The thesis deals with food preferences of elk (Alces alces) in Czech zoological gardens and contains observation of food preferences in zoo Hluboká nad Vltavou. The results are compared with food offer and preference in the wilds described in literature. The first part of the thesis is aimed at literary review about elk, its characteristic, categorization, distribution and biology. It is focused mainly on the studies dealing with feeding habits and food preferences of elk in the wilds and captivity. The second part describes observation of food preferences of three elks in zoo Hluboká nad Vltavou, provided during 10 feedings. The food quality and composition of feed ration was also evaluated not only in zoo Hluboká nad Vltavou, but also in zoo Praha and zoo Brno (based on information from local zoo keepers). Analysis of variance showed that there are differences in feeding latency between various types of food. Elks preferred dried herbs, carrot and oak bark. On the contrary, elks did not preferred oat flakes and both granules for giraffes and elks. Recommendation for feed ration improving, including also food preferences of the elk were also suggested.

Abstract Spatial and temporal variation is a universal feature in most organisms in nature, commonly reflecting the past evolutionary history of the species as well as the prevailing environmental conditions. The purpose of this doctoral thesis study was to investigate the genetic and phenotypic variation, and to assess the roles of the different processes affecting them in the moose (Alces alces). Altogether 809 DNA samples of moose, gathered throughout Finland and the Republic of Karelia in Russia, were analysed with a variety of population genetic methods. Furthermore, the shape of the moose mandible was investigated with the help of geometric morphometrics using a subset of samples gathered from 179 moose in Finland. This study showed that the Finnish and especially the Karelian moose population harboured relatively high genetic diversity, albeit with clear regional differences in its spatial distribution. In the northern half of Finland, a secondary contact of two diverged mitochondrial lineages was revealed. The presence of the two lineages was interpreted to reflect the existence of allopatric refugia of moose during the Last Glacial Maximum and the subsequent bi-directional recolonisation of Fennoscandia. Furthermore, a spatially explicit Bayesian clustering analysis suggested existence of three genetic clusters, which were estimated to have split after the post-glacial recolonisation. The results also showed that past declines in the moose numbers during the 18th and 19th centuries led to population bottlenecks, leaving a genetic imprint. Thus, the present moose population in eastern Fennoscandia carries the signs of both ancient and more recent events in its genetic composition. Finally, a significant latitudinal shift was revealed in the shape of the moose mandible. The pattern was considered independent of the genetic clustering of the population. The main changes included an enlargement of the attachment surfaces of the muscles controlling biting and mastication, implying more effective mastication in the north compared with the south, possibly an adaptive response to a longer period of hard wintertime diet. The results of this thesis encourage continuation of studies on the moose in order to fully reveal the impact of particular historical events and especially anthropogenic factors on the genetic and phenotypic variation of this species. They also provide the starting point for ‘genetically enlightened’ moose management and conservation in Finland. / Tiivistelmä Lähes kaikilla eliölajeilla esiintyy ajallista ja paikallista muuntelua, joka on seurausta lajin evolutiivisesta historiasta ja vallitsevista ympäristöoloista. Tässä väitöskirjatutkimuksessa tutkin hirven (Alces alces) geneettistä ja fenotyyppistä muuntelua sekä niitä selittäviä taustatekijöitä populaatiogeneettisillä ja geometrisen morfometrian menetelmillä. Geneettisen aineiston muodostivat Suomesta ja Venäjän Karjalasta kerätyt 809 hirven DNA-näytteet. Fenotyyppisenä ominaisuutena tutkittiin hirven leukaluun muotoa yhteensä 179 alaleuasta. Geneettinen monimuotoisuus oli tutkimuksen mukaan Suomen ja erityisesti Karjalan hirvipopulaatiossa verrattain korkea, joskin alueelliset erot olivat varsin selviä. Pohjoisesta Suomesta löytyi kahta erilaistunutta mitokondrion DNA:n sukulinjaa, joiden arvioin erilaistuneen viimeisen jääkauden aikana, todennäköisesti erillisissä refugioissa, ja saapuneen aikoinaan Suomeen eri reittejä pitkin. Tämän ohella tuman DNA paljasti lisää alueellisia rakenteita; bayesilainen ryhmittelyanalyysi havaitsi hirvellä kolme erillistä alapopulaatiota. Näiden ryhmien arvioin kehittyneen vasta Suomen uudelleenasuttamisen jälkeen. Tämän tutkimuksen tulokset osoittivat myös, että historiallisesti tunnetut kannanromahdukset 1700- ja 1800-luvuilla johtivat