Group and family living is an integral part of many animals’ ecology. Thus the behavior became associated with plenty of advantages as well as disadvantages. However, rarely has the actual concept of the group been investigated. Questions such as, “What constitutes a ‘group’?” and “Do the individuals within these ‘groups’ associate with each other frequently enough to actually enjoy the benefits of group living?” are seldom asked. With these questions in mind, the aim of this study was to use Siberian jay (Perisoreus infaustus) individuals in their territories to explore and shed more light on the issue of the extent of group living. A working definition of a ‘group’ was made and subsequently the birds were observed in their natural habitat in northern Sweden during several seasons. Consequently, a pair-wise coherence index (CI) was created to quantify the levels of association between the individuals within each territory in order to investigate which possible factors affect the extent of the observed cohesion. Results of the study indicate that pair-wise cohesion was strongly dependent on the kinship of the birds. Moreover, it was found that the alpha and kin birds had significantly higher coherence values than the non kin birds. Thus, within a territory, it was the alpha and the kin birds that formed the core of the group with the non kin birds being much less associated with the other individuals. This was in contrast to the hypothesis, which predicted alpha birds to form the core of the group solely, with kin birds having significantly lower coherence. Interestingly, no effect of habitat type and season was found on the cohesion of the territorial group. Moreover, further evidence of sub-grouping was found even on the kinship level, thus suggesting yet more complex interplay between group cohesion and the group members.  All in all, with the non kin birds having low coherence values, the study casts light on the fact that even if individuals are present in a territory, they might not necessarily be a tight unit and as such be unable to fully benefit from group living.

<p>Genetic variation within and among populations is a result of past and ongoing processes. Among the most important of such processes are dispersal, habitat fragmentation and selection. This thesis use neutral genetic variation as a tool to investigate these processes in three bird species.</p><p>In the Siberian jay, the timing of dispersal is dependent on social dominance among siblings. Mark-recapture data, radio-tracking and genetic variation was used to investigate whether timing of dispersal had an effect on dispersal distance. The results show that early dispersing individuals also disperse longer. In the same species, genetic correlation between neighbours was used to find areas with high production of philopatric individuals, which could be indicative of high habitat quality.</p><p>Great snipe populations in northern Europe have a breeding range divided into two regions. A Q_ST-F_STapproach was applied to study variation in selection between regions. Differentiation between the regions in neutral molecular markers was low, indicating high gene flow, or short time available for neutral divergence. Morphological divergence between the regions was high, and Q_ST > F_ST, which indicates divergent selection. Thus, neutral genetic markers can be misleading in identifying evolutionary significant units, and the Q_ST-F_ST approach might be valuable to identify targets for conservation.</p><p>Rock ptarmigan, or its ancestors, originated in Beringia, and spread throughout the Holarctic region. Their distribution has subsequently been affected by glaciations, most likely leading to withdrawals and re-colonisations. Neutral genetic variation among five populations around the northern Atlantic was investigated. There was strong genetic structure among the populations, and evidence that Scandinavian rock ptarmigan has been isolated from other populations for considerable time. Rock ptarmigan in Svalbard showed slightly lower genetic variation than others, and comparisons with other studies suggested an eastern colonisation route to Svalbard.</p>

Genetic variation within and among populations is a result of past and ongoing processes. Among the most important of such processes are dispersal, habitat fragmentation and selection. This thesis use neutral genetic variation as a tool to investigate these processes in three bird species. In the Siberian jay, the timing of dispersal is dependent on social dominance among siblings. Mark-recapture data, radio-tracking and genetic variation was used to investigate whether timing of dispersal had an effect on dispersal distance. The results show that early dispersing individuals also disperse longer. In the same species, genetic correlation between neighbours was used to find areas with high production of philopatric individuals, which could be indicative of high habitat quality. Great snipe populations in northern Europe have a breeding range divided into two regions. A QST-FST approach was applied to study variation in selection between regions. Differentiation between the regions in neutral molecular markers was low, indicating high gene flow, or short time available for neutral divergence. Morphological divergence between the regions was high, and QST &gt; FST, which indicates divergent selection. Thus, neutral genetic markers can be misleading in identifying evolutionary significant units, and the QST-FST approach might be valuable to identify targets for conservation. Rock ptarmigan, or its ancestors, originated in Beringia, and spread throughout the Holarctic region. Their distribution has subsequently been affected by glaciations, most likely leading to withdrawals and re-colonisations. Neutral genetic variation among five populations around the northern Atlantic was investigated. There was strong genetic structure among the populations, and evidence that Scandinavian rock ptarmigan has been isolated from other populations for considerable time. Rock ptarmigan in Svalbard showed slightly lower genetic variation than others, and comparisons with other studies suggested an eastern colonisation route to Svalbard.

