 are the eye parasites of birds and mammals, which use freshwater snails as their first intermediate hosts. Here we examined the presence of philophthalmids in a total of 1515 gulls (589 Larus fuscus and 926 Larus michahellis) admitted between January 2010 and October 2016 for rehabilitation at Olhão (Portugal), by the use of combined morphological and molecular analysis. We recorded the first infected L. fuscus and L. michahellis in July and November 2015, respectively. The philophthalmids were located in the conjunctival sac or under the nictitating membrane. Gulls infected with Philophthalmus lucipetus Rudolphi, 1819 presented no clinical signs, while those infected with Philophthalmus lacrymosus Braun, 1902 presented serious eye damage in the same host species. The prevalence of P. lucipetus reached 3.6% in L. fuscus and 0.8% in L. michahellis; the prevalence of P. lacrymosus reached 0.3% and 0.0%, respectively. The outbreak of P. lucipetus likely started in a narrowly defined area, since the first six cases, found between July and October 2015, originated from a single municipality, and only later more cases started to be retrieved from other municipalities of Portugal. These findings represent the first records of both philophthalmids in the Iberian Peninsula, their first records in L. michahellis and the first record of P. lacrymosus in L. fuscus. Further follow-up of the outbreak and the identification of intermediate hosts are needed.     

On the responses of laughing gull chicks (Larus atricilla) to the calls of adults I. Recognition of the voices of the parents

Twelve matched pairs of parent-raised and hand-raised laughing gull chicks from the same clutches were tested, at 6 to 8 days post-hatching, with successive presentations of playback of a recording of calls of their parents and recording of calls of parents from a neighbouring nest, in a situation in which they could approach or withdraw from the sound.The parent-raised chicks oriented towards the sound, moved towards the sound and called when the calls of their own parents were played; they tended to withdraw from the sound and sit silently when the calls of the neighbouring parents were played.The hand-raised chicks fled from the sound and crouched in silence in response to both calls of their parents and the calls of neighbours. They were more vocal and active in periods without sound than were the parent-raised chicks.It is concluded that laughing gull chicks in nature learn to recognize the individual characteristics of the calls of their parents, and to react positively to the calls, through interaction with their parents prior to the age of 6 days post-hatching.     

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.     

A physiological marker for quantifying differential reproductive investment between the sexes in Yellow-legged gulls (Larus michahellis)

Asymmetry between males and females in the energy they invest initially in reproduction has resulted in the evolution of differing reproductive strategies (caring females vs. competitive males). However, parental care in many birds is shared by both sexes suggesting that male energy expenditure in agonistic behaviors and courtship feeding might compensate female costs of clutch production. Here, we tested the hypothesis that initial investment in reproduction by both sexes in the Yellow-legged Gull (Larus michahellis), a species with biparental care, is similar from a physiological perspective. In this income breeder, female and male reproductive investment during early breeding can be ultimately related to muscular activity (local foraging effort required for clutch production in females and courtship feeding and agonistic behaviors in the case of males). Thus, we evaluated sex-specific patterns of creatine kinase (CK, IU/L) levels in plasma, an indicator of physical effort associated with muscular activity dependent behaviors, through incubation as a reflection of the physiological response of both sexes to the reproductive investment they made up to clutch completion. Raw levels of CK were related to plasma levels of total proteins (TP, g/dL) to account for the differential physiological state of individuals when sampled (i.e. differential dehydratation). Female costs of clutch production were associated with post-laying levels of CK/TP. We grouped females according to their relative investment in clutch production: < 15.8%, 15.8 to 17.3% and > 17.3% of field metabolic rate; which showed increasing values of CK/TP (24.6, 53.1 and 66.0 IU/g, respectively). Moreover, we found similar CK/TP trends throughout incubation for both sexes (CK/TP = 50.2− [3.3 × days from laying]) suggesting similar physiological responses to reproductive effort and, therefore, analogous sex-specific initial investment. Thus, male investment in agonistic behaviors and courtship feeding apparently equaled female investment in clutch production. The use of CK measurements is revealed as a useful approach to investigating overall investment in reproduction for both sexes, providing further insights into our comprehension of reproductive strategies in seabirds.     