populaation pullonkaulaan, joka jätti jälkensä hirven perimään. Itäisen Fennoskandian hirvipopulaation geneettiseen muunteluun ovat siis vaikuttaneet sen historian aikana niin jääkauden aikaiset kuin tuoreemmatkin tapahtumat. Tämän lisäksi hirven alaleuan muodossa havaittiin merkitsevä etelä-pohjoissuuntainen muutos. Tulosten mukaan purentaa ohjaavien lihasten kiinnityspinnat laajenevat pohjoista kohti siirryttäessä, mikä viittaisi siihen, että hirven leukojen puruvoima on pohjoisessa suurempi kuin etelässä. Ilmiö oli riippumaton populaation geneettisestä ryhmittyneisyydestä, ja se on mahdollisesti seurausta kovemman talviruokavalion aiheuttamasta adaptiivisesta vasteesta. Tämän väitöskirjan tulokset rohkaisevat jatkamaan aiheen tutkimusta, jotta eri historiallisten tapahtumien sekä eritoten ihmisvaikutuksen merkitys lajin geneettiseen ja fenotyyppiseen muunteluun voitaisiin selvittää perin pohjin. Lisäksi tulokset muodostavat lähtökohdan ’geneettisesti valistuneelle’ hirvikannan hoidolle Suomessa.

Browsing patterns in large herbivorous mammals are complex and decisions regarding food items occur at different levels depending on internal or external factors such as energy requirements and available food sources. A broader understanding of the behavior of animals living in habitats of economic interests for humans enables human exploitation with as little impact as possible on the animals' habitats. In this study, individual moose (Alces alces) tracks in the snow were surveyed in a mountainous environment with spruce-dominated forest, to map out what was browsed in relation to what was available in forests of two age classes: young and older. The following hypotheses were made: (1) every potential tree a moose passes will not be browsed and the browsed trees will be browsed to different degrees, and (2) the amount of available forage and the amount of used forage will differ between young and older forest. The moose tracks were followed on skis or on foot and all trees within browsing distance along a transect of 50 meters were counted and signs of browsing were recorded. Control transects (without fresh tracks) were also surveyed in the same forest areas where moose tracks were found. The data showed a higher proportion of available trees in young forest (60%) compared to older forest (40%) (all trees along the track transects), and a higher proportion of browsed trees in young forest (26%) compared to older forest (17%) (excluding Norway spruce, Picea abies). Downy birch (Betula pubescens) was the most abundant tree (56%) and Scots pine (Pinus silvestris) the least abundant (<1%). Only 20% of all available trees along the track transects were freshly browsed to some extent, and even among the more selected species no more than around 50% of all available trees were freshly browsed. In young forest, Eurasian aspen (Populus tremula), rowan (Sorbus aucuparia) and willow (Salix spp.) were the most selected by moose, while common juniper (Juniperus communis) was the least selected. In older forest, however, gray alder (Alnus incana) was the most selected, followed by willow and rowan, while downy birch was the least selected. No browsing on Ericaceous shrubs were recorded in any of the transects. The study underlines that many different factors influence moose browsing choices, but generally a varied diet is sought and forests of different age classes, not exclusively young forests, are used for winter foraging. / Betesmönster hos stora växtätande däggdjur är komplext och val kopplade till föda sker på olika nivåer beroende på inre och yttre faktorer så som energibehov och tillgänglig föda. En bredare förståelse om beteenden hos djur som lever i habitat där människan har ekonomiska intressen möjliggör mänsklig exploatering med så lite påverkan som möjligt på djurens livsmiljöer. I den här studien spårades individuella älgar (Alces alces) i snö i en fjällnära miljö med grandominerad skog för att kartlägga vad som betades i relation till vad som fanns tillgängligt inom två åldersklasser; ung och äldre skog. Jag ställde upp två hypoteser: (1) varje potentiellt födoträd som en älg passerar kommer inte att vara betat på och de betade träden kommer att vara betade i olika grad, samt (2) mängd tillgängligt foder och mängd nyttjat foder kommer skilja sig mellan ungskog och äldre skog. Älgspåren följdes på skidor eller till fots och alla träd inom betesavstånd längs en transekt om 50 meter räknades samt kontrollerades för bete. Även kontrolltransekter (utan färska spår) inventerades i samma skogsområden där älgspår återfunnits. Datan visade på en högre andel träd i ungskog (60%) jämfört med äldre skog (40%) (alla träd längs spårtransekterna), och en högre andel betade träd i ungskog (26%) jämfört med äldre skog (17%) (gran, Picea abies, undantaget). Glasbjörk (Betula pubescens) var det mest förekommande trädet (56%) och tall (Pinus silvestris) det minst förekommande (<1%). Enbart 20% av alla tillgängliga träd längs spårtransekterna var färskbetade i någon mån, och även hos de högre selekterade arterna var inte mer än runt 50% av alla tillgängliga träd färskbetade. I ungskog var asp (Populus tremula), rönn (Sorbus aucuparia) och sälg (Salix spp.) högst selekterade, medan en (Juniperus communis) var lägst selekterad. I äldre skog var däremot gråal (Alnus incana) högst selekterad följt av sälg och rönn, medan glasbjörk var lägst selekterad. Inget bete på bärris noterades längs någon av transekterna. Studien understryker att många olika faktorer påverkar betesval hos älg, men att en varierad diet generellt eftersträvas och skogar av olika åldersklasser, inte enbart ungskogar, nyttjas till födosök vintertid.

