Breeding phenology and winter activity predict subsequent breeding success in a trans-global migratory seabird.

Inter-seasonal events are believed to connect and affect reproductive performance (RP) in animals. However, much remains unknown about such carry-over effects (COEs), in particular how behaviour patterns during highly mobile life-history stages, such as migration, affect RP. To address this question, we measured at-sea behaviour in a long-lived migratory seabird, the Manx shearwater (Puffinus puffinus) and obtained data for individual migration cycles over 5 years, by tracking with geolocator/immersion loggers, along with 6 years of RP data. We found that individual breeding and non-breeding phenology correlated with subsequent RP, with birds hyperactive during winter more likely to fail to reproduce. Furthermore, parental investment during one year influenced breeding success during the next, a COE reflecting the trade-off between current and future RP. Our results suggest that different life-history stages interact to influence RP in the next breeding season, so that behaviour patterns during winter may be important determinants of variation in subsequent fitness among individuals.

The ecology and age structure of a highly pathogenic avian influenza virus outbreak in wild mute swans.

The first UK epizootic of highly pathogenic (HP) H5N1 influenza in wild birds occurred in 2008, in a population of mute swans that had been the subject of ornithological study for decades. Here we use an innovative combination of ornithological, phylogenetic and immunological approaches to investigate the ecology and age structure of HP H5N1 in nature. We screened samples from swans and waterbirds using PCR and sequenced HP H5N1-positive samples. The outbreak's origin was investigated by linking bird count data with a molecular clock analysis of sampled virus sequences. We used ringing records to reconstruct the age-structure of outbreak mortality, and we estimated the age distribution of prior exposure to avian influenza. Outbreak mortality was low and all HP H5N1-positive mute swans in the affected population were <3 years old. Only the youngest age classes contained an appreciable number of individuals with no detectable antibody responses to viral nucleoprotein. Phylogenetic analysis indicated that the outbreak strain circulated locally for ~1 month before detection and arrived when the immigration rate of migrant waterbirds was highest. Our data are consistent with the hypothesis that HP H5N1 epizootics in wild swans exhibit limited mortality due to immune protection arising from previous exposure. Our study population may represent a valuable resource for investigating the natural ecology and epidemiology of avian influenza.

Migration and stopover in a small pelagic seabird, the Manx shearwater Puffinus puffinus: insights from machine learning.

The migratory movements of seabirds (especially smaller species) remain poorly understood, despite their role as harvesters of marine ecosystems on a global scale and their potential as indicators of ocean health. Here we report a successful attempt, using miniature archival light loggers (geolocators), to elucidate the migratory behaviour of the Manx shearwater Puffinus puffinus, a small (400 g) Northern Hemisphere breeding procellariform that undertakes a trans-equatorial, trans-Atlantic migration. We provide details of over-wintering areas, of previously unobserved marine stopover behaviour, and the long-distance movements of females during their pre-laying exodus. Using salt-water immersion data from a subset of loggers, we introduce a method of behaviour classification based on Bayesian machine learning techniques. We used both supervised and unsupervised machine learning to classify each bird's daily activity based on simple properties of the immersion data. We show that robust activity states emerge, characteristic of summer feeding, winter feeding and active migration. These can be used to classify probable behaviour throughout the annual cycle, highlighting the likely functional significance of stopovers as refuelling stages.

The involvement of prolactin in avian molt: the effects of gender and breeding success on the timing of molt in Mute swans (Cygnus olor).

The aim of the study was to test the hypothesis that decreasing plasma prolactin stimulates or permits the initiation of avian molt. Changes in the concentration of plasma prolactin in Mute swans (Cygnus olor) were compared in non-breeding singletons and breeding pairs. In breeding swans, the onset of molt is delayed compared to non-breeders, and is delayed further in breeding males compared to their female partners. The seasonal decrease in prolactin in non-breeding birds of both sexes started at the end of May and was associated with the initiation of molt 4 weeks later. The decrease in plasma prolactin in incubating females was more pronounced, as a consequence of increased prolactin secretion associated with incubation behavior, but also started at end of May, and was associated the onset of molt 6 weeks later. In breeding males, plasma prolactin increased at the end of May when they started to care for their newly hatched cygnets. Correspondingly, prolactin began to decrease 3-5 weeks later in males than in females. These males started to molt in mid August, at least 4 weeks later than females. It is concluded that molt is related to decreasing plasma prolactin, and is inhibited when plasma prolactin is increasing or high.

Age-specific reproduction in a long-lived species: the combined effects of senescence and individual quality.

