Aspects of the ecology of black grouse (Tetrao Tetrix), a species of international conservation concern, in commercial plantation forests were investigated between 1996 - 1998 at three study areas in Scotland. The aim was to identify the species' habitat and area requirements in first and second rotation forestry. The distribution of males was assessed using lek surveys and studied at two spatial scales in the mixed rotation forest landscape of Cowal, Argyll and at two spatial and temporal scales in highland Perthshire - a less afforested region. In addition, a radio-tracking study was undertaken to examine the habitat selection of broods in two first rotation plantations in north Perthshire. Pre-thicket forestry formed a preferred habitat but, in terms of lek distribution, black grouse did not differentiate between first and second rotation pre-thicket habitat patches. Patch size, the total amount of forestry in the area, the proportion that was pre-thicket stock and its level of fragmentation, however, were all correlated with the probability of a location holding a lek and the number of males attending it. Lek isolation reduced the number of males in attendance and increased the likelihood of the lek declining over time. Brood habitat preferences differed from those of adult birds. Broods selected habitats that were 'open' enough to support a rich ground flora and presumably an adequate invertebrate fauna but also 'closed' enough to provide cover, representing a compromise between foraging potential and predation risk. Brood roost sites differed by having shorter trees but a higher degree of cover 1-1.5 metres above the ground. Results from the different study areas and age classes are compared and contrasted and the implications of the research findings for 'black grouse friendly' forestry management are discussed. Finally, suggestions for further work are made.

The aim of this report was to investigate how much people in general know about the bird black grouse and the possibilities for guiding on black grouse lek. I wanted to know how the possibilities where generally and specific at the tourist-facility Granö beckasin, which is situated in municipality of Vindeln in north of Sweden. I also wanted to get a better understanding about how the black grouse lek works and how the guiding would be best performed. Two different questionnaires were laid out, one with aim to reach a wide public and discover the knowledge and interest among people in general about black grouse lek. The aim with the second questionnaire was to reach people who is interested in bird- watching and get more quality answers about guiding on black grouse lek. While the questionnaires were collecting answers, market research about guiding and investigation of black grouse lekking spots possible for guiding around Granö was done. Most of the participants in the questionnaires showed a high level of awareness when it came to know what black grouse lek is, the majority also describe an active outdoorlife. The study showed a high level of black grouse lekking spots around Granö. Research and questionnaires showed that the target group for this type of guiding is small. The people in target for the type of guiding have big interest for birds and nature but doesn´t always have the time to research black grouse lekking spots themselves, therefore could a guide be helpful.

From an ecological perspective, knowledge of demographic history is highly valuable because population size fluctuations can be matched to known climatic events, thereby revealing great insight into a species’ reaction to past climate change. This in turn enables us to predict how they might respond to future climate scenarios. Prominently, with the advent of high-throughput sequencing it is now becoming possible to assemble genomes of non-model organisms thereby providing unprecedented resolution to the study of demographic history and speciation. This thesis utilises four species of grouse (Aves, subfamily Tetraoninae) in order to explore the demographic history and speciation within this lineage; the willow grouse, red grouse, rock ptarmigan and the black grouse. I, and my co-authors, begin by reviewing the plethora of methods used to estimate contemporary effective population size (Ne) and demographic history that are available to animal conservation practitioners. We find that their underlying assumptions and necessary input data can bias in their application, and thus we provide a summary of their applicability. I then use the whole genomes of the black grouse, willow grouse and rock ptarmigan to infer their population dynamics within the last million years. I find three dominant periods that shape their demographic history: early Pleistocene cooling (3-0.9 Mya), the mid-Brunhes event (430 kya) and the last glacial period (110-10 kya). I also find strong signals of local population history – recolonization and subdivision events – affecting their demography. In the subsequent study, I explore the grouse dynamics within the last glacial period in more detail by including more distant samples and using ecological modelling to track habitat distribution changes. I further uncover strong signals of local population history, with multiple fringe populations undergoing severe bottlenecks. I also determine that future climate change is expected to drastically constrict the distribution of the studied grouse. Lastly, I use whole genome sequencing to uncover 6 highly differentiated regions, containing 7 genes, hinting at their role in adaptation and speciation in three grouse taxa. I also locate a region of low differentiation, containing the Agouti pigmentation gene, indicating its role in the grouse plumage coloration.

