Diss. Uppsala : Uppsala universitet, 1746. / Antikva. Över titeln: "I. N. J." Se: Lidén, Catalogus disputationum I, s. 310.

Diss. Uppsala : Uppsala universitet, 1746. / Över titeln: "I. N. J." Se: Lidén, Catalogus disputationum I, s. 310.

The genetic diversity in modern species is strongly affected by contemporary gene flow between populations, which in turn is governed by individual dispersal capacities and barriers in the landscape. However, current patterns of variation have also been shaped by movement over longer time-scales, such as the successive shifts in species distributions that have occurred during past climate changes. This thesis is focused on cold-adapted species, and one parameter that has greatly influenced their current genetic diversity is how they coped with climate warming at the last glacial/interglacial transition, ca 11.7 thousand years ago. I examined this in three different small herbivore taxa; true lemmings (Lemmus), ptarmigan (Lagopus) and hares (Lepus), whose modern distributions stretch from the exposed tundra to the subarctic moorlands and taiga. In the first paper, I investigated contemporary genetic structure in the cyclic Norwegian lemming (Lemmus lemmus) and proposed that mass movements during peak years act as pulses of gene flow between mountain areas, which homogenise the gene pool over surprisingly vast geographic distances. However, when I used ancient DNA to analyse the lemmings’ ability for long-term directional movement, I found that the Ice Age populations that inhabited the former midlatitude European tundra-steppe appear to have been incapable of shifting their distribution northwards following post-glacial climate warming. Instead, the results suggest that the endemic Norwegian lemming descends from an isolated population that survived the last glacial maximum in situ in a restricted ice free refugium. In contrast to the glacial lemmings, as well the majority of previously studied mammals, the ptarmigan (L. lagopus and L. muta) and hare (L. timidus) analyses revealed a long-term genetic continuity in Europe, where the midlatitude populations were able to keep pace with the rapidly changing climate at the last glacial/interglacial transition, enabling them to shift their ranges to northern and high-alpine regions. These different outcomes might be explained by ptarmigans’ flight capability that allows a less restricted dispersal across fragmented landscapes, and that the generalist nature of mountain hares makes them less vulnerable to habitat alterations. Species distribution modelling, however, indicated that continued climate warming will make some isolated regions unsuitable in the future, thereby forcing populations to adapt the new environmental conditions in order to avoid local extinctions. / <p>At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 1: Manuscript. Paper 3: Manuscript. Paper 4: Manuscript.</p>

