Dissertations / Theses on the topic 'Bryophyte'

Includes bibliographical references (leaves 46-54).
Forests occur as fragmented patches throughout Africa, however the basis for the disjunction of afromontane forests remains uncertain. The genetic structure of organisms should reflect their hi story, and in turn the history of their environment. Thus a phylogeographical study of forest-faithful mosses could provide insight into the fragmentation of forests. In this study, patterns of genetic variation in four forest-faithful mosses (Leptodon smithii, Pyrrhobryum spiniforme, Aerobryopsis capensis and Neckera valentiniana) were investigated.

Spatial processes have a profound influence on the structure and function of community assemblages. The dispersal of organisms from their place of origin to the location in which they live out their reproductive life is particularly important for plant communities, which generally cannot adjust their location post-germination. Connection between communities at a landscape scale can also influence species persistence, local and regional diversity, and functional turnover at the metacommunity scale. Animals have been shown to disproportionately deposit propagules in particular microsites in many plant species, facilitating the arrival of plants to appropriate niche-space. Birds are particularly notable seed dispersers, given their ability to fly long distances and their behavioral inclination toward using specific microsites within their habitat for foraging and nest building. Despite the known influence of animal behavior on plant dispersal outcomes, little work has been done to investigate the role of animals in dispersing bryophyte (moss, hornwort, liverwort) propagules. In order to examine how birds may affect bryophyte dispersal, I conducted two studies focused on understanding how bird species identity and behavior influence the bryophyte propagules they carry. In addition, I conducted a study to understand how metacommunity structure across a landscape can be influenced by focal spatial scale. In the first study I examined how bird species and foraging behavior impact the topical load of bryophyte spores found on bird surfaces. In order to determine this, I captured passerine birds in mist nets and swabbed them for spores. I found that spores were more abundant on passerine tails than legs, and that overall spore load was higher on larger birds. Thrushes in particular carried more spores than other groups overall. Bark and foliage foraging birds had more spores on their tails than ground foraging birds. From these samples I was able to germinate 242 individual bryophytes, demonstrating that carried spores were readily viable. In the second study, I examined species-specific relationships between bryophytes and the birds carrying them. Swabs from captured birds were grown in the lab and bryophyte species were determined genetically. I used a bipartite network approach to determine the level of specialization of associations within the overall network, as well as how specialized the avian associations of individual bryophyte species were. I then used the phylogenetic distance of bryophytes found on individual bird species in order to assess how specialized the assemblages on a given bird species were compared with a null, random model. I found that bryophyte associations with birds were nonrandom, and that the extent to which those associations were specialized differed by bird foraging behavior. In addition, I found that the diversity of propagules on bird surfaces was significantly nonrandom, with the exception of those bryophytes found on Spotted Towhees. In the final study, I examined the metacommunity structure of bryophytes at both patch and landscape scales across a relict landscape of Valdivian forest in North-Central Chile. This landscape consists of distinct natural patches of forest maintained by coastal fog deposition, surrounding by a dry matrix inhospitable to patch-resident bryophytes. I used quadrats to sample bryophyte species abundance at the base and at breast height of ten trees in each patch, in 20 patches across the landscape. I found that when considering the whole park as one metacommunity, the bryophyte community exhibited a Gleasonian structure, in which individual species turnover was idiosyncratic. Considering assemblages from both heights separately, a Clemenstian pattern was observed, suggesting that within each height compartment, turnover of species tended to happen together. Treating each patch as a metacommunity of individual community trees resulted in a wide variety of metacommunity structures across the park that did not reflect either longitude or latitude. Low canopy cover and small DBH resulted in structures reflecting random species loss. Underlying Shannon diversity did not explain differences in the observed structures. This dissertation provides the first evidence that passerine birds carry bryophyte propagules, and that their individual species use of habitat and foraging behaviors are likely to influence the number and diversity of the bryophytes they are dispersing. This has implications for understanding disjunct species and genetic distributions observed in bryophytes that to date have lacked an explanatory mechanism for long distance directed dispersal. In addition, understanding how avian behavior may disperse propagules at a local to regional scale may provide better insight into the trajectory of bryophyte recruitment on impacted landscapes. I also found that assignation of metacommunity structure is sensitive to spatial scale in bryophytes. Together, these findings increase our understanding of the role that spatial processes play in forming bryophyte communities.