As planted forests mature and are clearfelled in patches, second rotation tree crops (restocks) become available to black grouse, a species of conservation concern in the UK. Currently, only a limited amount is known about the resources provided by this habitat to black grouse and their broods. The aims of this study therefore, were to investigate the recovery of field-layer vegetation and the invertebrate population from restock through to canopy closure of planted trees, assess the duration of habitat availability and food resources to black grouse, and understand how forest management could improve provision. Changes to the abundance of predators resulting from habitat management were also considered. The comparative habitat quality of restocks was assessed in a wider landscape context. Field-layer vegetation in 72 restocks in two afforested areas in the north-east and the south-west of the Scottish Highlands was surveyed. On average, only 60% of ground in restocks was re-planted with second rotation trees, with the remainder left unplanted. Initial vegetation recovery was generally impeded by timber harvest residues (mainly brash), which comprised up to 50% of total ground cover two years after restock. Increased cover of heather Calluna vulgaris, often an important component of black grouse habitat, and decreased brash cover were recorded in areas of restocks where first-rotation timber was removed by cable-winch (compared with harvester and forwarder removal) and in planted areas (compared with areas left unplanted). Bilberry Vaccinium myrtillus and cotton grass Eriophorum spp. occurrence was recorded infrequently irrespective of restock age or management. Heather generally dominated the field-layer six years after restock, reaching a height and density reported to be suitable for black grouse nesting and brood cover in other studies. The onset of tree canopy closure as early as eight years suggests that suitable black grouse habitat availability in restocks is likely to be severely limited in duration. Brash removal, or break-up and re-distribution of the brash layer, positively affected the recovery of field-layer vegetation species potentially of use to black grouse. Extending the fallow period prior to restock resulted in an extended period of suitable habitat available to black grouse prior to canopy closure. However, habitat created by extending the fallow period also attracted a higher number of mammalian predators of black grouse. In the longer term, areas of restocks left unplanted should provide a valuable open-ground resource after canopy closure of the planted crop, although results suggest that management to prevent encroachment of naturally regenerating non-native trees is likely to be necessary. Invertebrate taxa selected by chicks in previous black grouse studies were available in all ages of restock studied. Taxa abundance differed as restocks aged and field-layer vegetation developed, although contrasting habitat preferences of taxa meant that each was affected differently by restock management. No single forest management method positively increased abundance of all taxa. Abundance of Lepidoptera larvae, a key food item for black grouse chicks, was positively related to dwarf shrub cover. An extended fallow period prior to restock should prolong increased larvae availability to chicks. Provision of preferred field-layer vegetation habitat and invertebrate abundance in restocks was comparable to habitat surrounding leks - areas likely to be occupied and utilised by black grouse. Restocks had a comparatively low occurrence of key plant species, including bilberry Vaccinium myrtillus and cotton grass Eriophorum spp. Cover of the dwarf shrub bog myrtle Myrica gale, positively associated with Lepidoptera larvae abundance in habitat surrounding leks, was absent from restocks. The abundance of other invertebrate taxa considered was similar between leks and restocks. Study findings are discussed with reference to black grouse conservation and commercial forestry systems in Europe. Management recommendations to improve habitat for black grouse in second rotation planted forests in Britain are provided.

