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Attacks by wolves (Canis lupus) on dogs (C. familiaris) presumably are motivated both by preying and elimination of potential competitors. Regardless of these alternative motivations in wolves, the risk of attacks might be higher when the density of primary prey is low. We examined how many dogs do territorial wolves in Finland kill in relation to the population density of the most abundant ungulates, moose (Alces alces), white-tailed deer (Odocoileus virginianus), and roe deer (Capreolus capreolus). Most attacks by wolves on dogs take place in hunting with dogs. The number of wolf-killed dogs was in highly significant negative relationship to the population density of white-tailed deer and to total ungulate biomass per unit area which is largely determined by the density of white-tailed deer. Our results indicate that abundant wild prey would decrease the risk at which wolves attack dogs. On the other side of the coin prevail two hard facts which wildlife managers had to take a notice. White-tailed deer, although a potential mitigator of wolf–human conflict, is an alien species and a partner in > 6000 traffic collisions annually in Finland. One factor that seemed to increase the risk of wolf attacks on dogs is the low ungulate density in regions where moose is the only remarkable ungulate prey. Higher moose densities could decrease the risk of attacks, but on the other hand, higher densities could increase the risk of serious traffic collisions and browsing damages in forests.

Sociality in animal populations is a continuum, and interactions between conspecifics are meaningful for all vertebrates. Ignorance of social structures can lead to misunderstanding their ecology and, consequently, to unsuccessful species management. Here, we combined genetic and spatial data on radio-collared brown bears (Ursus arctos) to investigate kin-related home range overlap and kin-related centroid distance within central and eastern Finland. We found that the extent of home range overlap was positively correlated with relatedness among adult females. In addition, home range centroid distance decreased as relatedness increased. Moreover, there were significant differences between the two studied regions: female brown bears in central Finland were more closely related to each other, and the sizes of their home ranges were larger than those in eastern Finland. The smaller home ranges and lower degree of relatedness among bears in eastern Finland might be a result of the substantially higher hunting pressure in the area, combined with immigration of new unrelated individuals from Russia.

Transboundary connectivity is a key component when conserving and managing animal species that require large areas to maintain viable population sizes. Wolves Canis lupus recolonized the Scandinavian Peninsula in the early 1980s. The population is geographically isolated and relies on immigration to not lose genetic diversity and to maintain long term viability. In this study we address (1) to what extent the genetic diversity among Scandinavian wolves has recovered during 30 years since its foundation in relation to the source populations in Finland and Russia, (2) if immigration has occurred from both Finland and Russia, two countries with very different wolf management and legislative obligations to ensure long term viability of wolves, and (3) if immigrants can be assumed to be unrelated. Using 26 microsatellite loci we found that although the genetic diversity increased among Scandinavian wolves (n = 143), it has not reached the same levels found in Finland (n = 25) or in Russia (n = 19). Low genetic differentiation between Finnish and Russian wolves, complicated our ability to determine the origin of immigrant wolves (n = 20) with respect to nationality. Nevertheless, based on differences in allelic richness and private allelic richness between the two countries, results supported the occurrence of immigration from both countries. A priori assumptions that immigrants are unrelated is non-advisable, since 5.8% of the pair-wise analyzed immigrants were closely related. To maintain long term viability of wolves in Northern Europe, this study highlights the potential and need for management actions that facilitate transboundary dispersal.

Background: Understanding the processes that lead to hybridization of wolves and dogs is of scientific and management importance, particularly over large geographical scales, as wolves can disperse great distances. However, a method to efficiently detect hybrids in routine wolf monitoring is lacking. Microsatellites offer only limited resolution due to the low number of markers showing distinctive allele frequencies between wolves and dogs. Moreover, calibration across laboratories is time-consuming and costly. In this study, we selected a panel of 96 ancestry informative markers for wolves and dogs, derived from the Illumina CanineHD Whole-Genome BeadChip (174 K). We designed very short amplicons for genotyping on a microfluidic array, thus making the method suitable also for non-invasively collected samples. Results: Genotypes based on 93 SNPs from wolves sampled throughout Europe, purebred and non-pedigree dogs, and suspected hybrids showed that the new panel accurately identifies parental individuals, first-generation hybrids and first-generation backcrosses to wolves, while second- and third-generation backcrosses to wolves were identified as advanced hybrids in almost all cases. Our results support the hybrid identity of suspect individuals and the non-hybrid status of individuals regarded as wolves. We also show the adequacy of these markers to assess hybridization at a European-wide scale and the importance of including samples from reference populations. Conclusions: We showed that the proposed SNP panel is an efficient tool for detecting hybrids up to the third-generation backcrosses to wolves across Europe. Notably, the proposed genotyping method is suitable for a variety of samples, including non-invasive and museum samples, making this panel useful for wolf-dog hybrid assessments and wolf monitoring at both continental and different temporal scales.