Anthropogenic changes in climate and land use are causing a dramatic erosion of biodiversity. To understand this erosion, and predict future transformations of biodiversity, we need to understand better species’ response to these changes at different spatial and temporal scales. Modeling studies have identified correlations between physical parameters of the environment and species’ distribution at large spatial scales. However, this does not accurately characterize the response of a specific species, since this does not account for the constraints arising from the biology of the species. This thesis shall combine knowledge on the biology of species obtained from laboratory experiments with modeling studies. This will allow us (i) to identify life history traits and biotic interactions that influence species’ adaptive potential, and hence, explain possible differences in species’ distribution, and (ii) to consider, not only the ecological but also the evolutionary aspects of species’ response to changes. This integrative approach is likely to improve our predictions on species’ population dynamic in a changing environment. I focus on a community of butterflies in Sweden (Vanessa cardui, Polygonia c-album, Aglais urticae, Aglais io, Araschnia levana) that feeds on the stinging nettle (Urtica dioica). The available knowledge on the biology of these species and their short life cycles, which allow investigations of their response to changes on a short-time scale, make them a good system to study. Among three of these species, I showed great differences in organisms’ response to variation in food nutrient content. This is a potentially important finding considering the increased use of chemical fertilizers. These differences are to a large extent explained by differences among species in their degree of host plant specialization and voltinism (paper II). Thus, life history traits determine the response of species to environmental changes, but are themselves likely to evolve in response to such changes. Climate change, for instance, may alter the phenological synchrony between plant-feeding insects and their host plants, making it necessary for the insects to evolve their host plant range in order to ensure the availability of resources during larval development (paper I &amp; III). The biology of a species, including biotic interactions, helps to explain the observed shift in a species’ distribution and environmental niche that result from climate change. I have shown that the recent establishment of A. levana in southern Sweden has modified the niche of the resident species, A. urticae and A. io (Paper IV). Niche partitioning in this community is likely mediated by parasite-driven apparent competition. / Mänsklig påverkan på klimat och markanvändning har orsakat en dramatisk förlust av biologisk mångfald. Effekten av dessa förändringar på lokal och regional nivå är dock komplex, och kräver integrativa strategier för att kunna förstå och förutsäga förändringar, på individ-, art- och samhällsnivå. Experimentella studier har utforskat arters plastiska och evolutionära respons till främst abiotiska förändringar, och observationsdata har använts för att modellera skiften i fenologi och utbredning som en konsekvens av klimatförändringar. Trots detta är det fortfarande mycket kvar att förstå för att kunna förutsäga hur miljöförändringar ska påverka arters respons på olika rumsliga och tidsliga skalor. Denna avhandling undersöker i vilken utsträckning arters specifika livshistoria och artinteraktioner kan förklara deras ekologiska och evolutionära respons på miljöförändringar. För att angripa detta har jag fokuserat på ett samhälle av fjärilar i Sverige (Vanessa cardui, Polygonia c-album, Aglais urticae, Aglais io, Araschnia levana) som alla lever på brännässla (Urtica dioica). Den tillgängliga kunskapen om dessa arters biologi samt deras korta livscykler gör det möjligt att undersöka deras svar på förändringar över korta tidsskalor, vilket gör dem till ett lämpligt studiesystem. Huvudslutsatsen från denna avhandling är att för att beskriva hur en art svarar på en förändring måste man ta hänsyn till variation i livshistorieegenskaper och artinteraktioner. Till exempel har den ökade användningen av kemiska gödningsmedel förändrat näringstillgången även i naturliga ekosystem, vilket gynnar växtarter som är kapabla att växa under höga näringsnivåer, som brännässla. Variation i växternas näringsinnehåll kommer i sin tur att påverka herbivorerna som äter av dem, och artikel II visar att skillnader mellan fjärilsarter i hur de svarar på variation i näringstillgång till stor del beror på specialiseringsgrad och voltinism (antal generationer per år). Livshistorieegenskaper avgör således hur arter kommer att svara på förändringar i klimat och markanvändning, men sådana miljöförändringar påverkar i sin tur också evolution av livshistorieegenskaperna (artikel I &amp; III). Slutligen, förändringar i utbredning som ett resultat av klimatförändring kommer även att påverka den lokala sammansättningen av interagerande arter (resurser, predatorer, konkurrenter). Ett exempel på detta är hur den relativt nyliga koloniseringen av södra Sverige av A. levana har förändrat nischerna hos de inhemska arterna A. urticae och A. io (artikel IV). / Le réchauffement climatique et les changements d’occupation des terres d'origine anthropique provoquent une forte érosion de la biodiversité. Pour comprendre cette érosion, et prédire les transformations futures de la biodiversité, il nous faut mieux connaitre la réponse des espèces à ces changements, aux différentes échelles spatiales et temporelles. Grâce à des outils de modélisation statistique, des corrélations entre les paramètres physiques de l’environnement et la distribution des espèces à grande échelle spatiale ont été observées. Mais ceci ne suffit pas à caractériser finement la réponse d’une espèce donnée, car celle-ci dépend des caractéristiques biologiques propres de l’espèce. Cette thèse se propose donc d’associer les connaissances sur la biologie des espèces obtenues par des expériences en laboratoire à des études de modélisation. Ceci permettra (i) d’identifier des traits d’histoire de vie et les relations biotiques qui influencent le potentiel adaptatif des espèces, et donc expliquent d’éventuelles différences de répartition, et (ii) d’envisager, au-delà des aspects écologiques, la composante évolutive de cette réponse. Une telle approche intégrative est susceptible d’améliorer nos prédictions sur la dynamique des espèces dans un environnement changeant. Le système d’étude de cette thèse est une communauté de papillons en Suède (Vanessa cardui, Polygonia c-album, Aglais urticae, Aglais io, Araschnia levana), se nourrissant de l'ortie (Urtica dioica). La biologie de ces espèces est bien connue et leur cycle de vie rapide permet d’étudier leur réponse aux changements à une échelle de temps court. Chez trois de ces espèces, j’ai mis en évidence des réponses très différentes à une augmentation de la teneur en nutriments de leur nourriture, conséquence attendue de l’utilisation accrue d’engrais chimiques. Ces différences sont dans une large mesure expliquée par la gamme de plantes hôtes utilisées et le voltinisme (article II). Ces traits d'histoire de vie déterminent donc la réponse des espèces aux changements, mais sont eux-mêmes susceptibles d’évoluer. Par exemple, le réchauffement climatique modifie la synchronie entre les insectes herbivores et leurs plantes hôtes et pousse à l’évolution du régime alimentaire des papillons afin que la présence de ressources soit assurée durant le développement des larves (articles I &amp; III). Les connaissances sur la biologie des espèces, y compris leurs interactions biotiques, permettent de comprendre les variations de leur distribution spatiale et de leur niche environnementale. Ainsi, l’établissement récent d’A. levana dans le sud de la Suède, suite au réchauffement, a modifié les niches des espèces résidentes, A. urticae et A. io (article IV). / <p>At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 1: Manuscript. Paper 4: Manuscript.</p> / Ekoklim