Genetic variation is a prerequisite for the viability and evolution of species. Information on population genetic patterns on spatial and temporal scales is therefore important for effective management and for protection of biodiversity. However, incorporation of genetics into management has been difficult, even though the need has been stressed for decades. In this thesis population genetic patterns in continuous environments are described, compared among species, and related to conservation management. The model systems are moose (Alces alces) in Sweden and multiple species in the Baltic Sea, with particular focus on the Northern pike (Esox lucius). The spatial scope of the studies is large, and much focus is dedicated towards comprehensive sampling over large geographic areas. The moose population in Sweden is shown to be divided into two major subpopulations, a northern and a southern one. Both subpopulations show genetic signals of major population bottlenecks, which coincide with known population reductions due to high hunting pressure (Paper I). The Northern pike in the Baltic Sea shows relatively weak, but temporally stable population genetic structure. An isolation by distance pattern suggests that gene flow primarily takes place among neighboring populations, either over shortest waterway distance or along the mainland coast, with island populations acting as stepping stones (Paper III). In a comparative study of seven Baltic Sea species no shared genetic patterns were found, either in terms of genetic divergence among or genetic diversity within geographic regions. These results complicate the incorporation of genetic data into management, because it suggests that no generalization can be made among species in the Baltic Sea, but that species-specific management is needed (Paper II). Over the last 50 years, 61 species in the Baltic Sea have been studied with respect to spatial genetic patterns. For over 20 of these species information of direct relevance for management is available. Relevant information is synthesized into management recommendations (Paper IV). This thesis provides vital information on spatial and temporal genetic structure for a number of ecologically and socio-economically important species. It shows that such information is important to consider species by species and that both local and metapopulation approaches are needed to effectively manage genetic diversity in e.g. moose and pike. Further, it identifies for which organisms in the Baltic Sea genetic information exists, how it can be used, and where important information is lacking. In order to successfully make use of genetic data in management, effective communication channels between academia and policy-makers are needed. / <p>At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 3: Manuscript. Paper 4: Manuscript.</p>

Betesskador på skog kostar både skogsägare och industri stora summor pengar. Ett sätt att minska betesskador kan vara bättre kontroll över populationsstorleken av klövvilt. I Sverige används flera olika etablerade viltinventeringsmetoder. Svagheten är att de baseras på trender, index och avskjutningar mer än att få fram en siffra på antalet djur. En tillförlitlig inventeringsmetod är av stor vikt som beslutsunderlag till förvaltningsplaner. Drönare är ett obemannat luftfartyg som kan utrustas med olika kameror och sensorer. Tidigare fältstudier har visat att värme från vilt kan plockas upp av en drönare utrustad med termisk kamera. Svårigheten har varit att få tillförlitlighet vid artbestämning av vilt med enbart termiska kamera. I denna studie utfördes en viltinventering av ett 506 ha stort område norr om Ljungby i Kronobergs län under januari 2020. Med hjälp av en drönare utrustad med både termisk och RGB kamera utfördes en totalinventering av området. Resultatet visar att kombinationen av både termisk och RGB kamera ökar tillförlitligheten vid artbestämning av vilt. Metodens svagheter är täta krontak av gran, gällande regelverk från transportstyrelsen samt drönarens begränsande batterikapacitet. Slutsatsen är att vilt säkrare kan identifieras och artbestämmas med kombinationen av termisk- och RGB kamera jämfört med enbart termisk kamera men att metoden behöver vidareutvecklas.