Knowledge about the connectivity among natural populations is essential to identify management units for effective conservation actions. Conservation-minded management has led to the recovery of large carnivore populations in northern Europe, possibly restoring connectivity between the two separated, but expanding brown bear (Ursus arctos) populations on the Scandinavian peninsula to the west and Karelia, a part of the large Eurasian population, to the east. The degree of connectivity between these populations has been poorly understood, therefore we investigated the extent of connectivity between the two populations using autosomal microsatellites and Y chromosome haplotypes in 924 male bears (the dispersing sex), sampled during a period of 12 years (2005–2017) across the transborder area where these two populations meet. Our results showed that the two populations are not genetically isolated as reported in earlier studies. We detected recent asymmetrical gene flow at a rate (individuals per generation) of 4.6–5.5 (1%) from Karelia into Scandinavia, whereas the rate was approximately 27.1–34.5 (8%) in the opposite direction. We estimated historical gene flow of effective number of migrants to be between 1.7 and 2.5 between the populations. Analyses of Y chromosome markers supported these results. Successful recovery and expansion of both populations led to the restoration of connectivity, however, it is asymmetric, possibly due to different recovery histories and population densities. By aligning monitoring between neighboring countries, we were able to better understand the biological processes across the relevant spatial scale.

Hybridization and admixture can threaten the genetic integrity of populations and be of particular concern to endangered species. Hybridization between grey wolves and dogs has been documented in many wolf populations worldwide and is a prominent example of human‐mediated hybridization between a domesticated species and its wild relative. We analysed whole‐genome sequences from >200 wolves and >100 dogs to study admixture in Fennoscandian wolf populations. A principal component analysis of genetic variation and Admixture showed that wolves and dogs were well‐separated, without evidence for introgression. Analyses of local ancestry revealed that wolves had <1% mixed ancestry, levels comparable to the degree of mixed ancestry in many dogs, and likely not resulting from recent wolf–dog hybridization. We also show that the founders of the Scandinavian wolf population were genetically inseparable from Finnish and Russian Karelian wolves, pointing at the geographical origin of contemporary Scandinavian wolves. Moreover, we found Scandinavian‐born animals among wolves sampled in Finland, demonstrating bidirectional gene flow between the Scandinavian Peninsula and eastern countries. The low incidence of admixture between wolves and dogs in Fennoscandia may be explained by the fact that feral dogs are rare in this part of Europe and that careful monitoring and management act to remove hybrids before they backcross into wolf populations.

The threat that wolves (Canis lupus) pose to hunting dogs is one reason why Finnish hunters have negative attitudes towards wolves and one of the potential motivations for the illegal killing of wolves. During 2010–2017, wolves killed an average of 38 dogs (range 24–50) per year in Finland. Most of the attacks (91%) were directed at hunting dogs during the hunting season. To decrease the risk of attacks, the last seven positions (one position per hour) of GPS-collared wolves were accessible to the public with a 5 × 5 km resolution during the hunting seasons (from August 20th to February 28th) of 2013/2014 (from September 2nd onwards), 2015/2016, 2016/2017 and 2017/2018. The link was visited more than 1 million times in 3 of the 4 seasons. Fatal attacks on dogs occurred on 17% of the days during the hunting seasons of our study (n = 760 days). Both the attacks and visits peaked in September–November, which is the primary hunting season in Finland. According to the general linear model, the number of daily visits to the website was higher on days when fatal attacks occurred than on other days. Additionally, season and the number of days passed from the first day of the season were significantly related to the daily visits. Visits were temporally auto-correlated, and the parameter values in the model where the dependent variable was the number of visits on the next day were only slightly different from those in the first model. A two-way interaction between season and attack existed, and the least squares means were significantly different in 2017/2018. The change in daily visits between consecutive days was related only to the number of days from the beginning of the season. We examined whether this kind of service decreased dog attacks by wolves. Wolf attacks were recorded in 32% of the wolf territories, where at least one wolf had been collared (n = 22). However, within the territories without any GPS-collared wolves, the proportion of territories with wolf