Majoriteten av Sveriges vattendrag observeras i skogsmark, där en god skogsbruksplanering kan förbättra vattenkvaliteten. Världsnaturfonden (WWF) konstruerade således verktygen Naturvärden, Påverkan, Känslighet och Plusvärde (NPK+) och Blå målklassning, vilka har utvecklats av skogsstyrelsen. NPK+ innefattar en visuell naturvärdesinventering som poängsätter vattendrags befintliga egenskaper enligt ett fältprotokoll. Blå målklassning baseras på NPK+-poängsumman och beskriver ambitionsnivån för vattendragens hänsyn. År 2005 erhöll Kolarebäcken ett av Västra Götalands län största bestånd av den rödlistade flodpärlmusslan (M. margaritifera), men i dagsläget påvisas en markant minskning av arten. Studien ämnar undersöka om Blå målklassning, Känslighet och NPK+-protokollets underkategorier kan förklara var flodpärlmussla förekommer och var höga eller låga tätheter finns i ett vattendrag, samt huruvida variabler inom signifikanta kategorier eller underkategorier i NPK+ gynnar flodpärlmusslans förekomst och täthet. Kolarebäcken inventerades enligt skogsstyrelsens manual av Blå målklassning. Baserat på skogens och vattnets karaktär konstruerades segment med en NPK+-poängsumma och Blå målklass. Flodpärlmusslans förekomst och täthet hämtades från länsstyrelsens inventeringsstationer år 2011 och 2017. Varken Pearson’s Chi-squared test, Fisher’s exact test eller ANOVA påvisar att Blå målklassning kan förklara var förekomst eller höga, alternativt låga, tätheter finns. Oparade t-test och Welch t-test påvisar att flodpärlmusslan främst förekommer i segment som erhåller höga naturvärden i vattendraget och i kantzonen, i synnerhet död ved som genereras av en äldre kantzon. Multipel linjär regressionsanalys påvisar att högre tätheter främst förekommer i segment med höga naturvärden och känslighet samt låg påverkan, i synnerhet en äldre, blöt kantzon. Fler vattendrag behöver dock undersökas för bekräftelse av att Kolarebäckens samband representerar övriga vattendrag. / Streams in Sweden mainly occurs in woodland, where a decent forest management plan can improve the water quality. Thus, the World Wide Fund for Nature (WWF) constructed Conservation value, Impact, Sensitivity and Added values (CISA) and Blue targeting, which were further developed by the Swedish Forest Agency. CISA involves an inventory, which marks visual variables within streams according to a check list. Blue targeting is based on the CISA credits and describes the consideration ambition level. In 2005, Kolarebäcken contained one of the largest populations of the red-listed freshwater pearl mussel (M. margaritifera) in Västra Götaland County. However, a significant reduction of the species is currently detected. The study aims to investigate whether Blue targeting, Sensitivity and the subcategories of CISA checklist might explain where freshwater pearl mussels occur and where high or low densities are found in a stream, plus whether variables within significant categories or subcategories in CISA favor the occurrence and density. Kolarebäcken was inventoried according to the Swedish Forest Agency’s manual. Stream sections, containing one CISA credit and Blue target, were constructed based on fluctuations in forest or water. The mussels’ occurrence and density were obtained from the County Administrative Board’s inventory locations in 2011 and 2017. Neither Pearson's Chi-squared test, Fisher's exact test nor ANOVA indicates that Blue targeting can explain where occurrence or high, alternatively low, densities are found. Unpaired t-tests and Welch t-tests indicates that the occurrence is mainly affected by high conservation values in the stream and riparian zone, especially occurrence of dead wood generated by an old riparian zone. Multiple linear regression analysis indicates that high densities mainly occurs within segments including high conservation values, sensitivity and low impact, especially an old, wet riparian zone. Investigations of several streams are required to confirm that the association in Kolarebäcken represents other streams.