The presented thesis investigates the role of bryophytes in the deadwood and carbon (C) cycle of boreal black spruce forests in Labrador, Canada. All major forest C pools (live-tree, standing and downed deadwood, organic layer, mineral soil) were quantified for three old-growth, nine clearcut harvested, and three burned forest stands in order to characterize forest C dynamics of a high-latitude humid boreal forest ecosystem. Tree and aboveground deadwood C dynamics of Labrador black spruce forests were similar to those of drier or warmer boreal forests. However, due to bryophyte-driven processes such as woody debris (WD) burial and paludification, the studied forests contained high organic layer, mineral soil, and buried wood C stocks. The comprehensive field-measured data on C stocks was used to evaluate the CBM-CFS3, a Canadian national-scale C budget model, with respect to its applicability to Labrador black spruce and humid boreal forests elsewhere. After selected biomass estimation and deadwood decay parameters had been adjusted, the CBM-CFS3 represented measured live-tree and aboveground deadwood C dynamics well. The CBM-CFS3 was initially designed for well-drained upland forests and does not reflect processes associated with bryophytes and high forest floor moisture content, thus not capturing the large amounts of buried wood and mineral soil C observed in the studied forests. Suggestions are made for structural changes to the CBM-CFS3 and other forest ecosystem C models to more adequately represent the bryophyte-regulated accumulation of buried wood, organic layer, and mineral soil C. Accuracy of forest C models could be further improved by differentiating WD decomposition rates by disturbance history, because WD respiration reflects disturbance-induced changes in temperature and moisture regimes. In Labrador, WD respiration was limited by low WD moisture levels and high temperatures in burned stands, and by high WD moisture contents and low temperatures in old-growth stands. Following harvesting, residual vegetation prevents the desiccation of WD, resulting in significantly higher WD respiration compared to old-growth and burned stands. Moreover, the bryophyte layer recovers faster following harvest than following fire, which reduces WD desiccation due to moisture retention, water transfer, and moisture-induced cooling and results in higher WD decomposition rates. Bryophytes are thus a key driver of the deadwood and C cycle of humid boreal Labrador black spruce forests. The author recommends to classify these and similar boreal forests as a functional ecosystem group called “humid boreal forests”, preliminarily defined as “boreal forest ecosystems featuring a bryophyte-dominated ground vegetation layer associated with low soil temperatures, high moisture levels, low dead organic matter decomposition rates, and subsequently (in the absence of stand-replacing disturbances) an accumulation of buried wood embedded in a thick organic layer”. Bryophytes are also an integral component of many coniferous forests outside the boreal biome. Bryophyte-regulated processes such as WD burial or paludification are thus likely significant to the global C cycle. The potential climate change-induced release of large amounts of CO2 from buried wood and soil C pools necessitates an increased understanding of how bryophyte productivity and decomposition constraints will change with increasing temperature and varying moisture regimes. Ecosystems such as humid boreal forests with potentially high C losses to the atmosphere may thus be identified and counteractive forest management strategies can be developed and implemented
Cette thèse de doctorat s’intéresse à l’influence qu’exercent les mousses sur les cycles du bois mort et du carbone (C) dans des pessières noires boréales humides du Labrador, Canada. Toutes les réservoirs majeurs de C (arbres vivants, bois mort sur pied et effondré, l’horizon de matière organique, sol minéral) de trois pessières vierges, neuf coupes à blanc et de trois pessières brûlées ont été quantifiés pour caractériser le cycle du C des forêts humides boréales du nord. Les dynamismes de C des arbres vivants et du bois mort supraterrestre ressemblaient à ceux des forêts boréales plus sèches ou aux températures plus chaudes. À cause des processus régulés par les mousses (l’enterrement du bois mort ou la paludification), les forêts étudiées contenaient des stocks élevés de C au sein de l’horizon de matière organique, le sol minéral et le bois enterré. Les données ont aussi été utilisées pour évaluer le MBC-SFC3, un modèle national canadien du bilan du C, concernant son applicabilité aux pessières boréales humides de Labrador et d’ailleurs. Suite à l’ajustement de quelques paramètres, p.ex. des taux de décomposition, le MBC-SFC3 reproduisait bien le dynamisme mesuré des arbres vivants et du bois mort supraterrestre. Le MBC-SFC3 a initialement été développé pour les sites bien drainés et ne considère pas les processus associés avec les mousses ou l’humidité élevée du sol. Conséquemment, le MBC-SFC3 ne représentait pas les stocks élevés de C mesurés pour le bois enterré et pour le sol. Les modifications structurelles du MBC-SFC3 et d’autres modèles du C forestier sont nécessaires pour représenter adéquatement l’accumulation du C au sein de ces réservoirs. La précision des modèles du C forestier pourrait encore être améliorée par une différenciation des taux de décomposition selon le régime de perturbations, parce que la respiration du bois mort reflète les changements de la température et d’humidité associés avec une perturbation spécifique. Dans les pessières brûlés du Labrador, la respiration du bois mort était limitée par a faible humidité du bois et des températures élevées; dans les pessières vierges, par l’humidité élevée du bois et des températures basses. Dans les coupes à blanc, la végétation résiduelle empêchait le dessèchement du bois mort. Il s’y ensuivit que la respiration du bois mort y est nettement plus élevée en comparaison avec des pessières brûlés ou vierges. La décomposition du bois mort après coupe à blanc est aussi favorisée par la récupération plus rapide de la couche de mousses, diminuant conséquemment le dessèchement du bois mort par la conservation d’humidité, les transports vertical et horizontale d’eau et le refroidissement induit par l’humidité. Ainsi, les mousses sont les facteurs clés dans les cycles du bois mort et du C des pessières noires boréales au Labrador. L’auteur préconise la classification de ces pessières et des forêts semblables comme un groupe fonctionnel d’écosystèmes nommé : « pessières boréales humides » ; provisoirement définies comme « des écosystèmes forestiers avec une végétation terrestre dominée par les mousses et par conséquent associée avec des températures basses du sol, une humidité élevée, des taux de décomposition faibles et (en l’absence de perturbations) l’accumulation du bois enterré dans des couches organiques epaisses ». En outre, les mousses sont des éléments principaux des nombreuses forêts résineuses n’appartenant pas au biome boréal. Les processus régulés par les mousses tels l’enterrement du bois mort ou la paludification sont probablement importants pour le cycle global de C. La libération potentielle de grandes quantités de CO2 des réservoirs « bois enterré » et « sol » à la suite des changements climatiques exige une meilleure compréhension des transformations de la productivité des mousses et des limitations de la décomposition dues aux températures plus élevées et au taux d’humidités variables. Ainsi, les écosystèmes aux pertes potentielles de C élevées (p.ex. les pessières boréales humides) peuvent être identifiés et des mesures d’aménagement antagonistes peuvent être développées et implémentées. Traduction assistée par : Karl-Heinrich von Bothmer, Géry van der Kelen
Die vorliegende Arbeit untersucht die Einflüsse von Moosen auf den Totholz- und Kohlenstoff-(C)-Kreislauf in borealen Schwarzfichtenwäldern in Labrador, Kanada. Um den C-Kreislauf dieses humiden borealen Waldökosystems zu charakterisieren, wurden alle bedeutenden C-Speicher (lebende Bäume, stehendes und liegendes Totholz, organische Auflage, Mineralboden) von drei Primärwald-, neun Kahlschlags- und drei Brandflächen quantifiziert. Die C-Dynamiken der Bäume und des oberiridischen Totholzes der Untersuchungsflächen ähnelten denen von trockeneren und/oder wärmeren borealen Wäldern, während die organische Auflage, der Mineralboden und das begrabene Totholz bedingt durch von Moosen regulierte Prozesse wie Totholzeinlagerung und Paludifizierung besonders hohe C-Vorräte aufwiesen. Mit dem umfangreichen C-Datensatz wurde das CBM-CFS3, das nationale kanadische C-Modell, am Beispiel Labradors im Hinblick auf seine Anwendbarkeit in humiden borealen Wäldern evaluiert. Nach Anpassung ausgewählter Parameter, z.B. der Totholzabbauraten, wurden die gemessenen C-Dynamiken der Bäume und des oberiridischen Totholzes vom Modell abgebildet. Das CBM-CFS3 wurde ursprünglich für staunässefreie, terrestrische Waldstandorte entwickelt und berücksichtigt keine mit Moosen oder hoher Bodenfeuchte assoziierten Prozesse, so dass es die hohen C-Vorräte des begrabenen Totholzes und des Bodens nicht widerspiegelte. Eine adäquate Abbildung der Akkumulation von C in diesen Speichern erfordert strukturelle Änderungen des CBM-CFS3 und anderer Wald-C-Modelle. Die Genauigkeit von Wald-C-Modellen könnte darüber hinaus durch eine Differenzierung der Totholzabbauraten in Abhängigkeit vom Störungsregime verbessert werden, da störungsspezifische Veränderungen von Temperatur und Feuchte von der Totholzatmung widergespiegelt werden. Im Untersuchungsgebiet limitierten geringe Holzfeuchten und hohe Holztemperaturen die Totholzatmung auf Brandflächen. In Primärwäldern wirkten dagegen hohe Holzfeuchten und geringe Holztemperaturen hemmend. Auf Kahlschlägen verhinderte die verbleibende Vegetation die Austrockung des Totholzes, was zu signifikant erhöhten Atmungsraten im Vergleich zu Brand- und Primärwaldflächen führte. Zudem wird der Totholzabbau auf Kahlschlen durch eine schnellere Erholung der Moosdecke als auf Brandflächen gefördert, da Moose durch ihr hohes Wasserspeichervermögen, vertikalen und horizontalen Wassertransport und feuchte-induzierte Kühlung der Austrockung des Totholzes entgegenwirken. Moose sind somit ein Schlüsselfaktor im Totholz- und C-Kreislauf der humiden borealen Schwarzfichtenwälder Labradors. Die Autorin empfiehlt die Klassifikation dieser und ähnlicher borealer Wälder als eine funktionelle Ökosystemgruppe namens “humid boreal forests”; vorläufig definiert als “boreale Waldökosysteme mit durch Moose dominierter Bodenvegetation und damit assoziierten niedrigen Bodentemperaturen, hohen Bodenfeuchten, geringen Abbauraten und (in Abwesenheit großflächiger Störungen) der Akkumulation von begrabenem Totholz in mächtigen organischen Auflagen”. Auch außerhalb des borealen Bioms sind Moose ein wesentlicher Bestandteil vieler Nadelwälder. Durch Moose regulierte Prozesse wie Totholzeinlagerung und Paludifizierung sind daher wahrscheinlich relevant für den globalen C-Kreislauf. Die durch den Klimawandel bedingte potentielle Freisetzung von großen Mengen CO2 aus begrabenem Totholz und dem Boden macht ein besseres Verständnis der zu erwartenden Veränderungen von Mooswachstum und Abbauhemmnissen als Folge erhöhter Temperaturen und variabler Feuchteverhältnisse erforderlich. Somit können Ökosysteme mit potentiell hohen C-Verlusten, wie z.B. humide boreale Wälder, identifiziert und diesen entgegenwirkende Bewirtschaftungsmaßnahmen entwickelt und umgesetzt werden

