| 1. |
- Dahlberg, C. Johan, 1978-
(författare)
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The role of microclimate for the performance and distribution of forest plants
- 2016
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- Microclimatic gradients may have large influence on individual vital rates and population growth rates of species, and limit their distributions. Therefore, I focused on the influence of microclimate on individual performance and distribution of species. Further, I examined differences in how microclimate affect species with contrasting distributions or different ecophysiological traits, and populations within species. More specifically, I investigated the performance of northern and southern distributed forest bryophytes that were transplanted across microclimatic gradients, and the timing of vegetative and reproductive development among northern, marginal and more southern populations of a forest herb in a common garden. Also, I compared the landscape and continental distributions across forest bryophytes and vascular plants and, thus, their distribution limiting factors at different spatial scales. Lastly, I examined the population dynamics across microclimatic gradients of transplants from northern and southern populations of a forest moss. The effects of microclimatic conditions on performance differed among bryophytes with contrasting distributions. There were no clear differences between northern and southern populations in the timing of development of a forest herb or in the population dynamics of a moss. However, within each region there was a differentiation of the forest herb populations, related to variation in local climatic conditions and in the south also to proportion of deciduous trees. The continental distributions of species were reflected in their landscape distributions and vice versa, in terms of their occurrence optima for climatic variables. The variation in landscape climatic optima was, however, larger than predicted, which limit the precision for predictions of microrefugia. Probably, the distributions of vascular plants were more affected by temperature than the distributions of bryophytes. Bryophytes are sensitive to moisture conditions, which was demonstrated by a correlation between evaporation and the population growth rate of a forest moss. We might be able to predict species’ landscape scale distributions by linking microclimatic conditions to their population growth rates, via their vital rates, and infer larger scale distribution patterns.
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| 2. |
- Dahlgren, Johan Petter, 1978-
(författare)
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Linking plant population dynamics to the local environment and forest succession
- 2008
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- Linking environmental variation to population dynamics is necessary to understand and predict how the environment influences species abundances and distributions. I used demographic, environmental and trait data of forest herbs to study effects of spatial variation in environmental factors on populations as well as environmental change in terms of effects of forest succession on field layer plants. The results show that abundances of field layer species during forest succession are correlated with their functional traits; species with high specific leaf area increased more in abundance. I also found that soil nutrients affect vegetative and flowering phenology of the forest herb Actaea spicata. The effect of nutrients shows that a wider range of environmental factors than usually assumed can influence plant phenology. Moreover, local environmental factors affected also the demography of A. spicata through effects on vital rates. An abiotic factor, soil potassium affecting individual growth rate, was more important for population growth rate than seed predation, the most conspicuous biotic interaction in this system. Density independent changes in soil potassium during forest succession, and to a lesser extent plant population size dependent seed predation, were predicted to alter population growth rate, and thereby the abundance, of A. spicata over time. Because these environmental factors had effects on population projections, they can potentially influence the occupancy pattern of this species along successional gradients. I conclude that including deterministic, as opposed to stochastic, environmental change in demographic models enables assessments of the effects of processes such as succession, altered land-use, and climate change on population dynamics. Models explicitly incorporating environmental factors are useful for studying population dynamics in a realistic context, and to guide management of threatened species in changing environments.
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| 3. |
- Fogelström, Elsa, 1986-
(författare)
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Plant phenology in seasonal environments
- 2019
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- Phenology, or the seasonal timing life-history events such as emergence, reproduction and senescence will determine the outcome of interactions between plants and both abiotic and biotic aspects of the environment. Such timing is therefore of utmost importance for plants in seasonal environments. In this thesis, I first investigated the factors determining the start, end and length of the growing season for a perennial herb. Secondly, I estimated phenotypic selection on flowering time and investigated to which extent it corresponded to genotypic selection in a natural field setting. Thirdly, I estimated population differentiation in flowering time in a common garden and in the field. Lastly, I experimentally manipulated the synchrony of a perennial herb and its main herbivore to investigate the effects of herbivore phenological preference and plant-herbivore synchrony on the direction of selection on flowering time.I found that flowering individuals emerged earlier in spring than non-flowering individuals and that large individuals senesced later in autumn, suggesting that the length of the growing season is linked to individual condition and resource demands. Phenotypic selection favoured early-flowering individuals, but there was no genotypic selection. I found evidence for genetic population differentiation in flowering time in a common garden but not in the field. This suggests that, although flowering time has a genetic component, the observed variation in flowering time was mainly plastic under natural field conditions. Lastly, I show that constant herbivore preferences of plant phenology, in combination with environmentally driven variation in relative synchrony of the plant and the herbivore, leads to among-year variation in natural selection on flowering time. With this thesis, I contribute to identifying the factors affecting plant phenology as well as of the mechanisms shaping selection on flowering time in perennial plants. Such knowledge is essential for predicting species responses to climate change.