After decades, even centuries of persecution, large carnivore populations are widely recovering in Europe. Considering the recent recovery of the wolverine (Gulo gulo) in Finland, our aim was to evaluate genetic variation using 14 microsatellites and mtDNA control region (579 bp) in order (1) to determine whether the species is represented by a single genetic population within Finland, (2) to quantify the genetic diversity, and (3) to estimate the effective population size. We found two major genetic clusters divided between eastern and northern Finland based on microsatellites (FST = 0.100) but also a significant pattern of isolation by distance. Wolverines in western Finland had a genetic signature similar to the northern cluster, which can be explained by former translocations of wolverines from northern to western Finland. For both main clusters, most estimates of the effective population size Ne were below 50. Nevertheless, the genetic diversity was higher in the eastern cluster (HE = 0.57, AR = 4.0, AP = 0.3) than in the northern cluster (HE = 0.49, AR = 3.7, AP = 0.1). Migration between the clusters was low. Two mtDNA haplotypes were found: one common and identical to Scandinavian wolverines; the other rare and not previously detected. The rare haplotype was more prominent in the eastern genetic cluster. Combining all available data, we infer that the genetic population structure within Finland is shaped by a recent bottleneck, isolation by distance, human-aided translocations and postglacial recolonization routes.

Wolves’ attacks on hunting dogs are a major conflict between wolves and hunters in Northern Europe. The reasons affecting the risk of wolves’ (Canis lupus) attacks on hunting dogs are poorly known. We examined whether the number of wolves in territory, the territory size, and the space use of GPS-collared wolves affected the risk of fatal attacks during hunting season in Finland. We found that attacks were more frequent within the periphery than core of the territory. Based on locations of GPS-collared wolves, wolves spent 8% (range 2–12%, n = 17) of time within the border zone (11% of the territory area) of their territory. Forty-five percent of all attacks considered (n = 11) took place within this zone which was high proportion as compared to the distribution of wolf locations. This result might reflect wolves’ motivational status to eliminate potential competitors when they are moving near territory boundaries.