It is known that artificial light at night has a role in disrupting many birds’ routines when it comes to timing of dawn song. However, studies show various results on which species that are affected, how much and on what latitude. Birds at northern latitudes is experiencing longer and brighter natural nights during the breeding season as the spring is progressing, compared to birds living in areas more south. The bright nights might affect how the birds respond to the artificial light at night. This study's aim was to investigate how different species of songbirds in Linköping (58°N), Sweden, time their dawn song during breeding season, depending on presence or absence of artificial light at night coming from street lamps. The results show that early singing species are more affected than late singing species. The common blackbird for example, advanced its song because of artificial light, up to over an hour on average. Late singing birds was also affected by the urban lights, though not as strongly as the early singing species. Also shown was that the temperature and the amount of cloudiness, did not affect the difference in timing of dawn song between the locations. Future studies should focus on collecting data from more northern latitudes. The effect the advanced timing of dawn song, both regarding the birds on an individual scale and also regarding the ecosystems in total,should also be investigated further.

Due to increasing habitat loss, human-lion conflict, poaching and other reasons, African lion (Panthera leo) populations have suffered a drastic decline. The African Lion and Environmental Research Trust (ALERT) is working to stop this pattern and is the first organization with an ex-situ conservation project for lions. Before releasing lions raised by captive-bred adults, they must first be ensured to behave properly to make sure they have the highest chance of survival. One challenge in the wild is encountering and competition with unknown conspecifics. By conducting playback of unfamiliar lion roars, the behaviours of lions under this ex-situ reintroduction program were tested and compared with observations from earlier studies of wild lions. Social interactions were also collected and a social network analysis was done to give information about the social structure in the pride. This in turn was compared with boldness scores, calculated from behavioural responses in the playback experiments. Lastly, I searched for associations between age and sex with both boldness and social interactions.   The studied pride consisted of 12 lions. The lions were more vigilant when a playback consisted of numerous lions vocalizing, but playing more than three lions seemed to make them loose interest, suggesting either habituation or false information. One adult female and the alpha-male were most bold, followed by five sub-adults. Boldness did not vary according to sex or age differences, but the social network analysis showed that some social interactions were more dominated by one sex or age group. These behaviours were in agreement with comparisons of wild prides.   This study showed that captive-bred lions have developed natural social behaviours. Based on the behavioural responses observed by the captive-origin lions to the playbacks of unfamiliar lions, it is unclear whether these lions would appropriately respond when encountered with unfamiliar conspecifics in the wild post-release.