The dynamics of natural populations may be influenced by a variety of factors, ranging from feeding interference and territoriality to the risk of predation and climatic effects. The relative influence of these factors may be contingent upon the quality of the habitat in which individuals reside. A factor that can largely affect population dynamics and that often covaries with habitat structure is predation risk. However, the combined effect of habitat and predation risk can vary according to the social context and intrinsic characteristics that affect individual behavioural responses. This thesis investigates the effects of habitat quality at the level of the population and the individual, and examines how it relates to the above factors in Siberian jays (Perisoreus infaustus), a territorial, group-living species in which the main cause of mortality is predation. The results demonstrate a strong effect of habitat on survival, reproduction and behaviour. Mortality was generally higher in open areas and managed forests and reproductive success decreased after forest management. Breeding females were more sensitive to environmental factors than males, possibly because of higher reproductive costs. Estimates of spatial demography suggested that there were more sinks than sources, and that they were located in open, managed forests. Behavioural decisions confirmed that open forests were associated with higher predation risks. However, decisions depended on social context; immigrants took highest risks and were the recipients of most aggression, largely an effect of social subordination. Also, parents provide their offspring with benefits that are withheld from immigrants. As a result, first-year survival was higher in retained offspring. Investigating the effects of multi-scale habitat quality on individual behaviour and population dynamics has generated an increased understanding of the effects of forest management on the dynamics of this population. This approach should facilitate development of an effective conservation management policy for this species.

<p>This thesis examines risk management in breeding Siberian jays (<i>Perisoreus infaustus</i>), which is indigenous to the northern taiga. Parent behaviour and the nest are cryptic. A new nest is built each year. It is placed on spruce or pine branches close to the trunk and well insulated with lichens, feathers and reindeer hair.</p><p>Nest failure rate was the main factor driving annual variations in jay numbers. The probability for nesting attempts to be successful ranged annually between 0.08 and 0.70. Nest predation was rampant and a main cause of nest failure. Nest predators were mainly other corvids (primarily the Eurasian jay <i>Garrulus glandarius</i>). Habitat quality was the main factor determining the risk of predation. The risk for nest failure due to predation was higher in thinned forests with an open structure and with a high abundance of man-associated corvid species (jays, crows, raven). </p><p>Siberian jay parents show several strategic adjustments in life-history and behaviour to the risk of nest predation. Parents traded reduced feeding rates for a lower predation risk and allocated feeding to low risk situations. Chick provisioning imposes a cost by drawing the attention of visually hunting predators to the location of nests, and parents adjusted their daily routines and avoided exposure by allocating provisioning to times of low activity among nest predators. These strategic adjustments of feeding efforts were estimated to reduce the exposure to nest predators by 26 percent. Also, parents adjusted their reproductive efforts to the perceived presence of predators in a playback experiment. Siberian jays reduced their reproductive investment by laying a smaller clutch size when high risk of nest predation reduced the value of current reproduction, as predicted from life-history theory.</p>