The world's tundra regions can be considered as the last unexploited terrestrial biome on the earth's surface. Bryophytes contribute significantly to primary production and nutrient cycling in tundra habitats but received considerably less attention than the vascular plants during research for the Tundra Biome Project of the International Biological Programme in the 1970s. The IBP synthesis recognised this and called for more bryophyte production data and improved methodology in future studies. The synthesis also identitied the need for more information on within-site variability, energy/chlorophyll relationships and mineral nutrient studies where tundra bryophyte production patterns are concerned. Marion Island in the sub-Antarctic has an oceanic "tundra" type of vegetation, and the island offers the characteristics of an ideal "natural laboratory" for conducting fundamental ecological research particularly in the field of primary production and energy flow. Studies at Marion Island paralleled those of the IBP but also concentrated on the vascular component of the vegetation. Bryophytes contribute up to 60% of the biomass and 90% of plant cover in the mires that dominate the well-vegetated coastal plain of Marion Island. Following the termination of IBP research therefore it was decided to extend these studies during the 1980s by measuring growth and characterising the major factors influencing growth patterns in Marion Island bryophytes. In this thesis information is given on the physical and biotic environment of Marion Island. Data is presented on the cover, biomass, seasonal and annual production values, and turn-over rates (decomposition) of 15 bryophyte species found across the full range of habit at types at Marion Island. An analysis is also made of growth-associated variables viz. soils, climate, plant energy, chlorophyll. water and mineral nutrient contents. Production was found to be related broadly to moisture-related variables, overlain by nutrient factors in some instances. Higher absolute production figures were obtained than at other tundra sites and interesting phenological data are discussed. The results suggest that decomposition is controlled mainly by tissue degradability factors and that decomposition is more significant than production in controlling biomass accumulation. A model is given for production/decomposition patterns across the range of habitat conditions at Marion Island. Several growth measurement techniques were tested and recommendations are made concerning methodology for future bryophyte production studies. By reference to ecological data collected during the study.

Climate change is expected to have a strong impact on subarctic ecosystems. Increased temperatures as well as altered precipitation and snow cover patterns are predicted to change species distribution and affect biogeochemical processes in the subarctic tundra. Bryophytes are an essential vegetational component in northern ecosystems, due to their high abundance and importance in many ecological processes. In this study the effects of elevation and altered snow cover on the temporal dynamics of freezing damage in three subarctic bryophyte species (Hylocomium splendens, Ptilidium ciliare, and Sphagnum fuscum) were studied in a snow manipulation field experiment in Abisko, during early spring. Soil temperature and field moisture of moss shoots were collected. A freeze-thaw incubation experiment was conducted to investigate the freeze-thaw cycle resistance of H. splendens and P. ciliare originating from habitats with two differing snow-cover thicknesses. Freezing damage differed significantly between the bryophytes species with P. ciliare experiencing the least and S. fuscum the highest damage. Damage was higher at the low elevation, possibly attributable to acclimation effects. Snow removal led to higher damage in moss shoots, but no interactions of the different snow cover treatments with elevation, species or time were found. Freezing damage increased over time and no recovery occurred, likely due to temporal patterns in soil freeze-thaw cycles during early spring. Soil freeze-thaw cycles were the main factor influencing damage in bryophytes after snow melt. Measured environmental parameters could not explain the entire variation in damage. Damage might additionally be attributable to increased UV radiation or disturbances by herbivores.

Diss. (sammanfattning) Stockholm : Stockholms universitet, 2010.
At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 2: Submitted. Paper 3: Manuscript. Paper 4: Manuscript.

Abstract In this thesis, I studied the patterns in the assemblage composition as well as the biogeography and ecology of spring macroinvertebrates and bryophytes in Finland. My main objectives were to assess the importance environmental variables to macroinvertebrate and bryophyte assemblage composition in springs at the level of multiple spatial scales. In addition, I assessed the importance of springs in the boreal mire landscape, and sought the ecological and environmental determinants of a key species in boreal springs. In a large-scale study, I also examined the concordance between macroinvertebrates and bryophytes across boreal ecoregions, and assessed how macroinvertebrate assemblage variation corresponds to terrestrially-based ecoregions. Locally, spring macroinvertebrate assemblage structure displays high variation between different kinds of mesohabitats within springs, highlighting the importance of careful sampling of all habitat types in spring surveys. Helocrenes and other aquatic-terrestrial ecotone habitats harbour the highest species diversity and most spring-dependent species among spring habitat types. Further, spring-influenced mire patches were shown to have distinct cranefly assemblages in the mire landscape and to harbour higher cranefly diversity than mire types with lower trophic status, emphasising the importance of springs for mire biodiversity. Regionally, a red-listed spring-dependent caddisfly species appeared to be a surrogate for a high spring conservation value, indicating high overall species diversity and the occurrence of additional red-listed species. On a large geographical scale, intersecting the boreal ecoregions, a pattern of gradual change of benthic macroinvertebrate assemblage composition from south to north was detected, largely corresponding to terrestrially-derived ecoregions. However, the physical attributes of springs also need to be taken into account in bioassessment studies. Macroinvertebrate assemblage variation also correlated with physical habitat-scale variables, but not with changes in water chemistry. In contrast, spring bryophyte assemblages showed a distinct response to variation in water chemistry, but not to variation in physical habitat characteristics. Bryophytes and insect assemblages were concordant with each other on the large geographical scale, although the concordance was rather weak. Because of their different kind of responses to the physical and chemical variables, insects and bryophytes of springs are poor surrogates for each other in boreal springs.

Unequal sex ratios are widespread in dioecious plants and understanding their cause is important to understanding fundamental aspects of their population dynamics, and yet what causes biased sex ratios in plants is still poorly understood. Competition experiments have been used in plants to predict the outcome of species interactions, but they have rarely been used to help explain sex ratio bias. This study used a response surface competition design to measure the relative competitive abilities of the sexes of the bryophyte Marchantia inflexa (a thallus liverwort of Marchantiaceae) to predict the outcome of competition before the onset of sexual reproductive structures. In bryophytes, dioecy and sex ratio bias is especially common, making them effective organisms for studying sex ratio bias. Given the frequency of female bias in bryophytes, the hypothesis was that females will show a higher competitive ability relative to males. The experiment was conducted in greenhouse conditions at several densities and proportions over the course of seven months. As individuals grew and formed clumps, identities were tracked, and growth measurements were made using photographs and computer imaging software. Both sexes grew on average 41% more with the opposite sex relative to their single-sex cultures. A model predicting future sex ratios showed coexistence between the sexes and predicted a male biased sex ratio of 3.2 males to 1 female. A trade-off was observed for males where single-sex cultures contained more asexual structures than mixed-sex cultures and the reverse for growth rate. Higher levels of asexual reproduction in males in single-sex cultures might be selected for to increase male dispersal for a higher probability of encountering females. This pattern was not found for females. The overyielding results suggest an interaction effect may exist due to niche differentiation between the sexes. In addition, the results suggest that in some dioecious plants a change in sex ratio can occur before differences in their allocation to sexual reproduction.

