<strong>I. Introduction</strong> The family Paridae contains five important genera &ndash; Parus, Aegithalos, Psaltiparus, Auriparus and Remiz. Each of these genera includes a group of vary similar species, but the different genera appear to be widely separated from each other. The objects of the present study were:- To ascertain the annual cycle of behaviour of the Great Tit (Parus major), and to attempt an analysis of some aspects of it.</li> <li>To compare the behaviour of the Great Tit with that of other tits, particularly with other species of Parus. To this end parallel observations were made on the other members of the family present in the study area, and the relevant literature was reviewed. Comparative study is of value in facilitating the interpretation of behaviour patterns, and in indicating relationships between species and genera. The study was carried out in Wytham Wood, Berkshire. <strong>II. Methods</strong> A field study was made on a wild population of tits. Birds were trapped and ringed so that individuals were recognisable in the field. <strong>III. Flocking</strong> Outside the breeding season Great Tits live in flocks which show a moderate decree of integration and which move irregularly over a limited area at rates of up to 200&nbsp;yds. per hour. The flocks show two types of movement:- (i) Drifting movements occurring during feeding. (ii) Integrated movements, during which the birds stop feeding and move for some distance. In the latter the flocks are integrated by: (a) The 'Twink' call, which brings other individuals into a ready-to-fly mood. (b) The situation "other-bird-flying-away" which releases flying. The tendency to integration of the flock is balanced by certain disruptive tendencies, the most important of which are due to aggressive behaviour. <strong>IV. Winter Fighting</strong> Fighting over food is common in winter flocks. The most important behaviour patterns are:- (i) The supplanting attack, in which one bird flies at another and dispossesses it of its food. (ii) The wings-raised, and (iii) the head-forward postures, used to threaten nearby individuals, (iv) Avoiding behaviour: tits with food avoid others which approach too closely. (v) Flying away with food particles too large to be eaten immediately. The study of such behaviour can be amplified by allowing, the tits to feed in front of a mirror: they then behave towards their image as they would towards a rival. The image may provoke attack, or flight, or intermediate responses. Supplanting attacks are most common in mid-winter, and occur sufficiently often for such behaviour to be of definite ecological importance. <strong>V. Movements</strong> Adult Great Tits are completely sedentary, and remain on or near their previous year's territory the whole year round. Juvenile Great Tits disperse soon after becoming independent of their parents: they may wander up to several miles from their birth-place. Many juveniles hare settled in a restricted range by the Autumn, though some continue to wander until the nesting; season. The winter range is approximately 10&ndash;20 acres. <strong>VI. Pair Formation</strong> The Great Tit flocks begin to break up in January, but the process is gradual, and in cold weather some flocks say be seen as late as April. Some birds are paired during the winter, but most pair formation takes place when the flocks break up. The mechanism is not yet fully understood, but it appears that the male makes supplanting attacks on other members of the flock, and that his further behaviour depends on that of the attacked individual. Once pair formation has occurred, there is a marked difference in the behaviour of the sexes, the males being much more 'alert' and lass retiring than the females. Certain courtship displays appear at this time of year. <strong>VII. Territory</strong> Soon after the pairs separate out from the flock, the male Great Tits start to use certain sites in preference to others for singing. Fighting against other males is more intense in the neighbourhood of these sites (preferred stations) than elsewhere. In this way an area round the preferred stations (the preferred area) comes to be defended from other Great Tits. The boundaries of this area become defined through skirmishes with other pairs. The preferred area is at first unstable and its boundaries vague: it changes only gradually into a territory in the classical sense. Territorial behaviour is thus much less rigid in the titmice than in most species which have previously been studied. The average size of Great Tit territories in the study area was 2.0 acres. Territories seem to differ considerably in size for the same species in different areas, and also for different Parus species in the same area. The biological value of territorial behaviour is difficult to assess. <strong>VIII. Song</strong> Each Parus species has a characteristic song, though in each species there is considerable variation. The song period is roughly similar in all the Parus species studied: song starts in December, becomes common and well marked in early spring, and dies away at about the time when the young leave the nest. Song nay also be heard from juvenile birds in late summer, and during the autumn period of sexual behaviour. The singing of a male attracts females and warns off other males. The stimulus situations in which it occurs are similar to those in other Passerine species. <strong>IX. Reproductive Fighting</strong> Reproductive fighting occurs in relation to the defence of a female, a territory and a rest-site. The most important behaviour patterns are: (i) The supplanting attack, (ii) The head-up posture, (iii) Upright flight, and (iv) Displacement pecking. Although the behaviour is complex, it can be understood on the assumption that each of the combatants is activated by two drives &ndash; a fighting drive and an escape drive. The fighting drive is expressed primarily in the supplanting attack:, and the escape drive in flying away. In a skirmish the two drives are in approximate equilibrium. The strengths, and relative strengths, of these two drives determine which of the several possible responses will be given to each stimulus situation. In the Great Tit the head-up posture is very elaborate, and is accentuated by a black ventral stripe. In the Blue Tit the posture is less elaborate, and there is no corresponding structure. Although the displays used in intraspecific reproductive fighting are very similar in the different Parus species, they do not appear in the interspecific disputes over nest-sites. <strong>X. Copulation and Courtship Feeding</strong> In all Parus species it is usually the male who initiates copulation, though the female sometimes does so. The precopulatory display consists of a rapid shivering of the wings, a crouching posture, and a high pitched 'zeedling' note. This is given by both sexes. Courtship feeding often follows copulation. It becomes very common during the egg-laying period, the female begging almost continuously all day. The male continues tc feed the female during incubation. The begging display of the female is similar to the precopulatory display, and to the begging of the young. Wing shivering seems to be a ritualized intention movement of flight. <strong>XI. Nest Site</strong> All Parus species build in holes. The different species have slightly different requirements for nest-sites, though there is a very large overlap. During the pre-nesting season the male inspects suitable sites, and his display causes the female to shiver her wings and inspect the hole.