Šio darbo tikslas buvo išstudijuoti elninių žvėrių (briedžių, tauriųjų elnių ir europinių stirnų) gausumą 2009-2010 m. žiemojimo periodu Kuršėnų urėdijos miškuose ir palyginti skirtingų miškų gautus rezultatus. Tyrimai atlikti 18 miškų, naudojant McCain netiesioginę elninių žvėrių apskaitą pagal jų paliekamus ekskrementus bei Aldous medžių pažeidimo metodą. Pagal atliktus tyrimus, vidutinis elninių žvėrių tankumas Kuršėnų urėdijos miškuose 2009-2010 m. žiemojimo periodu buvo 14, 7 individų tūkstančiui hektarų. Briedžių tankumas tirtame regione buvo du kartus didesnis, lyginant su vidutiniu briedžių tankumu visoje Lietuvos teritorijoje, tauriųjų elnių – labai panašus, o stirnų buvo dvigubai mažesnis. Taigi, tirtuose miškuose yra gausu stambiųjų elninių, tačiau jie konkuruoja su stirnomis. Vidutinis briedžių ir tauriųjų elnių tankumas tirtuose miškuose atitinka ekologinį ir ūkiškai leistiną elninių žvėrių gausumą Lietuvos miškams, o stirnų yra du kartus mažesnis, tačiau, atsižvelgiant į mitybinę konkurenciją, taurusis elnias laikomas pagrindine elninių žvėrių rūšimi tirtoje vietovėje, tad pagrindinis dėmesys turėtų būti skiriamas jo populiacijų gausinimui. Elniniai žvėrys pagal mitybos specializaciją renkasi skirtingus biotopus. Didžiausias briedžių gausumas nustatytas krūmynuose, tauriųjų elnių – pievose, o europinių stirnų – dirbamų laukų biotopuose. Vidutinis dendrofloros panaudojimo pašarams intensyvumas Kuršėnų urėdijos miškuose buvo 18,5 %, bet netgi didžiausi pažeidimai... [toliau žr. visą tekstą] / The aim of this study was to find out the density of cervids during wintering period in Kušėnai forests and to compare gathered data with density in different forest. The research have been conducted in 18 forests, using McCain indirect record by excrement and Aldous trees damage method. Studies have shown that the average density of cervids in surveyed areas is 14.7 individuals per thousand hectares. Moose density in this region is twice higher than average moose density in whole Lithuania, red deer – very similar, roe deer density is half lower. Therefore, explored forests is rich of large ungulate mammals, but they compete with roe deers. The average abundance of red deer and moose satisfies commercially and ecologicaly acceptable ones, also roe deer density is more than two times smaller, bur in aspect of the competition diet, red deer is considered to be the main cervid species in studied forests, so more attention should be paid to increase their population. Cervids by specificity of their species in different forests choose similar biotopes. The highest abundance of moose were in scrubs, red deer – in meadows, roe deer – in cultivated fields. The average of tree damage in stydy areas is 18.5 percent, but even the highest nutrient load does not exceed the recommended rate (30–40 %), so there is no danger to natural forest regeneration.