Chez les plantes vasculaires, le métabolisme des phénylpropanoïdes conduit à la synthèse de précurseurs de biopolymères structuraux tels que la lignine, ainsi que de nombreux composés antioxydants et anti-UV. Ce métabolisme phénolique est apparu lors de la colonisation des terres par les plantes et a été critique pour leur adaptation à ce nouvel environnement. Physcomitrella patens, une bryophyte phylogénétiquement proche des premières plantes terrestres, est un bon modèle pour l'étude de certains caractères ancestraux. P. patens est dépourvue de lignine. En combinant des approches phylogénomique, génétique et biochimique, ce travail démontre le rôle essentiel de deux BAHD hydroxycinnamoyl tranférases dans le métabolisme phénolique de la mousse et la formation de précurseurs de la cuticule, une couche hydrophobe qui recouvre les parties aériennes de la plante et lui confère une imperméabilité. Il suggère également que deux hydroxycinnamoyl transférases sont requises pour la formation des composés phénoliques solubles accumulés par la mousse. Une exploration préliminaire du métabolisme des flavonoïdes chez ce modèle révèle d’autre part le caractère incomplet et primitif de cette voie métabolique
In vascular plants, the phenylpropanoid metabolism leads to the synthesis of precursors of structural biopolymers such as lignin and of essential antioxidants and UV screens. The phenolic pathway leading to these compounds appeared upon plant land colonization and is thought critical for their adaptation to this new environment. Physcomitrella patens is a bryophyte, an early-diverging land plant and thus a good model to reveal ancestral traits. P. patens is devoid of lignin. Combining phylogenomic, genetic and biochemical approaches, this work demonstrates the essential role of two BAHD hydroxycinnamoyl transferases in the moss phenolic metabolism and in the formation of precursors of the cuticle, a hydrophobic layer, covering and conferring impermeability to the aerial parts of the plant. It also suggests that two nonredundant hydroxycinnamoyl transferases are required for the formation of the soluble phenolic compounds accumulated in moss. A preliminary exploration of the flavonoid metabolism in this model in addition reveals primitive features of this metabolic route

Aim: To compare the α, β and phylogenetic diversity of the ground bryophyte communities along a tropical altitudinal gradient in order to determine which processes govern these assemblages. Location: La Réunion island (55°39'E; 21 °00'S), in the western Indian Ocean. Methods: The ground bryophyte communities were surveyed along the eastern slope of Piton des Neiges (350 - 3050 m). α, β and phylogenetic diversity along the altitudinal gradient was determined and graphically presented. Results: The α diversity had two distinct peaks at 1150 - 1350 m and at 2750 m, these corresponded to a dominance in liverwort and moss species, respectively. The phylogenetic diversity along the altitudinal gradient was far greater than is predicted from the null models suggesting that ground bryophyte communities are structured according to "environmental filters". The mid-domain effect and mass effect could not be used to describe the changing diversity along the altitudinal gradient, however a correlation analysis revealed temperature and relative humidity influences the changing a diversity. Main Conclusions: In conclusion both ecological and evolutionary processes structure the ground bryophyte communities.

Numbers of red deer (Cervus elaphus) in the Highlands have more than doubled in the last few decades and conservation organisations have expressed concern over their potential impacts on the natural heritage. Bryophytes and lichens represent a substantial component of Scottish biodiversity and northwest Scotland has cryptogam assemblages that are rare elsewhere in the world. Few studies have separated the impacts of red deer from those of sheep and none has focused on the response of important bryophyte and lichen assemblages to red deer management. This thesis has started to address this shortfall in scientific knowledge. Research was centred at Letterewe and neighbouring estates near Loch Maree, Wester Ross where there has been no sheep grazing for decades. Existing exclosures were used to compare blocks of cryptogam communities associated with specific habitats either side of the deer fence. In this way bryophyte and lichen species cover, diversity, dominance and richness associated with low-lying siliceous rocks, globally rare wet heath/bog and the internationally important epiphytic Lobarion lichen community were investigated. Other approaches involved comparing cryptogam communities found on boulder tops inaccessible to red deer with those that were accessible, attempting to correlate red deer dung densities with localised impact to liverwort-rich oceanic heath and by contrasting summit moss-heath vegetation between two estates that have different mean deer densities. Liverworts associated with the mixed hepatic mat were also compared either side of a long-standing deer fence at Knoydart. These studies have shown that red deer and their management (using exclosures) have both positive and negative effects on important bryophyte and lichen assemblages in northwest Scotland. This body of research is presented as a series of self-contained scientific papers.

Bryophytes form an important component of northern vegetation communities. Mosses efficiently capture aerially deposited nutrients, restricting nutrient availability to the soil. Given that key ecosystem processes of northern ecosystems are nutrient-limited, understanding nutrient cycling of the moss layer is key to understanding ecosystem nutrient and C cycling in these systems. However, the role of the moss layer in regulating ecosystem-scale nutrient and C cycling, while potentially significant, is largely unknown. The aim of this thesis is to investigate the effect of the relative availability of N and P on aspects of bryophyte nutrient uptake, retention and C acquisition. The hypothesis investigated is that the availability of one nutrient will influence the demand for the other and thereby moss nutrient acquisition and retention mechanisms. To test this hypothesis, various aspects of moss nutrient cycling in response to the relative availability of N and P were investigated. As the C cycle is tightly linked to the N and P cycles, the hypothesis extended to include bryophyte C assimilation and decomposition processes of an arctic tundra. Bryophyte nutrient demand was chiefly governed by the tissue N:P ratio. Consequently, nutrient uptake, both from aerially deposited nutrients and through moss-cyanobacteria N2 fixation, and nutrient losses after a simulated rainfall event were mostly in response to the relative availability of N and P rather than the availability of one nutrient alone. This thesis provides novel evidence that ectohydric mosses have the ability to internally translocate nutrients. In conjunction with efficient nutrient capture, this trait makes mosses strong nutrient sinks which are likely to exert considerable control over ecosystem nutrient cycling. The relative availability of N and P played a role in C uptake of mosses. Through the production of recalcitrant litter and their insulating effect on soil microclimate mosses exerted an influence over ecosystem C cycling.

Includes bibliographical references.
This study examines the effects of fragmentation on bryophyte diversity in renosterveld vegetation of South Africa's west coast. The main objectives of this study are to; Document bryophyte species assemblages in remnant fragments of renosterveld vegetation, investigate the effect of fragment size, isolation and heterogeneity on bryophyte diversity patterns in renosterveld, determine the extent to which fragments are nested and evaluate the extent to which existing conservation measures are appropriate for the long-term survival of the bryophyte flora in this vegetation type. Seventy-two bryophyte species were recorded from 23 fragments ranging from 0.1 to 102 hectares.