Illegal killing of wolves (hereinafter ’wolf poaching’) in human-inhabited areas where wolves are also legally harvested is a special case of wildlife crime. This doctoral thesis examines wolf poaching in Finland and Sweden from the ecological perspective. In the first paper, we examined the causes of mortality among collared Finnish wolves and the role of estimated poaching rates on population changes. The second paper related the likelihood of being poached to covariates expressing different dimensions of the wolf conflict at two spatial scales (territory and country level) in Finland. Third paper turns the focus to Sweden, where we examined the disappearances of adult wolves in relation to population size, legal harvest and inbreeding. The first two studies were done in collaboration with the Natural Resources Institute Finland (Luke) and the third with the Scandinavian wolf research project SKANDULV. Poaching outnumbered other causes of death. Most poaching cases were unverified. Other causes of death were legal harvest, traffic and natural mortalities. Both populations had a relatively high number of wolves with unknown fates. Inbreeding was not related to the disappearances of adult wolves in Sweden. Remoteness to human inhabitation and the detectability of the wolves from the forest roads (road crossings by wolves) increased the likelihood of poaching in Finland. Adult wolves suffered high risk of poaching in both populations. Risk was highest in early spring in Finland. Larger population size increased and the number of legally harvested wolves decreased poaching in both countries. Poaching seemed to limit the study populations despite the management efforts that used legal hunting as a tool to increase tolerance towards wolves., Tiivistelmä Suden ja ihmisen rinnakkainelo johtaa konflikteihin, joiden lievittämiseksi susikantoja rajoitetaan usein luvallisella metsästyksellä. Suteen kohdistuu lisäksi laitonta tappamista eli salametsästystä. Tämä väitöstutkimus selvittää salametsästyksen ekologisia vaikutuksia Suomen ja Ruotsin susipopulaatioihin. Ensimmäisessä osatyössä selvitimme suomalaisten pantasusien kuolinsyitä, arvioimme salametsästyksen voimakkuutta ja sen vaikutuksia populaatiotasolla. Toisessa osatyössä tarkastelimme susikonfliktiin liittyvien ennustetekijöiden vaikutusta laittomasti tapetuksi tulemisen riskiin reviiritasolla ja koko Suomen mittakaavassa. Kaksi ensimmäistä osatyötä tehtiin Luonnonvarakeskuksen (Luke) suurpetotutkimuksessa. Kolmas osatyö tehtiin osana skandinaavista susitutkimushanketta (SKANDULV). Siinä selvitimme populaatiokoon, luvallisen pyynnin ja sukusiittoisuuden vaikutuksia aikuisten susien katoamiseen Ruotsissa. Salametsästys oli susien yleisin kuolinsyy, mutta suurin osa tapauksista jää toteen näyttämättä. Muita kuolinsyitä olivat luvallinen metsästys, liikenne ja luonnolliset kuolinsyyt. Aineistoissa oli runsaasti kohtaloltaan tuntemattomaksi jääneitä yksilöitä. Yksilöiden sukusiittoisuusaste ei ollut yhteydessä susien katoamisiin Ruotsissa. Syrjäinen sijainti ja susien havaittavuus metsätiestöltä lisäsivät laittoman tapon todennäköisyyttä Suomessa. Salametsästysriski oli korkein kevättalvella. Aikuisilla susilla oli huomattavan korkea riski tulla laittomasti tapetuksi. Tutkimuksen perusteella salametsästyksen määrää selittävät erityisesti susipopulaation kulloinenkin koko ja luvalliset pyyntimäärät. Suurempi susikanta lisäsi salametsästystä ja metsästyslupien määrä vähensi sen riskiä. Salametsästys vaikuttaa säädelleen susikantoja siitä huolimatta, että susikonfliktia on pyritty lieventämään luvallisella metsästyksellä.

The number of effective breeders (Nb) and effective population size (Ne) are population parameters reflective of evolutionary potential, susceptibility to stochasticity, and viability. We have estimated these parameters using the linkage disequilibrium‐based approach with LDNE through the latest phase of population recovery of the brown bears (Ursus arctos) in Finland (1993–2010; N = 621). This phase of the recovery was recently documented to be associated with major changes in genetic composition. In particular, differentiation between the northern and the southern genetic cluster declined rapidly within 1.5 generations. Based on this, we have studied effects of the changing genetic structure on Nb and Ne, by comparing estimates for whole Finland with the estimates for the two genetic clusters. We expected a potentially strong relationship between estimate sizes and genetic differentiation, which should disappear as the population recovers and clusters merge. Consistent with this, our estimates for whole Finland were lower than the sum of the estimates of the two genetic clusters and both approaches produced similar estimates in the end. Notably, we also found that admixed genotypes strongly increased the estimates. In all analyses, our estimates for Ne were larger than Nb and likely reflective for brown bears of the larger region of Finland and northwestern Russia. Conclusively, we find that neglecting genetic substructure may lead to a massive underestimation of Nb and Ne. Our results also suggest the need for further empirical analysis focusing on individuals with admixed genotypes and their potential high influence on Nb and Ne.

Background: Carnivores are re-establishing in many human-populated areas, where their presence is often contentious. Reaching consensus on management decisions is often hampered by a dispute over the size of the local carnivore population. Understanding the reproductive dynamics and individual movements of the carnivores can provide support for management decisions, but individual-level information can be difficult to obtain from elusive, wide-ranging species. Non-invasive genetic sampling can yield such information, but makes subsequent reconstruction of population history challenging due to incomplete population coverage and error-prone data. Here, we combine a collaborative, volunteer-based sampling scheme with Bayesian pedigree reconstruction to describe the pack dynamics of an establishing grey wolf (Canis lupus) population in south-west Finland, where wolf breeding was recorded in 2006 for the first time in over a century. Results: Using DNA extracted mainly from faeces collected since 2008, we identified 81 individual wolves and assigned credible full parentages to 70 of these and partial parentages to a further 9, revealing 7 breeding pairs. Individuals used a range of strategies to obtain breeding opportunities, including dispersal to established or new packs, long-distance migration and inheriting breeding roles. Gene flow occurred between all packs but inbreeding events were rare. Conclusions: These findings demonstrate that characterizing ongoing pack dynamics can provide detailed, locally-relevant insight into the ecology of contentious species such as the wolf. Involving various stakeholders in data collection makes these results more likely to be accepted as unbiased and hence reliable grounds for management decisions.