<p>The moose (Alces alces) is very important both economically and ecologically, therefore all knowledge of moose is vital for future management of the moose population. Little is known about moose habitat selection in Sweden. In coastal southwestern Sweden growing human population and new infrastructure projects continuously threaten to fragment and isolate local moose populations. The habitat selection of 22 moose, 8 males and 14 females, in southwestern Sweden was studied from February 2002 until December 2005. The moose were captured and fitted with GPS-collars and positions were collected at 2-hour intervals. The number of moose positions totaled 71103 during the study period of 46 months. Data for individual animals were divided into four seasons: spring, summer, fall and winter based on climate and moose biology. A total of 125 moose seasonal home ranges were generated and habitat use within each of the generated home ranges was studied using Euclidean distance-based analysis. A reclassified digital landcover map was divided into the land use classes agriculture, clear-cut, coniferous forest, deciduous forest, mire and mountain. The results showed that there was a difference in habitat selection between males and females. Males were significantly closer to forest and clear-cuts compared to females. Both males and females selected clear-cuts and avoided agriculture within their home ranges.</p> / <p>Älgen är en viktig art, både ekonomiskt och ekologiskt, och all kunskap är viktig för att även i framtiden kunna sköta en sund älgstam. Trots flertalet studier finns det många frågetecken om älgens habitatval i Sverige. En ökad exploateringstakt och nya infrastrukturprojekt hotar att fragmentera och isolera populationer av älg. Habitatvalet hos 22 älgar, 8 tjurar och 14 kor, i sydvästra Sverige studerades mellan februari 2002 och december 2005. Älgarna sövdes och utrustades med GPS-sändare, deras positioner registrerades varannan timma och det totala antalet positioner under den 46 månader långa studietiden var 71103 stycken. Data från varje älg delades in i 4 säsonger; vår, sommar, höst och vinter, baserat på klimat och älgens biologi. Totalt genererades 125 hemområden baserade på säsong, och valet av habitat inom varje hemområde studerades med hjälp av Euclidean distance-based analysis. En omklassificerad digital marktäckedata användes som var indelad i 6 olika klasser; odlad mark, hygge, barrskog, lövskog, myrmark och berg i dagen. Resultaten visade att det var skillnad mellan könen i hur de väljer habitat. Tjurarna var signifikant närmare barrskog och hyggen än korna, men både tjurar och kor selekterade för hyggen och undvek odlad mark inom deras hemområden.</p>

The moose (Alces alces) is very important both economically and ecologically, therefore all knowledge of moose is vital for future management of the moose population. Little is known about moose habitat selection in Sweden. In coastal southwestern Sweden growing human population and new infrastructure projects continuously threaten to fragment and isolate local moose populations. The habitat selection of 22 moose, 8 males and 14 females, in southwestern Sweden was studied from February 2002 until December 2005. The moose were captured and fitted with GPS-collars and positions were collected at 2-hour intervals. The number of moose positions totaled 71103 during the study period of 46 months. Data for individual animals were divided into four seasons: spring, summer, fall and winter based on climate and moose biology. A total of 125 moose seasonal home ranges were generated and habitat use within each of the generated home ranges was studied using Euclidean distance-based analysis. A reclassified digital landcover map was divided into the land use classes agriculture, clear-cut, coniferous forest, deciduous forest, mire and mountain. The results showed that there was a difference in habitat selection between males and females. Males were significantly closer to forest and clear-cuts compared to females. Both males and females selected clear-cuts and avoided agriculture within their home ranges. / Älgen är en viktig art, både ekonomiskt och ekologiskt, och all kunskap är viktig för att även i framtiden kunna sköta en sund älgstam. Trots flertalet studier finns det många frågetecken om älgens habitatval i Sverige. En ökad exploateringstakt och nya infrastrukturprojekt hotar att fragmentera och isolera populationer av älg. Habitatvalet hos 22 älgar, 8 tjurar och 14 kor, i sydvästra Sverige studerades mellan februari 2002 och december 2005. Älgarna sövdes och utrustades med GPS-sändare, deras positioner registrerades varannan timma och det totala antalet positioner under den 46 månader långa studietiden var 71103 stycken. Data från varje älg delades in i 4 säsonger; vår, sommar, höst och vinter, baserat på klimat och älgens biologi. Totalt genererades 125 hemområden baserade på säsong, och valet av habitat inom varje hemområde studerades med hjälp av Euclidean distance-based analysis. En omklassificerad digital marktäckedata användes som var indelad i 6 olika klasser; odlad mark, hygge, barrskog, lövskog, myrmark och berg i dagen. Resultaten visade att det var skillnad mellan könen i hur de väljer habitat. Tjurarna var signifikant närmare barrskog och hyggen än korna, men både tjurar och kor selekterade för hyggen och undvek odlad mark inom deras hemområden.