L'adaptation des plantes à la vie terrestre il y a 500 millions d'années a joué un rôle majeur dans l'évolution de la vie sur Terre. Les plantes jouent toujours un rôle crucial à l'heure actuelle, en tant que base de la plupart des écosystèmes et, par conséquent, base de toutes les civilisations humaines. Comprendre leur adaptation aux modifications passées et présentes de leurs conditions de vie est une clef pour comprendre le passé et être capable de répondre aux défis agricoles futurs. Les sciences végétales ont fait de grands progrès dans la compréhension de la réponse des plantes à leur environnement, mais la plupart des études se sont concentrées sur la lignée des angiospermes, qui comprend les plantes cultivées. Néanmoins, pour avoir une vision plus large de l'adaptation des plantes terrestres (Embryophytes) à diverses conditions, dans le cadre de 500 millions d'années d'évolution sur la terre ferme, il est essentiel d'étudier d'autres lignées de plantes terrestres. Dans cette logique, ce travail se focalisera sur la plante modèle non vasculaire Marchantia polymorpha, dont la lignée a divergé des plantes vasculaires il y a environ 480 millions d'années. Nous avons développé un ensemble de données sur la diversité intraspécifique qui nous a permis de mettre en évidence certains mécanismes d'adaptation chez M. polymorpha. L'analyse des signatures de sélection sur les gènes nous a permis de distinguer les fonctions conservées soumises à une forte sélection purificatrice des fonctions variables soumises à une sélection équilibrante ou à des balayages sélectifs. En utilisant cet ensemble de données sur la diversité intraspécifique, des études d'association à l'échelle du génome (GWAS) ont pu être réalisées sur la réponse de M. polymorpha aux conditions climatiques mais aussi au champignon pathogène Colletotrichum nymphaeae (stress biotique). Enfin, un pangénome basé sur les gènes a été construit et a permis d'identifier des gènes avec une variation de présence-absence entre les accessions, qui sont souvent associés à la réponse au stress et à l'adaptation locale. En croisant ces trois approches, nous avons trouvé des familles de gènes qui semblent impliquées dans la réponse de M. polymorpha aux stress. Parmi elles, on peut citer les terpènes synthases, les peroxydases, les NBS-LRR (NLR), les lectines, les lipoxygénases ou les polyphénols oxydases. La plupart de ces fonctions sont partagées avec d'autres plantes terrestres, ce qui montre que la plupart des mécanismes généraux d'adaptation sont assez conservés chez les Embryophytes. Néanmoins, la plupart de ces familles de gènes présentent des caractéristiques propres à la lignée, telles que des gènes spécifiques, des expansions de famille ou des transferts horizontaux de gènes, qui différencient l'organisation des familles de gènes chez les Marchantia de celle connue chez les angiospermes. L'ensemble de ces résultats montre que les plantes terrestres partagent la plupart de leurs mécanismes d'adaptation, hérités de leur dernier ancêtre commun, et que ces fonctions générales ont subi des modifications spécifiques à chaque lignée, dépendant des différentes contraintes qui ont façonné les lignées de plantes terrestres
Plant adaptation to a terrestrial life 500 million years ago played a major role in the evolution of life on Earth. Plants still play a crucial role at present time, as bases of most ecosystems, and consequently base of all human civilizations. Understanding their adaptation to past and present modification of their living conditions is a key to understand the past and be able to respond to future agricultural challenges. Plants sciences made great advances in understanding plant response to their environment, but most studies focused on the angiosperm lineage which contains crops. Nevertheless, to get a broader picture of land plant (Embryophytes) adaptation to various conditions, in the framework of 500 million years of evolution on land, it is essential to study other land plant lineages. In line with this logic, this work will focus on the non-vascular plant model Marchantia polymorpha, whose lineage diverged from vascular plants around 480 million years ago. We developed an intraspecific diversity dataset that allowed us to uncover some mechanisms of adaptation in M. polymorpha. Analyses of selection signatures on genes enabled us to distinguish conserved functions under strong purifying selection from variable ones undergoing balancing selection or selective sweeps. Using this intraspecific diversity dataset, genome-wide associations studies (GWAS) could be performed on the response of M. polymorpha to climatic conditions but also to the fungal pathogen Colletotrichum nymphaeae (biotic stress). Finally, a gene-based pangenome was built and allowed identifying genes with a presence-absence variation between accessions, that are often associated with stress response and local adaptation. Crossing these three approaches, we found gene families that seem involved in M. polymorpha response to stresses. Among them can be cited the terpene synthases, the peroxidases, the NBS-LRR (NLR), the lectins, the lipoxygenases or the polyphenol oxidases. Most of these functions are shared with other land plants, showing that most general mechanism of adaptation are quite conserved in Embryophytes. Nevertheless, most of these gene families displayed lineage-specific characteristics, such as specific genes, family expansions or horizontal gene transfer, that differentiated gene family organization in Marchantia from the one known in angiosperms. Taken together, these results show that land plants share most of their mechanisms of adaptation, inherited from their last common ancestor, and that these general functions underwent lineage-specific modifications, that can hint at the different constraints that shaped the different lineages of land plants

Rapid climate change in Arctic regions is of concern due to important feedbacks between the Arctic land surface and the global climate system. A large amount of organic carbon (C) is currently stored in Arctic soils; if decomposition is stimulated under warmer conditions additional release of CO2 could result in an accelerating feedback on global climate. The strength and direction of Arctic C cycle - climate feedbacks will depend on the growth response of vegetation; if plant growth increases some or all of the extra CO2 emissions may be offset. Currently the Arctic is thought to be a small net sink for CO2, the expected balance of terrestrial C sinks and sources in the future is unknown. In this thesis I explore some of the critical unknowns in current understanding of C cycle dynamics in Arctic vegetation. Quantifying gross primary productivity (GPP) over regional scales is complicated by large spatial heterogeneity in plant functional type (PFT) in Arctic vegetation. I use data from five Arctic sites to test the generality of a relationship between leaf area index (LAI) and canopy total foliar nitrogen (TFN). LAI and TFN are key drivers of GPP and are tightly constrained across PFTs in Low Arctic Alaska and Sweden, therefore greatly simplifying the task of up-scaling. I use data from Greenland, Barrow and Svalbard to asses the generality of the LAI-TFN relationship in predicting GPP at higher Arctic latitudes. Arctic ecosystems are unique among biomes in the large relative contribution of bryophytes (mosses, liverworts and hornworts) to plant biomass. The contribution of bryophytes to ecosystem function has been relatively understudied and they are poorly represented in terrestrial C models. I use ground based measurements in Northern Sweden to fill an existing data gap by quantifying CO2 fluxes from bryophytes patches in early spring and summer, and develop a simple model of bryophyte GPP. Using the model I compare bryophyte GPP to that of vascular plants before, during and after the summer growing season, finding that productive bryophyte patches can contribute up to 90 % of modelled annual GPP for typical vascular plant communities at the same site, and that the relative magnitude of bryophyte GPP is greatest in spring whilst the vascular plant canopy is still developing. Understanding how GPP relates to plant growth is important in relating remotely sensed increases in Arctic ‘greenness’ to changes in plant C stocks. I use a 13C pulselabelling techniques to follow the fate of recently fixed C in mixed vascular and bryophyte vegetation, with a focus on quantifying the contribution of bryophytes to ecosystem carbon use efficiency (CUE). I show that bryophytes contribute significantly to GPP in mixed vegetation, and act to increase ecosystem CUE. I highlight the importance of including bryophytes, which do not have roots, in aboveground: belowground partitioning schemes in C models. To further explore C turnover in bryophytes, I use the results of a second 13C labelling experiment to develop a model of C turnover in two contrasting Arctic mosses (Polytrichum piliferum and Sphagnum fuscum). I find significant differences in C turnover between Polytrichum piliferum which respires or translocates about 80 % of GPP, while Sphagnum fuscum respires 60 %. This analysis is the first to explicitly model differences in C partitioning between Arctic bryophyte species. Finally, I discuss the implications of each chapter for our understanding of Arctic C dynamics, and suggest areas for further research.

Ecological restoration is the practice of assisting the recovery of a degraded, damaged or destroyed ecosystem. The aim of this study was to analyse the effects of two different restoration techniques on instream bryophyte abundance, species richness and diversity as well as community composition, in streams channelized for timber-floating. Instream bryophytes were collected from 10 tributaries of the Vindel River in boreal northern Sweden, from five stream reaches each of channelized reaches, which had not been restored; reaches restored in the early 2000s, using best-practice techniques; and reaches restored in the early 2000s and then re-restored in 2010 using the new “demonstration” techniques. A multitude of environmental variables were also measured at the sites. Bryophyte abundance was lower in demonstration restored sites than unrestored or best-practice restored sites but no significant difference was found in bryophyte species richness, diversity or species composition. Environmental variables correlated with bryophyte abundance, species richness, diversity and composition largely reflected effects of restoration, and probably the disturbance associated with restoration. Small sediment grain sizes also had a negative effect on species richness. Other environmental variables that influenced bryophyte species composition were the large-scale factors of latitude, longitude and elevation and reach-scale factors of potassium concentration and light absorbance. It is not yet possible to fully evaluate the effectiveness, in terms of bryophyte response, of the new demonstration restoration compared with best-practice since the recovery time between them differs and has not been sufficient. Long-term monitoring of the effects of restoration is needed in order to better evaluate success.