Osämja mellan människan och rovdjuren har funnits sedan vi slutade jaga och började bruka jorden och bedriva lantbruk. En av de absolut viktigaste faktorerna som bestämmer vart en art förekommer, hur stark en stam är och hur den förändras över tiden, är predation. I svensk historia har både rådjurs (Capreolus capreolus) och rödräven (Vulpes vulpes) båda populationer varit nära nationell utrotning. Syftet med uppsatsen är att ta reda på om räven har någon påverkan på rådjurspopulationen. Detta görs genom att analysera avskjutningsstatistik från viltdata.nu i tre olika län med liknande klimat och innehåller samma arter av predatorer. Analysen gjordes med korrelationsanalys. Dock var det inget utav resultaten som visade någon signifikans. Dessa olika resultat i de olika län tyder på att det finns andra faktorer än räven som påverkar rådjurspopulationen. / A conflict between humans and predators has existed since man stopped hunting and started with agriculture instead. We know today that one of the most important factors influencing the distribution and abundance of a species is predation. Over the last two? centuries, both the roe deer (Capreolus capreolus) population and the red fox (Vulpes vulpes) population have been very close to extinction in Sweden. The purpose of this study was to determine if the red fox population affects the roe deer population. This was done by analyzing hunting statistics from viltdata.nu in three different regions with similar climate and with the same type of carnivores. The analysis was performed using correlation analysis. None of the relationships between roe deer and the fox in the three different areas was significant. This indicates that there are factors other than predation by fox that influence the deer population.

Animals in captivity can be deprived of performing some of their natural behaviours. Using enrichments may allow them to express a larger part of species-specific behaviour repertoire and with a better frequency distribution. This study focuses on three species of the family Herpestidae which live in captivity at Bioparc Valencia (Spain). The project aims to study the effect of environmental enrichment on the behaviour of meerkats, banded mongooses and dwarf mongooses in human care. To achieve this goal two different types of enrichmentswere tested: (1) a food enrichment with several variations and (2) an olfactory enrichment with the presentation of two new odours. The food enrichment aimed to increase foraging behaviour and the olfactory enrichment aimed to test if captive animals behave differently in the presence of a predator’s odour compared to a non-predator’s odour. Results revealed that foraging can increase up to 16% implementing enrichments and that success depends on the presence and quantity of food. On the other hand, animals did not seem to behave differently in the presence of both odours. The frequencies of behaviours and time spent interacting did not differ between these olfactory enrichments. I conclude that implementing enrichment programmes may ensure better welfare for captive animals.

Parental care can be important especially in passerine birds and can depend on environment, food availability and behaviour. A behaviour that can affect the food supply for the offspring is neophobia, the fear of novel objects. The more fear the bird exhibits the less it visits the nest with food. In my study I aimed to examine neophobia and how it differed based on sex and if offspring number and weight influenced the behaviour. Video recordings of collared flycatchers nest boxes were used to see how the behaviour differed by placing a novel object on the nest box. The behaviours tested were duration of perching, hovering and visit including latency and frequency of visit. The results showed that there was a significant difference between the sexes in duration, latency, and frequency of visit but also duration of perching before and after placing a novel object on the nest box. In addition, my results showed that nestling weight significantly influenced frequency of visit and nestling number influenced duration of visit. The study shows that the collared flycatchers do express neophobia differently based on the sex and the nestling’s weight and number. Future studies could investigate if neophobia is a plastic behaviour and the effects of prolonged exposure to novel objects whichcould change the offspring’s survival chances or even the nestling’s behaviour.