Moose in Utah represent the southernmost naturally occurring populations of moose in the world. Concerns over possible numeric declines and a paucity of baseline data on moose in the state prompted the Utah Division of Wildlife Resources to initiate a study of moose demography in collaboration with Utah State University. The objectives of this study were to 1) determine reproductive rates of moose in Utah and the factors which influence them, and 2) combine aerial count data from multiple management units within the state to identify factors which influence interannual variation in population growth rates. We constructed generalized linear models to relate maternal body condition and age to reproductive success. We found that body condition (P = 0.01) and age (P = 0.02) contributed significantly to the probability of pregnancy and the best model describing this relationship was nonlinear. Body condition also related positively to subsequent calving (P = 0.08) and recruitment (P = 0.05), but model selection suggested the relationship for these metrics was best described by linear models. A meta-analysis of moose reproductive rates in North America suggested that reproductive rates declined significantly with latitude (P ≤ 0.01), i.e. as populations approached their southern range limit. We used Bayesian state-space models to combine moose count data from different management units to estimate statewide population dynamics between 1958 and 2013. This approach incorporated uncertainty in population counts arising from observation error. Population density and warm winter temperatures negatively influenced population growth rate with a high degree of confidence; 95% Bayesian Credible Intervals for these variables did not overlap zero. Short-term projections of moose abundance in the state suggested that the population will likely remain stable despite projected increases in winter temperature. Results from this study will aid managers in achieving management objectives as well as future decision making. The unique characteristics of the population also have application toward understanding the dynamics of populations of cold-adapted species at their southern range limit.

As human populations continue to grow and encroach into wildlife habitats, instances of human-wildlife conflict are on the rise. Increasing numbers of reported wildlife-vehicle collisions (WVCs) provide tangible evidence of anthropogenic impacts on wildlife as well as increasing threats to human health and safety. Increasing WVCs are of particular concern, especially those involving large-bodied ungulates such as moose (Alces spp.), because of the increased risk of property damage, personal injuries, and human fatalities. Motorists directly involved in a WVC are at risk of injury or mortality, but other motorists are also put at risk due to road obstructions and traffic congestion associated with WVCs. Mitigating these impacts on motorists and wildlife requires investigation into the temporal and spatial factors leading to WVCs. In Alaska, most WVCs involve moose (Alces alces), a large bodied ungulate capable of threatening human life when involved in a collision. Each moose-vehicle collision (MVC) in Alaska is estimated to cost $33,000 in damages. With this analysis, I analyzed the plethora of factors contributing to moose and motorist occurrence on the road system and motorist detection based on a historical dataset of MVC reports throughout Alaska from 2000 to 2012 and a dataset of field-derived measurements at MVC locations within the Matanuska-Susitna Borough from 2016 to 2018. My first analysis focused on the daily and annual trends in MVC rates as compared to expected moose and human behavioral patterns with a focus on guiding mitigation strategies. Fifty percent of the MVCs reported between 2000 and 2012 occurred where the commuter rush hours overlapped with dusk and dawn in winter, and the artificial lighting differences between boroughs suggest a link between artificial lighting and reduced MVCs. To focus more specifically on roadside features contributing to MVC risk, I collected and analyzed local and regional scale land cover and road geometry data at reported MVC sites in an area with a rapidly growing human population. I compared these data to similar data collected at random locations near documented MVC sites and at locations where moose that were fitted with global-positioning system (GPS) transmitters crossed highways. I used generalized additive mixed models to delineate which of the variables impacted the risk of both moose road crossings and MVCs. Moose road crossings were influenced by approximations of spatial, seasonal, and daily moose density as well as the proportion of deciduous-coniferous and coniferous forest in the area and the number of possible corridor or land cover types surrounding the site. The best MVC risk model was described by expected seasonal and daily changes in moose density and local scale measurements, including the sinuosity of the road, the height of vegetation near the road, and the angle between the road surface and the roadside. Together this information should guide transportation and urban planners in the Matanuska-Susitna Borough to use roadside vegetation removal, seasonal speed reduction, improved lighting strategies, dynamic signage, or partnerships with mobile mapping services to reactively reduce MVCs and to focus future road planning in areas with lower moose abundance and build roads that increase visibility and detection distances in areas where moose are common.