The blood vascular system in the head of the herring gull (Larus argentatus)

The structure and development of the blood vascular system in the head of the herring gull (Larus argentatus) have been studied using injection techniques and histological sections. Three different but interconnected divisions of the arterial system are recognized in the adult: the cerebral carotid artery system, the external ophthalmic artery system, and the external carotid artery system. Embryologically, the arterial system is characterized by changes in the relative development of these three divisions; the cerebral carotid system being the most prominent in the first half of the embryonic period. The venous system is divided into two parts, the rostral cephalic system and the caudal cephalic system, which drain separate regions of the head. The Rete ophthalmicum, which is an arteriovenous network associated with the external ophthalmic artery system, can be identified from the fifth day of incubation, and its development appears to be coupled with changes in the arterial supply to the eye. The possibility of a homology between the Rete ophthalmicum of birds and the Rete caroticum of mammals is briefly discussed.     

On the responses of laughing gull chicks (Larus atricilla) to the calls of adults II. Age changes and responses to different types of call

Three age-groups of laughing gull chicks were tested with playback of recordings of calls of their parents and calls of neighbouring adults. The chicks in each group showed response to the playback calls, and recognition of the voices of their parents. The chicks in the early group (1 to 3 days post-hatching) showed the least evidence of parent recognition: the chicks of the late group (12 to 28 days post-hatching) showed the lowest levels of response in general.Evidence of relationships between variation in the behaviour in the tests and variation in the test recordings was found from correlation analysis, the type of call initiating positive response, and analysis of temporal association. This evidence, and the results of tests which allowed responses to the parents' crooning and responses to the parents' ke-hah calls to be compared for chicks in the late group, indicate that whereas the filial approach behaviour of very young chicks is stimulated by crooning but suppressed by ke-hah and long-calls, this behaviour in older chicks requires the occurrence of ke-hah calls or long-calls for its elicitation. However the ke-hah calls and long-calls of adults other than the parents continue to have negative effects on the behaviour of older chicks.It is argued that this progression in discrimination of the voices of the parents and the changes in responsiveness to the different types of adult calls are consistent with functional requirements, both from the point of view of changes in a growing chick's social situation and from the point of view of the characteristics of the calls. The changes are also discussed from a developmental point of view.     

A declaration of intent in the kittiwake gull (Rissa tridactyla)

During the breeding seasons 1979–1982 observations were made of the vocal behaviour of the kittiwake gull (Rissa tridactyla). Attention was directed at the vocal behaviour emitted immediately preceding a bird's departure from the nest and partner. The data indicate that the kittiwake consistently makes only one type of vocalization (the Pre-departure Call) prior to departure and that the behaviour communicates intention to leave nest and partner. An analysis of the behavioural interactions reveals that the Pre-departure Call given by one bird elicits five different categories of response in the other. The data reveal that consequent partner behaviour either sanctioned or prevented absence. The functional significance of the results is discussed.     

Age differences in kleptoparasitism by laughing gulls (Larus atricilla) on adult and juvenile brown pelicans (Pelecanus occidentalis)

We recorded the choice of victims by a population of adult and first-winter laughing gulls (Larus atricilla) making attempts to steal food from adult and juvenile brown pelicans (Pelecanus occidentalis). Adult gulls made food theft attempts on juvenile pelicans with a disproportionately great frequency; first-winter gulls appeared to select pelican victims at random. Adult pelicans were more frequently successful in their foraging attempts than juvenile pelicans, suggesting that adults may be more reliable as potential food sources for gulls making theft attempts. However, juvenile pelicans attempted to evade kleptoparasitizing gulls less frequently than did adult pelicans, suggesting that the fish prey of juveniles may be more easily stolen. These patterns are discussed in relation to optimal victim choice by gulls, and to deferred maturity in both species.     

Effect of rapid modulation of circulating plasma testosterone concentration on begging,aggressive behavior and competition for food in black-headed gull (Larus ridibundus) chicks

Sibling competition mediated by begging behavior is extremely common in avian species and recent studies have highlighted the role of endogenous testosterone in regulating such phenomenon. However, current literature depicts an inconsistent pattern in altricial vs. semi-precocial species, with stimulating versus inhibitory effects of the hormone respectively. This is possibly due to a difference in the methodology of hormone treatment (short-term moderate dose versus a long-term stronger elevation, respectively) between the studies performed so far. In this study, we induced short-term moderate peaks in plasma testosterone levels, as applied in altricial bird species, and assessed the effects of our manipulation on begging, competitive and aggressive behavior in black-headed gull (Larus ridibundus) chicks, a semi-precocial species. Our results suggest that, unlike in altricial songbirds, temporary increase of plasma testosterone concentration suppresses begging and enhances aggressiveness towards intruders. However, it also increases aggression and the chances of getting priority while scrambling with nest mates to gain access to food. Thus, the inconsistencies in the hormonal control of begging behavior observed between altricial vs. semi-precocial birds seem real and perhaps related to species differences in complexity of the display and the nature of competition. These may be elucidated by future comparative studies.