Apparent changes in breeding performance with age measured at the population level can be due to changes in individual capacity at different ages, or to the differential survival of individuals with different capabilities. Estimating the relative importance of the two is important for understanding ageing patterns in natural populations, but there are few studies of such populations in which these effects have been disentangled. We analysed laying date and clutch size as measures of individual performance in a population of mute swans (Cygnus olor) studied over 25 years at Abbotsbury, UK. On both measures of breeding performance, individuals tended to improve up to the age of 6 or 7, and to decline after about the age of 12. Individuals with longer lifespans performed better at all ages (earlier laying, larger clutches) than animals that ceased breeding earlier. We conclude that the apparent mean increase in performance with age in mute swans is due to both individual improvement and differential survival of individuals who perform well, while the decline in older age groups is due to individual loss of function. Our results underline the need to take individual differences into account when testing hypotheses about life histories in wild populations.

Components of variance underlying fitness in a natural population of the great tit Parus major.

Traits that are closely associated with fitness tend to have lower heritabilities (h2) than those that are not. This has been interpreted as evidence that natural selection tends to deplete genetic variation more rapidly for traits more closely associated with fitness (a corollary of Fisher's fundamental theorem), but Price and Schluter (1991) suggested the pattern might be due to higher residual variance in traits more closely related to fitness. The relationship between 10 different traits for females, seven traits for males, and overall fitness (lifetime recruitment) was quantified for great tits (Parus major) studied in their natural environment of Wytham Wood, England, using data collected over 39 years. Heritabilities and the coefficients of additive genetic and residual variance (CVA and CVR, respectively) were estimated using an "animal model." For both males and females, a trait's correlation (r) with fitness was negatively related to its h2 but positively related to its CVR. The CVA was not related to the trait's correlation with fitness in either sex. This is the third study using directly measured fitness in a wild population to show the important role of residual variation in determining the pattern of lower heritabilities for traits more closely related to fitness.

Is clutch size in birds affected by environmental conditions during growth?

Only environmental conditions occurring at the time of breeding have been shown to affect clutch size in birds, even though conditions experienced during growth are known to affect body size or egg size. We show here that environmental conditions experienced during early life can affect clutch size in captive zebra finches (Taeniopygia guttata) and wild great tits (Parus major). Not only do factors outside the immediate breeding season affect clutch size, but clutch size control mechanism is permanently influenced by conditions experienced during ontogeny.

Poxvirus infection of the Manx shearwater (Puffinus puffinus).

During studies on the etiology of puffinosis, a disease of the Manx shearwater, 1 to 4% of full-grown birds were found to have dry, non-pigmented lesions on the webs of the feet. Poxvirus infection was detected in six of seven full-grown birds with such lesions. The lesions contained large encapsulated inclusions which were packed with mature and immature poxvirus particles. Poxvirus infection was not apparent in shearwater fledglings during puffinosis epizootics, and its spatial distribution was not related to that of puffinosis. The results indicate that poxvirus infection produces a mild, self-limiting disease in shearwaters and is not the cause of puffinosis.

Hematology of fledgling Manx shearwaters (Puffinus puffinus) with and without 'puffinosis'.

Hematological parameters were measured in 14 fledgling Manx shearwaters (Puffinus puffinus), with the disease puffinosis and in 10 birds that did not have the disease, on the Island of Skomer between 2 and 11 September 1991. The mean plasma fibrinogen concentration was significantly higher in the diseased birds and some of these had abnormally elevated monocyte counts. No other significant differences were observed.

Bird biodiversity.

Endemic Bird Areas of the World. Priorities for Biodiversity Conservation by A.J. Stattersfield, M.J. Crosby, A.J. Long and D.C. Wege, Birdlife International, 1997. £37.00 pbk (860 pages) ISBN 0946 888 33 7 CDROM: Birds of the World-A Multimedia Encyclopedia, Ransom, 1997. £14.99 ISBN 1 863 89 1748.

A survey of blood lead levels in Mute Swans Cygnus olor.

Following bans on the use of most lead angling weights, the incidence of lead poisoning cases in Mute Swans started to fall and the population started to increase. However, surveys of lead levels in blood of rescued swans continue to show that a high proportion of the birds are carrying levels in excess of 1.21 mol/l. Since rescued swans, although rescued for many different reasons, might be a biased sample, a survey was made of apparently healthy birds living in flocks in the summer. These too showed that a high percentage of the birds in most of the flocks sampled had blood lead levels in excess of 1.21 mol/l. No source of this lead has been identified other than lead fishing weights; these may be long-lost leads, current, but illegally used weights or "dust-shot" which it is still legal to use. Although many of the birds sampled are carrying lead burdens that are probably not harmful, others are seriously affected. Except in the most serious cases, it is not possible to recognise birds with elevated lead levels without taking a blood sample for assay.