Biodiversity in forests depends on long canopy continuity and existence of different elements which function as substrates for varying organisms. Bryophytes often occupy specific substrates with a patchy distribution. The aim of this thesis was to increase the ecological knowledge about two threatened moss species; Buxbaumia viridis, inhabiting decaying wood, and Neckera pennata, inhabiting bark of base-rich deciduous trees. Establishment from spores was investigated in Buxbaumia viridis and Neckera pennata and models were created to predict germination of spores as a function of pH and water potential. The effects of pH, phosphorus and nitrogen concentration were studied in Buxbaumia viridis, both on spore germination and on sporophyte occurrences in the field. Colony growth in relation to precipitation and microhabitat variables was studied in Neckera pennata, and a model was used to predict growth of colonies over time. Metapopulation dynamics of Buxbaumia viridis were analyzed as an effect of precipitation, habitat quality and patch quantity. A spatial explicit patch occupancy model was constructed to simulate metapopulation sizes and extinction risk over 100 years. The quality of the substrate was very important for spore establishment. Germination success increased with increasing pH in both species. Buxbaumia viridis was less sensitive to low pH than Neckera pennata when water was freely available. However, there was a strong interaction between pH and water potential in prediction of the final cumulative germination: the spores reacted positively to one factor only when the other factor was in a favourable range. Precipitation, moisture holding capacity and interference competition were the main factors affecting colony growth of Neckera pennata. Buxbaumia viridis showed large fluctuations in number of occupied patches among years. Both colonizations and extinctions were highly related to precipitation. Spore germination and sporophyte occurrences in the field were positively related to phosphorus concentration and pH of the substrate.

Le feu dynamise la forêt boréale. De part son long cycle de feu, la forêt boréale de l'est du Québec présente une proportion élevée de vieilles forêts. Les perturbations secondaires y maintiennent une grande hétérogénéité dans l'habitat, permettant la présence d'une diversité d'espèces végétales. La coupe à blanc, qui a également eu lieu sur ce territoire, produit des impacts sur la végétation et le sol qui diffèrent de ceux du feu de forêt. Ceci peut conduire à une dissociation entre les successions végétales après coupe et après feu, en termes de diversité, de composition en espèces ou de variabilité dans les communautés végétales de sous-bois. Afin d'étudier ceci, nous avons comparé des chronoséquences après coupe et après feu. Notre dispositif se situe dans la Côte-Nord et comprend deux chronoséquences formées de 25 peuplements issus de feux et de 28 peuplements issus de coupes à blanc. La chronoséquence après feu comprend 5 classes d'âge, la dernière >200 ans, alors qu'après coupe, les classes sont 20-50 ans et 50-80 ans. Les variables environnementales ont été étudiées à l'intérieur d'une placette circulaire de 400 m2 et portent sur la structure et la composition du peuplement, le bois mort, le parterre forestier et le sol. Les plantes vasculaires et les bryophytes ont été inventoriées dans 20 quadrats de 1 m2. Des indices de biodiversité, des analyses de redondance (RDA) et des analyses de correspondance détendancées (DCA) ont été utilisés. Après un feu, une plus grande hétérogénéité de l'habitat s'observe en début de succession et après 150 ans, associée à une richesse plus élevée en plantes vasculaires et non-vasculaires. À la fermeture de la canopée, une diminution des espèces intolérantes à l'ombre se produit ainsi qu'une augmentation des mousses hypnacées au sol. Le parterre forestier présente alors moins de micro-habitats disponibles, ce qui explique la diminution de la richesse des bryophytes. Après 150 ans, les espèces plus sensibles et dépendantes des micro-habitats, telles les hépatiques, sont davantage riches et répandues. Ceci semble dû à la présence du sapin baumier et des perturbations secondaires. La variabilité de la composition des plantes vasculaires devient maximale en vieille forêt, ce qui peut s'expliquer par les divergences successionnelles des peuplements. De son côté, la coupe à blanc ne diminue pas la diversité des plantes vasculaires et non-vasculaires de sous-bois par rapport au feu pour les peuplements de même âge. Cependant, nous n'avons pas observé les mêmes changements successionnels pour les mêmes groupes d'espèces après coupe. La dominance des espèces résiduelles de forêt mature après coupe empêche la présence de changement chez les plantes vasculaires à la fermeture de la canopée. Toutefois, la richesse des hépatiques diminue dans le temps après coupe. Les limitations en quantité et en qualité du bois mort dans les peuplements coupés à blanc peuvent limiter les micro-habitats disponibles et constituer une atteinte au maintien des espèces sensibles sur ces territoires. ______________________________________________________________________________ MOTS-CLÉS DE L’AUTEUR : feu de forêt, coupe à blanc, forêt boréale, plantes de sous-bois, bryophytes, diversité, hétérogénéité de l'habitat