Šio darbo tikslas buvo išanalizuoti literatūros šaltinius ir kitus prieinamus duomenis apie akumuliuotos taršos kiekius ir galimą jos genotoksinį poveikį paukščiams (individai, populiacijos), ir įvertinti atliktus tiriamųjų rūšių konkrečius citogenetinius tyrimus, aplinkos taršos ir citotoksinių efektų atskiruose individuose kontekste. Atlikus citogenetinius tyrimus, pateikti rezultatų analizės ir statistinio apdorojimo metodus. Mikrobranduolių analizės metodas sėkmingai taikytas skirtingų sistematinių taksonų gyvūnams- moliuskams, žuvims ir žinoma paukščiams. Taigi, šiame darbe tiriamų paukščių populiacijų tyrimams buvo taikytas mikrobranduolių analizės metodas. Didžiausias mikrobranduolių dažnis konstatuotas, P. major 10,5%, o mažiausias A. arundinaceus 0,55% individuose. Tai gali rodyti skirtingą rūšių jautrumą citotoksiniam ir genotoksiniam aplinkos poveikiui. Didžiausia tikimybė, kad tokius mikrobranduolių skaičiaus eritrocituose skirtumus tarp rūšių nulemia didžiąją dalimi skirtinga mityba, skirtinga maisto objektų akumuliuojama tarša. / The purpose of this diploma paper was to analyse the sources of literature and other available materials (reports of monitoring, data bases), valuating investigative birds species organisms' impurity and possible genotoxic effect of this. And finally to exercise methods of cytological micronuclei experiments and adduce results analysis and statistical processing methods. Cytogenetic experiments (micronuclei test) were made in distinct species in Paridae and Sylviidae families. The highest frequency of micronuclei was found in populations of Parus major – 10,5%, the lowest – in populations of A. arundinaceus 0,55% micronuclei. This can show different species sensitivity to cytogenetic and genetic environmental influence. The highest possibility that the distinctions of micronuclei frequency in erythrocytes between species are mostly determined by the difference of nourishment and the accumulative pollution of nutrition objects. It particularly depends on the quality conditions in their ecological niches.

Inter-specific and intra-specific variation in reactions towards novel and aposematic prey was found in several species of tits (Paridae). This Ph.D. thesis is focusing on various factors influencing reactions towards novel and aposematic prey in three European species of tits. We tested differences in exploration behaviour, neophobia, dietary conservatism, personality, age and experience as well as ability of avoidance learning and generalisation. We found no difference in exploration behaviour and in reaction towards novel prey in two different populations of great tits (Parus major). But the birds from the Finnish population were more neophobic than Czech birds, but they attacked aposematic firebug (Pyrrhocoris apterus) more often and faster than Czech birds. The difference can be explained by a different experience with local aposematic prey communities. Than we studied initial wariness in naive juveniles of great tits (P. major), coal tits (Periparus ater) and blue tits (Cyanistes caeruleus), and we tested how the initial wariness towards novel and aposematic prey can be deactivated by experience with palatable prey. Great tits and coal tits from experienced groups significantly decreased their neophobia towards both types of prey while blue tits did not change their strongly neophobic...