<p>Evolution is fuelled by independent reproduction events. Yet, the offspring of at least three percent of all bird species postpone dispersal and forego independent reproduction. The Siberian jay, <i>Perisoreus infaustus, </i>is such a species where some<i> </i>offspring are philopatric and remain in their natal territory for up to three years, forming family groups. The main finding of this thesis is that nepotistic anti-predator behaviour displayed by parents provided philopatric offspring benefits, which could be an incentive to stay and forego independent reproduction. Predation, (hawks - 80 % and owls - 15% of deaths observed) is the main cause of mortality. Parents increased their vigilance nepotistically; they were more vigilant against surprise predator attacks, and gave alarm calls when attacked when feeding together with offspring. However, the two parents differed in their behaviour. Mothers gave calls only when together with their offspring, while males also warned unrelated immigrants. Sitting predators were approached and mobbed more intensely by parents in the presence of philopatric offspring. The vocalisation of Siberian jays provides information about predation risk. Specific calls are given for hawks and owls, and calls also varied with hawk behaviour. The nepotistic anti-predator behaviour of parents is a benefit, which the offspring can gain only “at home”, and such behaviour appears to promote offspring to forego dispersal and independent reproduction. This was confirmed in an experimental manipulation; philopatric offspring dispersed when fathers were removed and replaced by a despotic, immigrant stepfather. From a life-history perspective, parents have an incentive to protect their reproductive investment. Nepotistic anti-predator behaviour create a safe haven in the natal territory for philopatric offspring and provides direct fitness benefits. Without such direct fitness benefits offspring may disperse and wait for a breeding opening elsewhere. </p>

This thesis examines risk management in breeding Siberian jays (Perisoreus infaustus), which is indigenous to the northern taiga. Parent behaviour and the nest are cryptic. A new nest is built each year. It is placed on spruce or pine branches close to the trunk and well insulated with lichens, feathers and reindeer hair. Nest failure rate was the main factor driving annual variations in jay numbers. The probability for nesting attempts to be successful ranged annually between 0.08 and 0.70. Nest predation was rampant and a main cause of nest failure. Nest predators were mainly other corvids (primarily the Eurasian jay Garrulus glandarius). Habitat quality was the main factor determining the risk of predation. The risk for nest failure due to predation was higher in thinned forests with an open structure and with a high abundance of man-associated corvid species (jays, crows, raven). Siberian jay parents show several strategic adjustments in life-history and behaviour to the risk of nest predation. Parents traded reduced feeding rates for a lower predation risk and allocated feeding to low risk situations. Chick provisioning imposes a cost by drawing the attention of visually hunting predators to the location of nests, and parents adjusted their daily routines and avoided exposure by allocating provisioning to times of low activity among nest predators. These strategic adjustments of feeding efforts were estimated to reduce the exposure to nest predators by 26 percent. Also, parents adjusted their reproductive efforts to the perceived presence of predators in a playback experiment. Siberian jays reduced their reproductive investment by laying a smaller clutch size when high risk of nest predation reduced the value of current reproduction, as predicted from life-history theory.

Evolution is fuelled by independent reproduction events. Yet, the offspring of at least three percent of all bird species postpone dispersal and forego independent reproduction. The Siberian jay, Perisoreus infaustus, is such a species where some offspring are philopatric and remain in their natal territory for up to three years, forming family groups. The main finding of this thesis is that nepotistic anti-predator behaviour displayed by parents provided philopatric offspring benefits, which could be an incentive to stay and forego independent reproduction. Predation, (hawks - 80 % and owls - 15% of deaths observed) is the main cause of mortality. Parents increased their vigilance nepotistically; they were more vigilant against surprise predator attacks, and gave alarm calls when attacked when feeding together with offspring. However, the two parents differed in their behaviour. Mothers gave calls only when together with their offspring, while males also warned unrelated immigrants. Sitting predators were approached and mobbed more intensely by parents in the presence of philopatric offspring. The vocalisation of Siberian jays provides information about predation risk. Specific calls are given for hawks and owls, and calls also varied with hawk behaviour. The nepotistic anti-predator behaviour of parents is a benefit, which the offspring can gain only “at home”, and such behaviour appears to promote offspring to forego dispersal and independent reproduction. This was confirmed in an experimental manipulation; philopatric offspring dispersed when fathers were removed and replaced by a despotic, immigrant stepfather. From a life-history perspective, parents have an incentive to protect their reproductive investment. Nepotistic anti-predator behaviour create a safe haven in the natal territory for philopatric offspring and provides direct fitness benefits. Without such direct fitness benefits offspring may disperse and wait for a breeding opening elsewhere.