How long has the extant flora been present in the Antarctic? Glaciological reconstructions propose that most areas in Antarctica were covered by thick ice sheets throughout the Last Glacial Maximum (LGM; ~22-18 kya) as well as previous glaciations, suggesting terrestrial life must have been extremely limited during these periods. In contrast, recent biogeographic and genetic studies support most extant groups of Antarctic terrestrial fauna having survived past glaciations in situ. However, studies on the origin and age of the Antarctic flora remain sparse. Applying population genetic, phylogeographic and divergence time analyses I assessed the global biogeography, origin and age of several abundant Antarctic moss species, including: four Polytrichaceae mosses, characterised by having bipolar distributions, the most common (~45% of species) distribution pattern amongst Antarctic mosses; the globally widespread moss Ceratodon purpureus; the bank-forming moss Chorisodontium aciphyllum, also known for its old sub-fossils in Antarctica and long-term viability from revival experiments; and, lastly, the genus Schistidium, the most species-rich moss genus in Antarctica, including many endemic species. Genetic analyses revealed evidence of long-term (multi-million year) survival of plants in Antarctica (several species of Schistidium, Ceratodon purpureus, and possibly Polytrichum juniperinum). However, evidence for a likely more recent (< 100 ky) arrival of Chorisodontium aciphyllum was also found. Some species revealed multiple separate dispersal events to the Antarctic, suggesting the region may be less isolated for spore-dispersed organisms than previously thought. Evidence for increased genetic diversity in the northern maritime Antarctic compared to other regions point at it including potential refugial areas. Furthermore, genetic patterns revealed geographic features that enable and limit the connectivity of bryophytes globally as well as in Antarctica. This study suggests that, despite the harsh polar climate during glaciation periods, many bryophytes have had a much longer presence in Antarctica than previously thought.

In the 1850s when the timber floating began in Sweden many streams were channelized, which meant that the streams were straightened out, side channels were closed, and stones, trees and boulders were moved out to the banks of the streams. These measures altered the morphology of the streams, causing impediments to water moving in from the stream to the riparian zone, and reduced flooding frequency. When the era of timber floating was over humans began to restore the channelized streams to restore the damaged systems, mainly to improve fish habitats. Channelization and restoration involve major disruptions to the stream ecosystem, and the purpose of this study was to investigate how the bryophyte communities in the riparian zone differed in diversity, species richness and areal coverage, depending on the form of disturbance they were exposed to and the time passed since the disturbance occurred. This study was conducted along tributaries to the Vindel River in northern Sweden. There were three types of tributaries: partly channelized streams, best-practice restored streams and demonstration restored streams. At all tributaries included in the study, bryophyte samples were collected at two heights from the water surface, 0 and 40 cm. These samples were then identified to family or species level and their areal coverage was estimated. The results show that the demonstration restored streams had the lowest species richness and, there were no significant differences in diversity and species richness between the channelized streams and the best-practice restored streams. Regarding areal coverage, there was no significant difference between the stream types. The results also show that the time plays an important role for recovery of the bryophyte communities since diversity and species richness increased with the time since the disturbance occurred. / Under 1850-talet då flottning av timmer påbörjades i Sverige startade kanaliseringen av många vattendrag, vilket innebar att vattendragen rätades ut, sidokanaler stängdes, och stenar, träd och stenblock flyttades ut till vattendragens kanter. Dessa åtgärder förändrade vattendragens morfologi, orsakade fördämningar mellan vatten och strandzon och minskade översvämningsfrekvensen. När timmerflottningens era var slut började människan restaurera de kanaliserade vattendragen, främst för att det visat sig att fisket försämrats. Kanalisering och restaurering innebär stora störningar för ekosystemet de utförs i och denna studie syftar till att titta hur mossamhällen i strandzonen skiljer sig åt i diversitet, artrikedom och täckning beroende