Animals exhibit a wide variety of social behaviours that are shaped by the external group social structure. Thus, understanding social behaviours and processes requires examining the individual social associations that form the basis of a group’s social network. The first objective of this thesis was to assess the consistency of social position within wild networks of black-capped chickadees (Poecile atricapillus) and to evaluate the effects of individual behavioural traits (exploratory personality and social dominance) on network position. Intra-annual social position was found to be repeatable and centrality increased with dominance rank, suggesting that dominant individuals occupy more central positions. The second objective of this thesis was to evaluate how network position and individual traits influence the transmission of social information through groups; the use of information acquired by other group members is expected to be an important benefit to group living. Social information regarding the location of novel foraging patches was observed to be transmitted through all eight chickadee groups. The rate of information transmission was found to be positively associated with dominance rank, but was not influenced by exploratory personality, indicating that dominant individuals may have greater access to social information than more subordinate individuals. The final aim of this thesis was to assess if social information transmission varied between urban and rural environments, as increased resource variability in more rural sites was expected to lead to higher reliance on social foraging cues. However, no effect of level of urbanisation was detected on the transmission of social information regarding novel food sources, which may indicate a habitat-independent strategy of social information use in chickadees. In general the results from this thesis indicate the importance of dominance status on individuals’ position within a social group, which can lead to differential exposure to social processes, such as social information transmission.

Abstract Anthropogenic climate change and habitat loss and deterioration affect populations worldwide. Climate warming has changed phenologies of many species across trophic levels. Some predator populations now experience temporal mismatches with their prey, as timings of peak prey abundance and of the predator’s highest food demands no longer meet. The temporal mismatch hypothesis suggests that the population’s recruitment rate is related to its degree of synchrony with the food resources needed to feed offspring. However, species’ and populations’ responses to climate warming differ. Human land use alters and destroys habitats of countless species. For example, many boreal forest bird populations have declined, presumably due to intensive forestry. It has decreased the amount of dead wood, causing a threat to saproxylic species. Identification of the key characteristics of high-quality habitats is essential for conservation planning and for developing sustainable forestry. As individuals are suspected to settle in habitats that maximize their fitness, analysis of nest site selection can be used to identify the key habitats. My dissertation concerns the impacts of climate change and habitat deterioration on boreal populations. I use hole-nesting passerines as model species. By utilizing long-term data I show that breeding phenologies of Parus major and Poecile montanus, but not of Cyanistes caeruleus, have shifted earlier. Also, the timing of the food peak has advanced, improving the synchrony between P. montanus and caterpillars. In P. major and C. caeruleus, synchrony has remained good. However, the positive effect of good synchrony on breeding success seems to be conditional, arising only in certain circumstances, such as in years of high caterpillar abundance. I suggest that in boreal populations temperature affects timing of breeding mostly as a proximate factor. The availability of standing decaying trees used for nesting sites was the most important habitat characteristic determining the nest site selection of P. montanus. Remote sensing data alone was insufficient to produce reliable models, as the ecologically important small-scale factor can only be determined by direct field surveys. Omission of forest thinning in selected forest sites would ensure the continuous availability of decaying wood with positive influence on biodiversity in managed forests. / Tiivistelmä Ihmisen aiheuttama habitaattien katoaminen ja huononeminen sekä ilmastonmuutos vaikuttavat populaatioihin kaikkialla maailmassa. Ilmaston lämpeneminen on muuttanut monien lajien fenologioita eri trofiatasoilla. Osalla saalistajalajien populaatioista suurin ravinnontarve ei enää ajoitu samaan aikaan saaliin runsaushuipun kanssa. Ajoituksen eriaikaisuus -hypoteesin mukaan todennäköisyys populaatioon rekrytoitumiselle riippuu synkronian asteesta saaliin kanssa. Ilmaston lämpenemisen vaikutuksissa lajeihin ja populaatioihin on kuitenkin vaihtelua. Ihmisen maankäyttö muuttaa ja tuhoaa lukuisien lajien elinympäristöjä. Esimerkiksi useiden boreaalisten metsien lintupopulaatioiden pienentymistä on selitetty intensiivisellä metsätaloudella. Lahopuun määrä metsissä on vähentynyt, mikä on uhka lahopuusta riippuvaisille lajeille. Korkealaatuisten habitaattien keskeisten piirteiden tunnistaminen on tärkeää luonnonsuojelun ja kestävän metsätalouden suunnittelulle. Koska yksilöiden