Wild populations of large carnivores in Europe were almost wiped out during the last centuries. Nowadays, the number of brown bears in North and Eastern Europe has increased, and the current situation suggests that these populations have recovered or are in the process of recovery. Knowledge of the population genetic consequences of demographic recovery in large carnivores, especially across national borders and on broader geographical scales, is still limited. In this study, we collected 3,757 fecal and hair samples as well as 881 tissue samples from brown bears across Northern Europe, with a focus on the Finnish population and neighboring areas, to investigate the population structure, connectivity, and genetic diversity on a spatial as well as a temporal scale. Bayesian clustering analysis of the population structure suggested the division of brown bears in Northern Europe into several genetic clusters, and the subdivision of the Finnish population into a northern and southern subpopulation. The estimation of gene flow pointed to better connectivity of the bears between Southern Finland and Western Russia, while migration between Scandinavia and Northern Finland as well as between Scandinavia and Southern Finland/Western Russia appeared to be restricted. Genetic clusters identified in Finland, Russia and Northern Norway displayed high genetic diversity, which was among the highest reported in wild brown bears. Recovery of the Finnish population has been accompanied by a detected range expansion towards the north, while genetic differentiation between clusters has decreased and genetic diversity has increased in the southern population, suggesting expansion from the south. Our results demonstrated that the immigration of bears from Russia still plays a major role in the Finnish bear population; however, connectivity between the Finnish-Russian population and Scandinavian bears appears to be restricted and should be improved, as well as regularly monitored, Tiivistelmä Suurpetojen luonnonpopulaatiot hävisivät Euroopasta melkein kokonaan viimeisten vuosisatojen aikana. Ruskeakarhujen määrä on viime aikoina kasvanut Pohjois- ja Itä-Euroopassa, ja karhupopulaatiot ovat toipuneet tai toipumassa. Tieto demografisen toipumisen geneettisistä seurauksista populaatioissa on varsin rajoittunutta etenkin laajemmassa maantieteellisessä mittakaavassa, yli valtiorajojen. Keräsimme tätä tutkimusta varten 3757 uloste- ja karvanäytettä ja 881 kudosnäytettä Suomesta ja sen lähialueilta. Tarkoituksenamme oli kartoittaa Pohjois-Euroopan karhupopulaatioiden geneettistä rakennetta ja monimuotoisuutta, sekä populaatioiden välisiä yhteyksiä huomioiden ajallinen ja maantieteellinen ulottuvuus. Bayesiläisen ryhmittelyanalyysin perusteella Pohjois-Euroopan karhut jakaantuvat useaan geneettiseen ryhmään. Suomen populaatiossa erottuivat eteläinen ja pohjoinen alapopulaatio. Analyysit geenivirran määrästä osoittivat, että Etelä-Suomen ja Länsi-Venäjän karhupopulaatiot ovat yhteneväisemmät, kun taas migraatio Skandinavian ja Pohjois-Suomen sekä Etelä-Suomen ja Länsi-Venäjän välillä vaikuttaisi olevan rajoittunutta. Suomesta, Venäjältä ja Pohjois-Norjasta tunnistetut alaryhmät olivat geneettisesti hyvin monimuotoisia, ja muuntelu oli korkeampaa kuin koskaan aiemmin karhuilla havaittu. Suomen karhupopulaation toipuessa ja levitessä pohjoiseen, geneettinen erilaistuminen maan sisällä on vähentynyt ja eteläisen alapopulaation monimuotoisuus kasvanut. Tämä viittaa populaation laajentumiseen etelästä käsin. Tulosten perusteella karhujen tulomuutto Venäjältä on yhä tärkeää Suomen populaatiolle. Suomen ja Venäjän karhupopulaatioiden yhteyttä Skandinavian karhupopulaatioihin tulisi seurata ja parantaa.