på vilken form av störning de utsatts för och tiden sedan de utsattes för störningen. Studien är gjord längs biflöden till Vindelälven i norra Sverige. Biflödena är dels kanaliserade vattendrag, bästa-praxis restaurerade vattendrag och demonstrationsåtgärdade vattendrag. Vid samtliga sträckor som ingår i studien har mossprover samlats in på två höjder från vattenytan, 0 och 40 centimeter. Dessa prover har sedan identifierats till lägsta möjliga taxonomiska nivå (art eller familj), och deras täckningsgrad har uppskattats. Resultaten visade att de demonstrationsåtgärdade vattendragen hade lägst artrikedom och diversitet och mellan kanaliserade vattendrag och bästa-praxis vattendrag kunde ingen signifikant skillnad i diversitet eller artrikedom påvisas.  Gällande täckningsgraden fanns ingen signifikant skillnad mellan vattendragstyperna. Resultatet visade också att tiden spelar en viktig roll för återhämtningen hos mossamhällena då artrikedom och diversitet ökar med tiden sedan inträffad störning.

<p>The main objectives of this thesis are to study if a topographic wetness index (TWI) could be used as a tool for predicting the spatial distribution of vascular plant species richness in the boreal forest as well as to study congruence in species richness between vascular plants, liverworts, mosses and lichens. A wetness index ln(a/tanβ) based on topography was used to assign a specific TWI-value to every 20 x 20m grid in two 25 km2 boreal forest landscapes (differing in average soil pH) in northern Sweden. Soil pH is known to be influenced by groundwater and to affect plant species richness in other biomes. Therefore, the relationships between plant species richness, TWI and soil pH were also studied. </p><p>The results showed that the majority of the investigated boreal forest landscapes were relatively dry and species-poor, whereas interspersed patches linked to areas with relatively high TWI had species-rich vegetation including the species of the drier parts of the landscape. There was a positive relationship between species richness of vascular plants and the TWI in both landscapes, but varied with average soil pH. TWI explained 30 % and 52 % of the variation in plant species richness in the landscape with lower and higher pH, respectively. The proportion of regionally uncommon plants also increased with TWI. Testing different calculation methods of the TWI resulted in a large variation in correlation strengths between the various TWI-values and different measured variables (species richness of vascular plants, soil pH, groundwater flow and soil moisture). The relationship between plant species richness and TWI could be further improved with some of the calculation methods. </p><p>When studying correlations in species richness using data sets from boreal forest in northern Sweden, strong positive correlations among vascular plants, mosses and liverworts were found, but no significant correlation between macrolichens and any of the other groups. This result could be explained by that the species number of each of the three related groups increases with ambient moisture, whereas the species number of macrolichens is weakly associated with moisture. </p><p>In conclusion, the TWI could become a useful tool in conservation management for identifying areas of special interest prior to field inventories. Since vascular plants can be used as an indicator taxon for species richness of mosses and liverworts, high TWI-values indicate areas of species hotspots of these taxa.</p> / <p>Syftet med avhandlingen är att dels studera om ett topografiskt fuktighetsindex skulle kunna vara användbart för att förutsäga fördelningen av kärlväxters artrikedom i boreal skog, dels att studera om den rumsliga fördelningen av artrikedom hos kärlväxter, blad- och levermossor samt lavar sammanfaller.</p><p>Ett fuktighetsindex, ln(a/tanβ), som bara är baserat på topografi användes för att beräkna ett indexvärde för varje 20 x 20 m grid i två 25 km2 stora boreala skogslandskap (med i medeltal olika mark-pH) i norra Sverige. Det är känt att mark-pH påverkas av grundvatten och att pH i sin tur påverkar artrikedom hos kärlväxter i andra biom. Därför studerades även sambanden mellan kärlväxters artrikedom, mark pH och TWI.</p><p>Resultaten visade att större delen av det studerade boreala landskapet var relativt torrt och artfattigt, medan mindre utspridda områden med höga TWI-värden var artrika på kärlväxter och här växte även arter som återfanns i de torra delarna av skogen. Sambandet mellan artrikedom hos kärlväxter och TWI var positivt i båda landskapen, men påverkades av de olika nivåerna på mark-pH. TWI förklarade 30 % av variationen i artrikedom i området med lägre mark-pH respektive 50 % i området med högre mark-pH. Andelen kärlväxter som klassificeras som icke vanliga i respektive region ökade också med TWI. Med andra beräkningsmetoder för TWI visade det sig att styrkan på korrelationerna mellan TWI och olika uppmätta variabler (artrikedom hos kärlväxter, mark-pH, grundvattennivå och markfuktighet) varierade mycket. Sambandet mellan artrikedom hos kärlväxter och TWI kunde förbättras ytterligare med vissa beräkningsmetoder.