Abstract Parasites are an inseparable part of the life of wild birds. They may cause morbidity, mortality or reduction in fecundity. Parasite distribution in hosts is typically not uniform and many host factors (e.g. age) may affect the pattern of distribution. Under certain conditions, parasites even have the potential to regulate the host population. The grouse species of Finnish forests — the capercaillie Tetrao urogallus , the black grouse Lyrurus tetrix and the hazel grouse Tetrastes bonasia — harbour several species of intestinal helminth parasites. The populations have fluctuated in cyclic manner but the mechanisms behind the cycles are largely unknown. I studied the interactions of forest grouse and their intestinal helminth parasites by using intestinal samples collected by hunters in five game management districts during eight years (1995–2002). The most common parasite species in the samples was the nematode Ascaridia compar. Also, three species of cestodes (Skrjabinia cesticillus, Paroniella urogalli and Hymenolepis sp.) were found. Large size, male gender and age over 1 year were connected with an increased probability and intensity of A. compar infection. Juvenile grouse were commonly infected with cestodes while in adults infections were quite rare. The influence of inbreeding on the susceptibility to parasite infections was studied in the capercaillie by analysing microsatellite heterozygosity. The less heterozygous birds were more likely to be infected with A. compar and were more intensely infected suggesting negative influence of inbreeding on parasite resistance. An indirect negative effect of parasites was found by comparing bags hunted with a trained dog or without a dog. Grouse infected by cestodes were significantly more common in the dog-assisted bag. Thus, cestode infection seemed to make grouse more vulnerable to canine predation. The interaction between grouse population dynamics and parasites was studied by analyzing the grouse densities obtained from annual wildlife counts and parasite indices. A. compar was most common and most abundant in the years of grouse population decline. The grouse population growth rate was negatively correlated with the annual mean abundance of A. compar. Relative survival but not breeding success decreased as the abundance of A. compar increased. The findings suggest that A. compar influences the dynamics of Finnish grouse even though regular cyclic dynamics are no longer evident. / Tiivistelmä Loiset kuuluvat erottamattomana osana luonnonvaraisten lintujen elämään. Ne voivat aiheuttaa sairautta, kuolleisuutta tai hedelmällisyyden alentumista. Tyypillisesti loiset ovat levinneet isäntäpopulaatioon epätasaisesti ja monet isännän ominaisuudet (esim. ikä) vaikuttavat levinneisyyteen. Tietyissä oloissa loiset voivat jopa säädellä isäntäpopulaatiotaan. Suomalaiset metsäkanalinnut — metso Tetrao urogallus, teeri Lyrurus tetrix ja pyy Tetrastes bonasia — ovat useiden suolistoloismatolajien isäntiä. Metsäkanapopulaatiot ovat vaihdelleet syklisesti, mutta syklejä aiheuttavat mekanismit ovat yhä tuntemattomia. Tutkin metsäkanalintujen ja niiden suolistoloisten välisiä vuorovaikutuksia käyttäen metsästäjien vuosina 1995–2002 viidestä eri riistanhoitopiiristä keräämiä suolistonäytteitä. Yleisin loislaji näytteissä oli kanalintusuolinkainen, Ascaridia compar. Myös kolme heisimatolajia (Skrjabinia cesticillus, Paroniella urogalli ja Hymenolepis sp.) todettiin. Suuri koko, koirassukupuoli ja yli yhden vuoden ikä olivat yhteydessä suurempaan kanalintusuolinkaistartunnan todennäköisyyteen ja voimakkuuteen. Nuorilla (alle 1 v.) linnuilla heisimadot olivat yleisiä, kun taas aikuisilla tartunnat olivat varsin harvinaisia. Sisäsiittoisuuden vaikutusta loistartuntaherkkyyteen tutkittiin metsolla mikrosatelliittiheterotsygotian perusteella. Vähemmän heterotsygoottiset metsot olivat todennäköisemmin ja voimakkaammin suolinkaisten infektoimia, mikä viittaa sisäsiittoisuuden negatiiviseen vaikutukseen loisten vastustuskykyyn. Loisten epäsuora haitallinen vaikutus havaittiin, kun verrattiin koiran kanssa ja ilman koiraa metsästettyä lintusaalista. Heisimadot olivat selvästi yleisempiä linnuilla, jotka oli metsästetty koiran kanssa kuin ilman koiraa metsästetyillä. Heisimatotartunta näytti siis altistavan metsäkanoja koiraeläinten saalistukselle. Metsäkanalintu- ja loispopulaatioiden välistä vuorovaikutusta tutkittiin analysoimalla vuosittaisia metsäkanatiheyksiä ja loisten runsautta. Kanalintusuolinkainen oli yleisimmillään ja runsaimmillaan metsäkanatiheyden laskuvuosina. Metsäkanapopulaation vuosittainen kasvuvauhti korreloi negatiivisesti kanalintusuolinkaisen vuosittaisen runsauden kanssa. Suhteellinen elossasäilyvyys laski kanalintusuolinkaisen runsauden lisääntyessä, mutta lisääntymistuloksen suhteen ei ollut samaa ilmiötä. Löydökset viittaavat siihen, että kanalintusuolinkaisella on vaikutusta suomalaisten metsäkanalintukantojen vaihteluihin, vaikka syklisiä kannanvaihteluja ei enää havaitakaan.

Black grouse (Tetrao tetrix) is a bird species with a lek mating system found in the Palearctic boreal taiga. It is assumed that it has a continuous distribution along Scandinavia and Siberia, whereas in Central Europe it has declined during the last decades. The primary objective of this thesis was to obtain a deeper understanding of the history, systematic classification and the genetic structure of black grouse on different geographical scales using microsatellites and control region mtDNA sequences (CR). I determined how much the mating system, habitat fragmentation and historical population processes have influenced the partitioning of genetic diversity in this species. Phylogeographical results are consistent with a demographic population expansion, and the patterns of postglacial dispersal suggest that a glacial refugium was located somewhere in central Asia, and from there black grouse spread out to Europe following the retreat of glacial ice sheets. I suggest that the two European black grouse subspecies, T. t. Tetrix and T. t. britannicus correspond to only one subspecies: T. t. tetrix, and that this lineage has diverged from T.t. viridanus, a subspecies found in Kazakhstan. The British population is significantly divergent from the remaining Eurasian samples for microsatellites but it is not for mtDNA. Therefore, they should regard as a separate Management Unit and not as a subspecies. Furthermore, British black grouse occur in three independent genetic units, corresponding to Wales, northern England/southern Scotland and northern Scotland. There was also genetic structure within Sweden. Habitat fragmentation is the main cause of population genetic structure in southern Swedish black grouse. In contrast, low levels of genetic differentiation and high connectivity were found in northern Sweden due to female-biased dispersal. On a finer geographical scale, I found genetic differences between leks due to a mixture of related and unrelated individuals within leks. However, mean relatedness values hardly differed from zero. Some leks were similar to one another and I interpret this as a result of variation in local reproductive success and philopatry. These factors would cause genetic structuring but this by itself would not reveal that kin selection is operating within black grouse leks.