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.

Peat mosses (Sphagnum) are widely distributed and are a major component of mire vegetation and peat throughout the boreal and temperate regions. Most boreal Sphagnum species regularly produce sporophytes, but the ecological role of the spore has been questioned. This study shows that the spores can form a spore bank and have the ability to germinate and contribute to moss establishment whenever suitable conditions occur. The results suggest that spore production is important for explaining the wide distribution and omnipresence of Sphagnum in nutrient-poor wetlands. The results further imply that initial recruitment from spores predominates in Sphagnum after disturbance or formation of suitable habitats.

A series of experiments showed that addition of phosphorus-containing substrates, such as fresh plant litter or moose dung, resulted in spore establishment on bare, moist peat. A field experiment indicated establishment rates of about 1% of sown, germinable spores on peat with added substrates. Plant litter on moist soil, without a closed cover of bryophytes, is an important safe site for the establishment of Sphagnum spores. The results fit the observed pattern of colonisation by Sphagnum beneath Eriophorum vaginatum tussocks in mires severely disturbed by peat extraction. Successful long-distance dispersal was indicated by the occurrence of several regionally new or rare Sphagnum species in disturbed mires.

Spore number per sporophyte ranged among Sphagnum species from 18 500 to 240 000, with a trade-off between spore number and spore size. Annual spore production was estimated at 15 million spores per square metre on two investigated mires. Sporophyte production showed a large interannual variation. Sporophyte production was positively related to the amount of precipitation the preceding summer. This was probably because a high water level promoted gametangium formation. Spore dispersal occurred in July and August. The earlier timing of spore dispersal in the more drought-sensitive, hollow-inhabiting sphagna should reduce the risk of sporophytes drying out prematurely during summer droughts.

Spores kept refrigerated up to 13 years retained high germinability. A field experiment showed that Sphagnum can form a persistent spore bank, with a potential longevity of several decades.

Kinabalu National Park (KNP) has the status as forest reserve class VI: virgin jungle forest in Malaysia. In 2000, it also has been recognized by UNESCO for its “outstanding universal values”. With an area size of 754 Km² and the highest peak at 4,095.2 metres, it provides a diversed habitat for ecologically important flora and fauna. Thus, from the Dipterocarp forest (lowland) to the sub-alpine forest (highland), each forest type offers different benefits to us. Ecologically speaking, montane forest (1200 - 2700m) is essential primarily as water catchment area though, this ecosystem often being forgotten due to its “low economic values” perceived by ordinary people. In fact, its economic value is high but since less people know about its free ecosystem services, it is unfairly assumed as less important in comparison to other type of forest, such as mangrove forest. In addition to that, the dominant communities inside the montane forest are bryophytes or in laymen’s term as mosses. In a nutshell, the functioning component of the moist and foggy forest is the mossy forest. As a reference to the bryodiversity in KNP, there are around 8000 species of bryophytes checklisted in the Malesian region.
Kinabalu National Park (KNP) ha lo status di riserva forestale classe VI: giungla, foresta vergine in Malesia. Nel 2000, inoltre è stato riconosciuto dall'UNESCO per i suoi "valori universali" in circolazione. Con una dimensione di un'area di 754 km ² e la cima più alta a 4,095.2 metri, fornisce un habitat per la flora diversed ecologicamente importanti e fauna. Così, dalla foresta dipterocarp (pianura) al sub-alpino forestale (altopiano), ogni tipo di foresta offre diversi vantaggi a noi. Ecologicamente parlando, foreste montane (1200 - 2700) è essenziale in primo luogo come bacino idrografico, però, questo ecosistema essendo spesso dimenticata a causa dei suoi "valori bassi economici" percepiti dalla gente comune. Infatti, il suo valore economico è alto ma dal momento che meno gente conoscere i suoi servizi ecosistemici gratuiti, è ingiustamente assunto come meno importante rispetto ad altro tipo di foresta, come la foresta di mangrovie. In aggiunta a ciò, le comunità dominanti all'interno della foresta di montagna sono briofite o in termini laici di come muschi. In poche parole, il componente funzionamento della foresta umida e nebbiosa è la foresta di muschio. Come riferimento alla bryodiversity in KNP, ci sono circa 8000 specie di briofite checklisted nella regione Malese.

Dispersal, especially long-distance dispersal, is an important component in many disciplines within biology. Many species are passively dispersed by wind, not least spore-dispersed organisms. In this thesis I investigated the dispersal capacity of bryophytes by studying the colonization patterns from local scales (100 m) to landscape scales (20 km). The dispersal distances were measured from a known source (up to 600 m away) or inferred from a connectivity measure (1–20 km). I introduced acidic clay to measure the colonization rates over one season of a pioneer moss, Discelium nudum (I–III). I also investigated which vascular plants and bryophytes that had colonized limed mires approximately 20–30 years after the first disturbance (IV). Discelium effectively colonized new disturbed substrates over one season. Most spores were deposited up to 50 meters from a source but the relationship between local colonization rates and connectivity increased with distance up to 20 km (I–III). Also calcicolous wetland bryophyte species were good colonizers over similar distances, while vascular plants in the same environment colonized less frequently. Common bryophytes that produce spores frequently were more effective colonizers, while no effect of spore size was detected (IV). A mechanistic model that take into account meteorological parameters to simulate the trajectories for spores of Discelium nudum fitted rather well to the observed colonization pattern, especially if spore release thresholds in wind variation and humidity were accounted for (III). This thesis conclude that bryophytes in open habitats can disperse effectively across landscapes given that the regional spore source is large enough (i.e. are common in the region and produce spores abundantly). For spore-dispersed organisms in open landscapes I suggest that it is often the colonization phase and not the transport that is the main bottle-neck for maintaining populations across landscapes.

At the time of the doctoral defence the following papesr were unpublished and had  a status as follows: Paper 2: Epubl ahead of print; Paper 3: Manuscript; Paper 4: Manuscript

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.

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.

Les conséquences sanitaires et environnementales de la pollution atmosphérique sont devenues au cours des dernières décennies un des problèmes majeurs de notre société. Sensibilisée à cette problématique, la région Nord Pas-de-Calais a été l’une des premières régions à soutenir et à valoriser la biosurveillance végétale de la qualité de l’air. Les travaux scientifiques réalisés dans le cadre de cette thèse ont pour objectif d’étudier les phénomènes d’accumulation et les impacts des composés organiques volatils (COV) chez les bryophytes. Articulées autour de deux axes, les recherches menées visaient à étudier, d'une part, l'accumulation des benzène, toluène, ethylbenzène et xylènes (BTEX), d'autre part, des effets au niveau cellulaire, du benzène chez Tortula ruralis, par l'analyse des variations de l'expression de biomarqueurs de stress oxydant. Des expérimentations complémentaires in situ , sur le littoral dunkerquois, et en conditions contrôlées ont permis d'établir des phénomènes d'équilibre des concentrations en BTEX entre notre modèle végétal et l'air. In situ, les teneurs en BTEX chez les mousses sont fortement conditionnées par les paramètres météorologiques, et principalement la force des vents. De plus, les concentrations atmosphériques en particules PM10 favoriseraient l'accumulation des BTEX chez la mousse. Mais d'une manière générale, les mousses n'accumulent pas les BTEX de façon corrélée aux teneurs dans l'atmosphère. Concernant l'expression des biomarqueurs de stress oxydant, des variations naturelles des paramètres biologiques mesurés sont observées chez la mousse. Ces variations seraient liées à la nature poïkilohydrique des mousses, leur capacité de reviviscence et aux aptitudes qu'elles ont développées pour résister aux variations de leur état d'hydratation. Ainsi, les biomarqueurs de stress oxydant souvent recherchés dans les études d'écotoxicologie, ne seraient pas adaptés pour l'étude des effets chez les bryophytes terrestres.