</p><p>Då korrelationer i artrikedom studerades användes ett dataset från boreal skog i norra Sverige. Resultaten visade på starka, positiva korrelationer mellan kärlväxter, blad- och levermossor, men inga korrelationer mellan någon av dessa grupper och lavar. Detta kunde förklaras med att artrikedom hos de tre korrelerande organismgrupperna ökar med ökad fuktighet, medan artrikedom hos lavar inte är kopplat till fukt.</p><p>Huvudslutsatsen i avhandlingen är att TWI, som endast är baserad på topograpfiskt data, skulle kunna bli ett värdefullt redskap i naturvårdsplanering för att identifiera särskilt intressanta skogsområden innan man gör fältinventeringar. Eftersom studien visar att kärlväxter kan användas som en indikator grupp för artrikedom hos blad- och levermossor indikerar höga TWI-värden områden med hög artrikedom även vad gäller dessa taxa.</p>

<p>This thesis examines the relative importance of local conditions, dispersal and dynamics of the trees on epiphyte distribution patterns and colonization-extinction dynamics. Study species are the mosses <i>Orthotrichum speciosum </i>and <i>O. obtusifolium</i>, and the red-listed <i>Neckera pennata</i>. The thesis also includes an attempt to parameterize a model for a lichen metapopulation (<i>Lobaria pulmonaria</i>) in a dynamic landscape, based on only presence/absence data of the epiphyte and its host trees. </p><p>The results show that epiphyte colonization of trees is affected by both local conditions, and by connectivity to occupied trees. The positive effect of connectivity, implying a restricted dispersal range, was established by both demographic and genetic studies. The important local conditions were tree diameter and vitality, and shade. Local extinctions from trees occurred among small trees with low local epiphyte abundance, but more often, were the results of tree fall.</p><p>The observed importance of connectivity on epiphyte colonization agrees with the assumptions of the classic metapopulation model. However, the classic metapopulation model assumes that the landscape is static, and that local extinctions occur for stochastic reasons. The dynamics of epiphytes are different. A new conceptual model is therefore suggested, the patch-tracking metapopulation model. It differs from the classic metapopulation model in that it includes dynamics of the patches, and in that local extinctions only occur as patches are destroyed.</p><p>Simulations of the dynamics of <i>N. pennata </i>showed that its future metapopulation size will be overestimated unless the dynamics of the trees are accounted for. The simulation results further suggest that the dynamics of <i>N. pennata </i>can be characterised by the patch-tracking metapopulation model. </p><p>The attempt to parameterize the <i>L. pulmonaria </i>metapopulation model showed that more information are required for rigorous parameterization, preferably of the past historic fire regime.</p>

This thesis examines the relative importance of local conditions, dispersal and dynamics of the trees on epiphyte distribution patterns and colonization-extinction dynamics. Study species are the mosses Orthotrichum speciosum and O. obtusifolium, and the red-listed Neckera pennata. The thesis also includes an attempt to parameterize a model for a lichen metapopulation (Lobaria pulmonaria) in a dynamic landscape, based on only presence/absence data of the epiphyte and its host trees. The results show that epiphyte colonization of trees is affected by both local conditions, and by connectivity to occupied trees. The positive effect of connectivity, implying a restricted dispersal range, was established by both demographic and genetic studies. The important local conditions were tree diameter and vitality, and shade. Local extinctions from trees occurred among small trees with low local epiphyte abundance, but more often, were the results of tree fall. The observed importance of connectivity on epiphyte colonization agrees with the assumptions of the classic metapopulation model. However, the classic metapopulation model assumes that the landscape is static, and that local extinctions occur for stochastic reasons. The dynamics of epiphytes are different. A new conceptual model is therefore suggested, the patch-tracking metapopulation model. It differs from the classic metapopulation model in that it includes dynamics of the patches, and in that local extinctions only occur as patches are destroyed. Simulations of the dynamics of N. pennata showed that its future metapopulation size will be overestimated unless the dynamics of the trees are accounted for. The simulation results further suggest that the dynamics of N. pennata can be characterised by the patch-tracking metapopulation model. The attempt to parameterize the L. pulmonaria metapopulation model showed that more information are required for rigorous parameterization, preferably of the past historic fire regime.