Rare and elusive species are difficult to study, because they are usually secretive, solitary, occur at low densities and have large home ranges. Wolverines (Gulo gulo) can both hunt and scavenge for food. In Fennoscandia, wolverines co-exist with either wild or semi-domesticated reindeer, which constitute their most important winter food. Approximately half of the 180–220 Finnish wolverines are found in northern Finland within the reindeer management area. However, the other half of the population is distributed in eastern and central Finland, and the ecology of wolverines especially in this area is poorly known. This research examined the habitat selection, diet and population genetics of wolverines in northern and eastern Finland. The results suggest that wolf presence is one of the most important variables influencing the habitat selection of wolverines. This finding supports the speculative idea that wolverines might benefit from being sympatric with wolves through increased scavenging opportunities. Furthermore, both the reproductive status of wolverines and the availability of different prey items were found to affect the wolverine diet. In northern Finland, semi-domesticated reindeer and mountain hare were the most frequently utilized prey species for breeding female wolverines. In eastern Finland, the most important food source for breeding females was moose carrion, whereas males and non-breeding females heavily utilized mountain hares. These results support the predictions of the optimal foraging theory, suggesting that wolverines opportunistically utilize the food source that is most energy-efficiently available. In areas with a low density of medium-sized ungulates, scavenging of wolf- and human-killed carrion plays an essential role in food acquisition by wolverines. According to the results of a population genetics investigation, two wolverine subpopulations exist in Finland: a northern and an eastern one. The overall genetic variability was, Tiivistelmä Ahma (Gulo gulo) tunnetaan sekä haaskansyöjänä että keskikokoisia hirvieläimiä ja pienriistaa saalistavana petona. Fennoskandian alueella peuran kesy tai villi muoto on ahman merkittävin saalislaji. Noin puolet Suomen 180–220 ahmasta elää Pohjois-Suomessa poronhoitoalueella, ja loput Itä- ja Keski-Suomessa. Poronhoitoalueen ulkopuolella elävien ahmojen ekologiaa on tutkittu erityisen vähän. Väitöstyössäni tarkastelin ahman habitaatinvalintaa, ruokavaliota ja populaatiogenetiikkaa pääasiassa Pohjois- ja Itä-Suomen alueilla. Tutkimukseni tulokset osoittivat, että suden läheisyys oli yksi tärkeimmistä ahman habitaatinvalintaan vaikuttavista tekijöistä. Tämä havainto tukee hypoteesia, jonka mukaan ahma saattaisi hyötyä susien läheisyydestä suuremman haaskatiheyden ansiosta. Todennäköisesti ahmat elävät mielellään samoilla seuduilla susien kanssa, mutta kaihtavat läheistä kanssakäymistä välttääkseen killansisäisen saalistuksen. Tutkimusteni perusteella sekä ahman lisääntymistila että alueen saaliseläinten saatavuus vaikuttivat ahman ruokavalioon. Poro ja metsäjänis olivat lisääntyvien ahmanaaraiden tärkein ravintokohde Pohjois-Suomessa. Itä-Suomessa merkittävin lisääntyvien naaraiden ravintokohde oli hirvi, jota ahmat hyödyntävät lähinnä haaskojen muodossa, mutta urokset ja ei-lisääntyvät naaraat sen sijaan saalistivat eniten metsäjänistä. Ahmat siis näyttävät hyödyntävän opportunistisesti sitä ravintokohdetta, joka kullakin alueella on energiatehokkainta saavuttaa. Susien ja ihmisten jälkeensä jättämät haaskat ovat merkittävä ravintolähde ahmoille alueilla, joilla keskikokoisten hirvieläinten tiheydet ovat alhaisia. Ahman populaatiogeneettinen tutkimus osoitti, että Suomen ahmat ovat geneettisesti jakautuneet kahteen alapopulaatioon, pohjoiseen ja itäiseen. Ahmakannan geneettinen monimuotoisuus oli pientä, ja molemmissa alapopulaatiossa oli nähtävissä merkkejä äskettäisestä pullonkaulailmiöstä. Populaatioiden välisen geenivirran määrän tulisi olla nykyistä k