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.

De par leurs propriétés morphologiques et physiologiques (l'absence de système racinaire et de tissus conducteurs développés faisant du dépôt atmosphérique la principale source en eau et éléments nutritifs), les mousses sont couramment utilisées comme sentinelles de la pollution métallique atmosphérique. Cependant, l'analyse des mousses ne permet pas d'estimer directement la valeur réelle du dépôt atmosphérique. Afin de convertir les concentrations mesurées dans les mousses en flux de retombées atmosphériques, divers paramètres doivent être renseignés (efficacité de capture et de rétention des métaux par la mousse, production annuelle de biomasse) ; tel fut l'objet de ce travail de recherche. Cette étude de processus est menée sur un site atelier en milieu rural, loin de sources ponctuelles de pollution (Vouzon, forêt domaniale de Lamotte Beuvron, Loire et Cher), sur un cycle annuel (octobre 2001-2002) et s'intéresse à une espèce de mousse : Scleropodium purum (Hedw. ) Limpr. Au niveau de l'organisme, le suivi temporel du développement morphologique (allongement du brin et production de biomasse) et physiologique (dégradation de la chlorophylle) a contribué à mieux renseigner la biologie de l'espèce échantillonnée. Un allongement moyen de 2,2 cm. Anˉ¹ du brin principal et un accroissement de biomasse de 103,7±29,5 g. Mˉ². Anˉ¹ ont ainsi été mesurés. Une caractérisation des diverses sources d'erreur (du prélèvement à l'analyse) a montré la faible variabilité associée à l'échantillonnage des mousses, validant ainsi le protocole usité. Le suivi régulier des retombées atmosphériques totales et des concentrations élémentaires mesurées dans les mousses a permis de confronter la valeur réelle du flux de dépôt annuel à la valeur estimée à partir des mousses. Pour chaque élément, un pourcentage d'efficacité de capture et de rétention par la mousse a ainsi pu être calculé. Le classement des éléments, en fonction de la valeur du pourcentage d'efficacité, conduit à la séquence suivante : Mn > K > P >Ca > Hg >Mg = Zn > Cu > Li > Sr > Ba > V > Pb > Fe > Na = Al > Si, où le flux de dépôt annuel réel est surestimé pour Mn et sous-estimé pour Si en utilisant les mousses. Divers phénomènes mis en évidence au cours de cette étude peuvent influer sur l'efficacité de capture et de rétention des éléments par le brin de mousse et ainsi expliciter les différences relevées entre le dépôt réel et estimé par les mousses.

La restauration écologique des tourbières se définit comme l’ensemble des procédés permettant d’assister à la régénération d’une tourbière qui a été dégradée, endommagée ou détruite. La gamme des tourbières perturbées qui nécessitent des actions de restauration est très diversifiée. L’étude de la régénération naturelle des écosystèmes perturbés a fréquemment été utilisée pour développer des méthodes de restauration qui sont adaptées aux types spécifiques de ces écosystèmes. Une compréhension plus complète des trajectoires de régénération, des contraintes environnementales et de dispersion, ainsi que des interactions interspécifiques, est directement reliée au développement de méthodes de restauration et à l’amélioration de nos connaissances sur la réponse des espèces et des communautés à des changements environnementaux. Cette thèse étudie la régénération de la végétation, plus spécialement des bryophytes, ainsi que les facteurs influençant leur régénération dans trois cas problématiques de restauration des tourbières perturbées. Premièrement, la régénération des communautés de bryophytes a été étudiée en fonction des conditions environnementales dans 28 tourbières minérotrophes des Territoires du Nord-Ouest et de l’Alberta ayant brûlé il y a un, deux ou cinq ans. Un changement de dominance a été observé entre les bryophytes pionnières (couvert moyen passant de 16 à 11 %) et les bryophytes de succession tardive (3 à 27 %) entre la 2e et la 5e année après le feu. La répartition et l’abondance des espèces de bryophytes semblent être contrôlées par leurs différences de capacité de régénération en lien avec leur stratégie d’histoire de vie, leur niche écologique et l’effet variable de la biomasse carbonisée. L’intégration de la notion de niche écologique chez les bryophytes de succession tardive ainsi que l’utilisation des espèces ayant un fort potentiel de régénération (comme Aulacomnium palustre et Ptychostomum pseudotriquetrum) devraient être considérées lors de l’élaboration des plans de restauration des tourbières minérotrophes. Deuxièmement, la régénération des sphaignes a été étudiée dans 25 sites donneurs (tourbières ombrotrophes où la végétation de surface a été récoltée pour servir de matériel de réintroduction) sur une chronoséquence de 17 ans en fonction des conditions environnementales et des pratiques de récolte. Les sphaignes dans les sites donneurs se régénèrent en moyenne en 10 ans, quoique des différences mineures dans la composition et l’abondance des espèces ont été observées entre les sites donneurs et les tourbières naturelles. Les espèces pionnières (p. ex., Eriophorum vaginatum), compétitives (p. ex., Sphagnum rubellum) et préférentielles des microhabitats plus humides (p. ex., Chamaedaphne calyculata) étaient plus abondantes dans les sites donneurs. Les pratiques de récolte avaient l’effet le plus important sur la régénération de la végétation, suggérant qu’il est possible d’accélérer la reprise des sphaignes en adoptant des pratiques de récolte appropriées. Des pratiques responsables consistent à éviter les sites donneurs très humides et la récolte durant le milieu de l’été lorsque les mousses ont un plus faible potentiel de régénération et lorsque le risque d’enlisement de la machinerie est élevé. Troisièmement, la régénération et le développement de tapis de sphaignes ont été étudiés dans deux types de tourbières restaurées et colonisées par des plantes pionnières de type graminoïde (Cypéracées). Dans le premier cas, le Carex aquatilis dominait un fen modérément riche restauré et formait des communautés denses avec beaucoup de litière. Dans le deuxième cas, l’Eriophorum angustifolium était dominant avec un faible couvert de litière dans une tourbière à sphaignes restaurée où la densité de la plante graminoïde avait été contrôlée par coupe répétée. Il a été démontré que l’effet des plantes graminoïdes sur les tapis de sphaignes était déterminé par leur litière plutôt que par la structure de la plante elle-même. En raison de l’ombre créée par la litière compacte, la présence d’une communauté dense d’une plante graminoïde accumulant beaucoup de litière comme Carex aquatilis ne semble pas être bénéfique pour développer des tapis de sphaignes productifs. La décision de contrôler les plantes graminoïdes pour accroitre le développement des tapis de sphaignes devrait prendre en considération, bien sûr, le couvert de la plante (non requis si le couvert moyen < 30 %), mais aussi sa forme de croissance (moins essentiel lorsque la plante pousse en tige unique), de même que ses potentiels d’accumulation de litière et d’envahissement (p. ex., Phragmites australis, Scirpus cyperinus). En somme, en plus de répondre spécifiquement à trois cas problématiques de restauration de tourbières perturbées, cette thèse fournit une compréhension améliorée des processus de régénération naturelle des écosystèmes de tourbières et des facteurs qui les influencent. Les résultats ont un lien direct avec le développement d’outils pour la restauration et avec l’approfondissement de nos connaissances sur les réponses des organismes et des communautés végétales aux changements de l’environnement.
The ecological restoration of peatland is defined as all the processes permitting the regeneration of a peatland that has been degraded, damaged or destroyed. The landscape settings and types of peatlands that require restoration actions is very diverse. The study of natural regeneration of disturbed ecosystems has frequently been used to develop restoration methods that are adapted to specific types of ecosystems. A comprehensive assessment of regeneration trajectories and the environmental and dispersal constraints, as well as the interspecific interactions is directly related to the development of restoration methods and the improvement of our knowledge on the response of species and communities to environmental changes. This thesis studies the regeneration of vegetation, especially of bryophytes, and the factors influencing their regeneration in three problematic cases of restoring disturbed peatland. Firstly, the regeneration of bryophyte communities was studied against a range of environmental conditions in 28 minerotrophic peatlands in the Northwest Territories and Alberta that had burned one, two or five years ago. A shift of dominance between pioneer (mean cover from 16 to 11 %) and late successional bryophytes (from 3 to 27 %) was observed between the second and fifth year after the fire. Bryophyte distribution and abundance were controlled by the species difference in terms of regeneration capacity related to their life history strategy, habitat niche, and the variable effect of the burned biomass. The integration of the ecological niche concept for late successional bryophytes and the use of species with a high potential of regeneration (such as Aulacomnium palustre and Ptychostomum pseudotriquetrum) should be considered during the elaboration of restoration plans of minerotrophic peatlands. Secondly, the regeneration of Sphagnum mosses was studied in 25 donor sites (peatlands where surface vegetation was harvested to supply reintroduction material) on a chronosequence of 17 years according to environmental conditions and harvesting practices. It takes on average 10 years for Sphagnum mosses in donor sites to recover, though minor differences in the species composition and abundance was observed between donor sites and natural peatlands. Pioneer species (e.g., Eriophorum vaginatum), competitive species (e.g., Sphagnum rubellum) and preferential species of wet microhabitats (e.g., Chamaedaphne calyculata) were more abundant in donor sites. Harvesting practices had the greatest effect on the regeneration of donor sites, suggesting that it is possible to speed up Sphagnum regeneration by choosing appropriate management practices. Best practices would be avoiding very wet donor sites and harvesting in the midsummer should be avoided when mosses are at their lowest regeneration potential and when the risk of machinery sinking is high. Thirdly, the regeneration and development of the Sphagnum carpets were studied in two experiments conducted in two types of restored peatlands colonized by pioneer graminoid plants (Cyperaceae). In the first experiment, Carex aquatilis dominated a moderately rich restored fen and formed dense communities with a lot of litter. In the second experiment, Eriophorum angustifolium was dominant with a low cover of litter in a restored cutover Sphagnum-dominated bog where the graminoid plant cover was controlled by repeated mowing. It was demonstrated that the effect of graminoid plants on the Sphagnum carpet was determined by the litter rather than by the structure of the plant itself. Because of the shade created by the dense litter, the presence of a community of a graminoid plant that accumulates a lot of litter such as Carex aquatilis is not beneficial for the formation of productive Sphagnum carpets. The decision to control graminoid plants to enhance the growth of Sphagnum carpets should consider the cover of the plant (not necessary when mean cover < 30 %), but also its life form (less necessary when the plant grows in unique stem), its litter accumulation and potential for invasion (e.g. Phragmatis australis, Scirpus cyperinus). In conclusion, this thesis, in addition to specifically address these three problematic cases, provides a more comprehensive assessment of the processes of natural regeneration of peatland ecosystems and of driving factors. The results have a direct link with the development of tools for the restoration of peatlands and with a deeper understanding of the responses of plants and communities to environmental changes.