The main objectives of this thesis are to study if a topographic wetness index (TWI) could be used as a tool for predicting the spatial distribution of vascular plant species richness in the boreal forest as well as to study congruence in species richness between vascular plants, liverworts, mosses and lichens. A wetness index ln(a/tanβ) based on topography was used to assign a specific TWI-value to every 20 x 20m grid in two 25 km2 boreal forest landscapes (differing in average soil pH) in northern Sweden. Soil pH is known to be influenced by groundwater and to affect plant species richness in other biomes. Therefore, the relationships between plant species richness, TWI and soil pH were also studied. The results showed that the majority of the investigated boreal forest landscapes were relatively dry and species-poor, whereas interspersed patches linked to areas with relatively high TWI had species-rich vegetation including the species of the drier parts of the landscape. There was a positive relationship between species richness of vascular plants and the TWI in both landscapes, but varied with average soil pH. TWI explained 30 % and 52 % of the variation in plant species richness in the landscape with lower and higher pH, respectively. The proportion of regionally uncommon plants also increased with TWI. Testing different calculation methods of the TWI resulted in a large variation in correlation strengths between the various TWI-values and different measured variables (species richness of vascular plants, soil pH, groundwater flow and soil moisture). The relationship between plant species richness and TWI could be further improved with some of the calculation methods. When studying correlations in species richness using data sets from boreal forest in northern Sweden, strong positive correlations among vascular plants, mosses and liverworts were found, but no significant correlation between macrolichens and any of the other groups. This result could be explained by that the species number of each of the three related groups increases with ambient moisture, whereas the species number of macrolichens is weakly associated with moisture. In conclusion, the TWI could become a useful tool in conservation management for identifying areas of special interest prior to field inventories. Since vascular plants can be used as an indicator taxon for species richness of mosses and liverworts, high TWI-values indicate areas of species hotspots of these taxa. / Syftet med avhandlingen är att dels studera om ett topografiskt fuktighetsindex skulle kunna vara användbart för att förutsäga fördelningen av kärlväxters artrikedom i boreal skog, dels att studera om den rumsliga fördelningen av artrikedom hos kärlväxter, blad- och levermossor samt lavar sammanfaller. Ett fuktighetsindex, ln(a/tanβ), som bara är baserat på topografi användes för att beräkna ett indexvärde för varje 20 x 20 m grid i två 25 km2 stora boreala skogslandskap (med i medeltal olika mark-pH) i norra Sverige. Det är känt att mark-pH påverkas av grundvatten och att pH i sin tur påverkar artrikedom hos kärlväxter i andra biom. Därför studerades även sambanden mellan kärlväxters artrikedom, mark pH och TWI. Resultaten visade att större delen av det studerade boreala landskapet var relativt torrt och artfattigt, medan mindre utspridda områden med höga TWI-värden var artrika på kärlväxter och här växte även arter som återfanns i de torra delarna av skogen. Sambandet mellan artrikedom hos kärlväxter och TWI var positivt i båda landskapen, men påverkades av de olika nivåerna på mark-pH. TWI förklarade 30 % av variationen i artrikedom i området med lägre mark-pH respektive 50 % i området med högre mark-pH. Andelen kärlväxter som klassificeras som icke vanliga i respektive region ökade också med TWI. Med andra beräkningsmetoder för TWI visade det sig att styrkan på korrelationerna mellan TWI och olika uppmätta variabler (artrikedom hos kärlväxter, mark-pH, grundvattennivå och markfuktighet) varierade mycket. Sambandet mellan artrikedom hos kärlväxter och TWI kunde förbättras ytterligare med vissa beräkningsmetoder. Då korrelationer i artrikedom studerades användes ett dataset från boreal skog i norra Sverige. Resultaten visade på starka, positiva korrelationer mellan kärlväxter, blad- och levermossor, men inga korrelationer mellan någon av dessa grupper och lavar. Detta kunde förklaras med att artrikedom hos de tre korrelerande organismgrupperna ökar med ökad fuktighet, medan artrikedom hos lavar inte är kopplat till fukt. Huvudslutsatsen i avhandlingen är att TWI, som endast är baserad på topograpfiskt data, skulle kunna bli ett värdefullt redskap i naturvårdsplanering för att identifiera särskilt intressanta skogsområden innan man gör fältinventeringar. Eftersom studien visar att kärlväxter kan användas som en indikator grupp för artrikedom hos blad- och levermossor indikerar höga TWI-värden områden med hög artrikedom även vad gäller dessa taxa.