oletetaan valitsevan niiden kelpoisuutta maksimoivia elinympäristöjä, pesäpaikanvalinta-analyysiä voidaan käyttää tärkeiden habitaattipiirteiden tunnistamiseen. Tarkastelen väitöskirjassani ilmastonmuutoksen ja habitaattien laadun vaikutuksia boreaalisiin populaatioihin. Mallilajeina käytän koloissa pesiviä varpuslintuja. Hyödyntämällä pitkäaikaisaineistoja osoitan, että lisääntymisen ajoittuminen on aikaistunut tali- ja hömötiaisella, mutta ei sinitiaisella. Myös ravintohuippu on aikaistunut, mikä on parantanut synkroniaa hömötiaisen ja sen pääasiallisen ravinnon eli toukkien välillä. Tali- ja sinitiaisella synkronia on pysynyt hyvänä. Hyvän synkronian myönteinen vaikutus lisääntymismenestykseen vaikuttaa kuitenkin ehdolliselta: se tulee esiin vain tietyissä olosuhteissa, kuten vuosina jolloin toukkia on runsaasti. Kevään lämpötilat näyttävät vaikuttavan pesinnän ajoittumiseen erityisesti proksimaattisena tekijänä. Pesäpaikkoina toimivien seisovien lahopuiden määrä on tärkein hömötiaisen pesäpaikanvalintaa määräävä tekijä. Kaukokartoitusaineisto yksinään ei riitä luotettavien mallien tuottamiseen, sillä ekologisesti tärkeät pienen skaalan tekijät voidaan kartoittaa vain suorin maastomittauksin. Metsien harventamatta jättäminen valituilla laikuilla turvaisi lahopuun jatkuvan saatavuuden, mikä vaikuttaisi myönteisesti talousmetsien biodiversiteettiin.

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Social behaviour is shaped by complex relationships between evolutionary and ecological processes interacting at different scales. Benefits gained from social associations can range from predator dilution to collective sensing, but little is known about how these can be influenced by social structure and phenotypic composition. In this thesis, I investigated how individual decision-making affects phenotypic social structure, and how this mediates social behaviour through emergent properties of collective group behaviour. First, using mixed-species flocks as a model system, I showed individual tits (Paridae, chapter 2) and thornbills (Acanthizae, chapter 3) varied significantly in their social positions. Within-species variation in network position was as large as between-species variation, sug- gesting that prescribing functional roles at the species level may not sufficiently account for potential differences in fitness operating at the individual level. Rather, this suggested that structure may be driven by phenotypic traits, underpinning network structure (chapter 4). Next, I used an extensive data set of foraging records to explore factors determining the composition, of flocks of great tits (Parus major, chapter 5). For example, assortment by dispersal phenotype (immigration status) was the result of spatial disaggregation, and I showed that this may facilitate social selection for breeding territories (chapter 6). Finally, I investigated how decision-making shaped mixed-species social structure. I found that tits used a common strategy for managing pressures of predation and starvation by shifting from exploration to exploitation at different times of the day (chapter 7). I then found that a very simple interaction rule successfully replicated mixed-species group structure (chapter 8). Strikingly, the same rule was applied to both conspecifics and het- erospecifics, potentially playing an important role in the maintenance of flock structure. Through experimental manipulation of ecological conditions, I found that heightened per- ceived predation resulted in stronger social attraction overall, whereas increased competition led to a reduction in attraction to conspecifics (chapter 9). Simulations suggested this could be one potential mechanism underpinning fission-fusion dynamics in these species. Together, the results in this thesis form a framework linking social behaviour to individ- ual fitness where natural selection is shaped by the social environment. This approach may prove useful for testing whether following common social rules reduces variance in benefits accrued by individuals, and how within-species variation in social behaviour can impact emergent properties of groups.

Neophobia and exploratory behaviour are two processes applied in great measure in birds' reactions towards novel stimuli. The specific demonstration of these two processes may differ between different species of birds as well as within them. We compared neophobia and exploratory behaviour in two species of tits (Paridae). Adult Great tits (Parus major) and Blue tits (Cyanistes caeruleus) did not differ neither in exploring novel environment and novel object nor in neophobia towards novel object near food. We have found great differences between juveniles of these two species. Juvenile Blue tits were more neophobic towards both novel objects. Age, sex and personality affected some differences within tested species. We tested whether there exist any differences in reactions towards novel food between and within species. We used red painted mealworm (larvae of Tenebrio molitor), house cricket (Acheta domestica) and aposematic bug (Pyrrhocoris apterus) as novel food. Adult wild-caught tits with plentiful experience with various feeds did not differ in reaction towards these types of novel food. In contrast hand-reared juvenile tits differed quite a lot. Juvenile Blue tits were more neophobic than juvenile Great tits towards all three types of novel food. We found dieraty conservatism, phenomenon...