The grey wolf is the most widely distributed of all land mammals and is a habitat generalist that inhabits all the vegetation types of the Northern Hemisphere. Wolves also breed well and have the potential to rapidly expand to new areas. In Finland, the wolf is a game species, and as a result of Finland’s membership of the European Union, the wolf population is subject to very limited hunting due to the obligation to protect the species. Mainly for this reason, the wolf population in Finland has increased significantly in recent years. In particular, the birth rate has developed favourably and the number of litters increased from just four in 1996 to 20 in 2005. It also seems at present that the wolf population in Finland is no longer following the fluctuations in wolf numbers in Russian Karelia. The general aim of this dissertation is to provide applicable knowledge for wolf management and conservation purposes, and especially to examine the effects of human-modified landscapes on wolf population expansion in Finland. Various aspects of habitat selection were investigated in four sub-studies to gain a thorough insight into the space use and habitat needs of wolves. Species data came from a long-term wolf population study that included location information from 85 radio- and GPS-GSM-collared wolves from 1998 onwards, as well as track location data based on about 30 000 annual observations recorded with geographical coordinates by a local network of experts on large carnivores. I found that adaptability makes it possible for the wolf to live in the multiple-use, semi-wild forests of Finland and that no restrictions are imposed by the landscape on wolf population growth and expansion. In general, the results of my dissertation provide evidence that wolves tend to avoid the presence of human influence when establishing a territory and also when selecting their den site. However, as wolf numbers increase, conflict situations will more frequently occur between w, Tiivistelmä Susi on habitaattigeneralisti, jonka levinneisyys kattaa kaikki pohjoisen pallonpuoliskon kasvillisuustyypit. Sudet myös lisääntyvät hyvin ja voivat näin ollen levittäytyä nopeasti uusille alueille. Suomessa susi on riistalaji, jonka metsästys on rajoitettua Euroopan unionin jäsenyyden myötä. Lähinnä tästä syystä Suomen susikanta on kasvanut voimakkaasti 1990-luvun ja 2000-luvun alun aikana. Erityisesti syntyvyys on kasvanut: pentueiden määrä lisääntyi neljästä 20:een vuosien 1996 ja 2005 välillä. Näyttäisi siltä, ettei Suomen nykyinen susipopulaatio enää seuraa Venäjän Karjalassa tapahtuvia kannanvaihteluita. Väitöskirjan tavoitteena on ollut tuottaa susikannan hoidossa ja suojelussa käytettävissä olevaa tietoa ja erityisesti tutkia susikannan levittäytymistä ihmisen muokkaamassa ympäristössä Suomessa. Neljässä osatutkimuksessa on käsitelty suden habitaatinvalintaa eri näkökulmista, jotta saataisiin kokonaiskuva suden tilankäytöstä Suomessa. Tutkimusaineisto rakentui pitkäaikaisesta seuranta-aineistosta, jota on kerätty vuodesta 1989 lähtien. Vuosien 1989 ja 2010 välillä 85 sutta on pannoitettu radio- tai GPS-GSM -pannalla. Tutkimuksessa käytettiin hyväksi myös noin 30 000 vuosittain tehtyä jälkihavaintoa. Tutkimustulosten perusteella susi selviää hyvin Suomen monikäyttömetsissä, eivätkä maisematyypit itsestään rajoita susipopulaation kasvua ja levittäytymistä. Toisaalta tulokset osoittavat myös sen, että sudet välttelevät ihmistoimintoja valitessaan reviiri- tai pesäpaikkoja. Kuitenkin susien lukumäärän kasvaessa ristiriitatilanteet susien ja ihmisten välillä tulevat lisääntymään, vaikka susivahinkoriskin suuruus vaihteleekin Suomessa huomattavasti. Esimerkiksi lammastilojen susivahinkoriski riippuu sekä ympäristötekijöistä, että tilan sisäisistä tekijöistä. Kaiken kaikkiaan lisääntyvän susipopulaation esiintymisalue on Suomessa vähitellen kasvanut leviten itärajan tuntumasta kohti länttä. Tämän myötä maantieteellinen etäisyys Skandinavian susipopulaat