Why evaluations of the ecological outcomes of stream and river restoration have largely reported inconclusive or negative results has been the subject of much debate over the last decade or more. Understanding the reasons behind the lack of positive results is important for bettering future restoration efforts and setting realistic expectations for restoration outcomes. This thesis explores possible explanations for why researchers have failed to find clear and predictable biotic responses to stream restoration: recovery time has been too short, that restoration of habitat complexity is not clearly linked to instream biodiversity, that one monitored organism group is not representative of the entire community, that restoration effort was not intense enough to restore the potential habitat complexity of a system, and that reach-scale restoration done in the presence of catchment-scale degradation obscures restoration results. The overarching goal of this thesis is to study the holistic effect of reach-scale restoration of historic reach-scale simplification, due to timber floating in northern Swedish streams, thus avoiding the added pressure of catchment-scale degradation typically found at most restoration sites (e.g., non-point-source pollution and impervious cover). Using this model system, I was able to show that it took 25 years for riparian plant species richness at restored sites to increase above that of channelized sites. Furthermore, it was clear that restoration of these streams caused a large and rapid change in N-processing in the riparian zone and this alteration persists for at least 25 years. Additionally, multiple metrics of geomorphic complexity were needed to explain some of the more subtle responses of organism groups. Macroinvertebrates, diatoms, and macrophytes did not respond concordantly and cannot serve as surrogates or indicators for each other. I found that older best practice methods of restoration rarely restored the large-scale features needed to bring the sites up to their potential complexity because these elements were destroyed or removed from the system. Advanced restoration techniques used in more recent restorations added big boulders and instream wood and increased complexity to a level that elicited a biological response. By combining surveys of multiple metrics of structure, diversity of multiple organism groups, and process in this thesis I was able to get a holistic view of the effects of restoration of streams after timber floating. We now know that it takes at least 25 years for riparian plants and N-cycling to recover, we understand that multiple metrics of geomorphic complexity should be measured to be able to explain biotic responses, and that restored complexity should better match the potential complexity of the site in order to elicit a biological response. Finally, we know that multiple organism groups need to be assessed when evaluating the response of biodiversity to restoration.

In recent time, forests in Sweden have lost habitat quality in the loss of deadwood due to the practices in productions forestry. This means that substrates for epixylic species has decreased both in quality and quantity. The aim with this study was to find which preferences the epixylic liverwort, Crossocalyx hellerianus has on its substrate and environment. C. hellerianus requires coarse logs with rotten wood, which usually is more available in forests with long forest continuity. This study was performed to see if forest continuity, isolation of habitats and if the substrate was of significance for a demanding species such as C. hellerianus. The study was performed in six sites and in each site several 50x10 meter transects were surveyed for logs. Each log was measured for diameter, sun exposure, ground contact, stage of decay and if the log was colonized by C. hellerianus. The result showed that C. hellerianus preferred coarser logs and that the species only appeared on an intermediate or later stage of decay. A high forest continuity and low habitat isolation turned out to be positive factors for the abundance of C. hellerianus