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

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A família Rhabdoweisiaceae Limpr. inclui musgos acrocárpicos, de tamanho pequeno a médio, que crescem, predominantemente como tufos e ocorrem, geralmente, sobre rochas e solo. A circunscrição da família é controversa, sendo sua monofilia sustentada por dados moleculares. O presente trabalho tem como objetivo realizar o estudo palinológico de espécies de Rhabdoweisiaceae Limpr. ocorrentes nas Américas e interpretar as relações entre as características palinológicas e as estratégias ecológicas das espécies estudadas, que pertencem aos gêneros Amphidium Schimp.; Arctoa Bruch & Schimp.; Cynodontium Schimp; Dichodontium Schimp.; Dicranoweisia Lindb. ex Milde; Kiaeria I. Hagen; Oncophorus (Brid.) Brid.; Oreas Brid.; Oreoweisia (Bruch & Schimp.) De Not.; Rhabdoweisia Limpr. Os esporos foram observados sob microscópio de luz antes e após acetólise e sob microscópio eletrônico de varredura e de transmissão. Para a avaliação das relações entre a morfologia dos esporos e as estratégias ecológicas foi realizada a análise de agrupamento, considerando os tipos de elementos de ornamentação, espessura do esporoderma polaridade, condição de tamanho, área apertural, forma de vida e substrato. As espécies estudadas apresentam esporos de tamanho pequeno a médio, com simetria radial e âmbito subcircular, são heteropolares ou apolares. O esporoderma é formado por intina, exina e perina. Os elementos de ornamentação permitiram a caracterização de quatro tipos morfológicos. Das 23 espécies estudadas, nove, incluídas em três gêneros apresentam esporos anisomórficos, sendo aqui relatados pela primeira vez. Os esporos das espécies de Amphidium Schimp. apresentam características palinológicas que permitem diferenciá-las das demais espécies integrantes de Rhabdoweisiaceae Limpr. Os padrões de ornamentação da superfície do esporoderma mais frequentes foram o baculoide e o granuloide; cerca de 80% das espécies analisadas apresentam área apertural diferenciada; dentre as carcterísticas ecológicas, rocha foi o substrato mais frequente, seguido por solos e a maioria das espécies apressentm tufo como forma de vida. As espécies de Rhabdoweisiaceae Limpr. analisadas neste trabalho foram agrupadas através de características palinológicas e ecológicas.
The family Rhabdoweisiaceae Limpr. includes small to medium sized acrocarpic mosses that grow predominantly as tufts and usually occur on rocks and soil. The circumscription of the family is controversial, and its monophyly is supported by molecular data. The present work aims to carry out the palynological study of species of Rhabdoweisiaceae Limpr. occurring in the Americas and to interpret the relationships between the palynological and ecological strategic characteristics of the species studied, which belong to the genera Amphidium Schimp.; Arctoa Bruch & Schimp.; Cynodontium Brid.; Dichodontium Schimp.; Dicranoweisia Lindb. ex Milde; Kiaeria I. Hagen; Oncophorus (Brid.) Brid.; Oreas Brid.; Oreoweisia (Bruch & Schimp.) De Not.; Rhabdoweisia Limpr. The spores were observed under light microscopy before and after acetolysis and under scanning and transmission electron microscopy. For the evaluation of the relationships between the spore morphology and the ecological strategies, the grouping analysis was performed considering the types of ornamentation elements, sporoderma polarity thickness, size condition, aesthetic area, life form and substrate. The species studied present small to medium spores, with radial symmetry and subcircular extent, are heteropolar or nonpolar. The sporoderm is formed by intina, exina and perina. The ornamentation elements allowed the characterization of four morphological types. Of the 23 species studied, nine, included in three genera present anisomorphic spores, being reported here for the first time. The spores of Amphidium Schimp. species have palynological characteristics that allow them to differentiate them from the other Rhabdoweisiaceae Limpr species. The most frequent patterns of ornamentation of the sporoderm surface were baculoid and granuloid; about 80% of the analyzed species present a distinct aperural area. Among the ecological characteristics, rock was the most frequent substrate, followed by soils and most species apressentm tuff as a way of life. The species of Rhabdoweisiaceae Limpr. analyzed in this work were grouped through palynological and ecological characteristics.