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| 4. |
- Andersson, Petter, 1975-
(författare)
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The importance of search behavior and movements for spatial distributions of herbivorous insects
- 2011
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- Insect populations commonly show large spatial variation in density, and much variation have been shown to be explained by the search behavior applied by the insect when locating habitat patches. This thesis explores the importance of odor-mediated attraction for immigration rates of herbivorous insects in relation to the size of the patches and the density of host plants within the patches. By using electroantennogram and measuring moth antennal responses to sex pheromones and thereby estimating the relative odor-mediated attraction from odor patches in field experiments (Paper I, III), I show that the length of the odor plumes emanating from the patches increase proportional with the square-root of the number of odor sources. In laboratory and field experiments (with the weevils Cionus scrophulariae and C. tuberculosus and the host plant figwort Scrophularia nodosa; Paper II) and meta-analyses on multiple insect herbivores (Paper III), I examined whether the relative increase in plume length could also predict the immigration rates of olfactory searching insects in relation to patches with increasing area and increasing density of host plants. The experiments (Paper II) and meta-analyses (Paper III) showed that the observed immigration rates of olfactory searching insects was well predicted by the relative increase in plume length, as estimated from the electroantennogram measurements (Paper I, III). The importance of immigration rates, relative to the effect of emigration and local growth was also investigated for the Cionus weevils in natural S. nodosa patches (Paper IV). This study showed that the density-patch size relationships of the weevils during early season were predicted by the net scaling of emigration and immigration rates, and differences in density-patch size relationships between the two species could be explained by inter-specific differences in their emigration rates from small patches. In conclusion, this thesis shows that search behavior can be used to predict immigration rates and spatial distributions of insects, with implications for pest control, conservation ecology and general ecological theory.
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| 5. |
- Arnell, Matilda, 1987-
(författare)
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Distribution patterns of fleshy-fruited woody plants at local and regional scales
- 2022
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- Fleshy-fruited woody plants share a long history with humans, providing us with food and wood material. Because of this relation, we have actively moved some of these plants across landscapes and continents. In Sweden, these species are often found in open and semi-open habitats such as forest edges, their fruits are most often dispersed by birds and their flowers are, with some exceptions, pollinated by insects. In this thesis my overall aim was to map and analyse distribution patterns of fleshy-fruited woody plants in Sweden to expand our knowledge on the mechanisms governing their distributions. First, I mapped a population of the early flowering, fleshy-fruited shrub Daphne mezereum (common mezeron, tibast) and surveyed the reproduction and fruit removal of all individuals (chapter I). My main aim was to investigate to what extent reproduction and fruit removal was affected by local distribution patterns. Secondly, I mapped local distribution patterns of fleshy-fruited woody species and analysed spatial associations between life stages and species (chapter II). My main aim was to relate these spatial associations to predictions of how bird dispersal would shape the local distribution patterns and the hypothesis that birds create ‘wild orchards’. Thirdly, I digitized historical maps and surveyed fleshy-fruited woody species along transects across landscapes (chapter III). My aim was to examine the hypothesis that these species accumulate in open and semi open habitats created by human land use. Fourthly, I estimated range filling of woody plants in Sweden at a 1 km2 resolution (chapter IV). My aim was to compare these estimates among species with different dispersal systems to understand the effect of dispersal on the occupancy of woody species at regional scales.I found the distribution patterns of these species to be affected by past and present land use, supporting the hypothesis that these plants accumulate in open habitats. Occurrences of species in this guild in todays’ forest are positively related to past human land use (chapter III) and the density of D. mezereum increases with decreasing distances to forest edges (chapter I). This accumulation may in part be explained by the positive effect of forest edges on reproduction and fruit removal (chapter I). I further found local distribution patterns of this guild and the individual species to be aggregated (chapter I and II), and spatial associations between saplings and reproductive individuals to support the ‘orchard’ hypothesis (chapter II). The aggregated pattern of fruit-bearing individuals was positively related to fruit removal whereas aggregated flowering individuals was negatively related to fruit set (chapter I). On the regional scale, I found these species to occupy climatically suitable areas, or fill their potential ranges, to a less extent that wind dispersed trees and shrubs (chapter IV), which may indicate dispersal limitation.In conclusion, the behaviour of birds and humans have shaped, and still shape the current distribution of fleshy-fruited trees and shrubs in Sweden, resulting in accumulation in open habitats and locally aggregated distribution patterns. Changing land-use practices and potential mismatches between fruit maturation and bird dispersal with a changing climate may thus result in even lower chances of these species to fill their potential ranges, due to habitat losses and dispersal limitations at local and regional scales.
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| 6. |
- Arvanitis, Leena, 1959-
(författare)
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Plant polyploidy and interactions with insect herbivores
- 2007
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- Polyploidization has been suggested to be a common mechanism for plant speciation. Polyploidy is associated with changes in plant traits and altered habitat preference. Antagonistic and mutualistic animals are known to discriminate between plants based on variation in such plant traits, suggesting that interactions may have an important role in divergence of plant polyploids after the polyploidization. In this thesis, I investigated the effect of insect herbivores on divergence of plant polyploids in a system consisting of the predispersal seed predator butterfly Anthocharis cardamines, the bud gall forming midge Dasineura cardaminis, and tetraploids and octoploids of the herb Cardamine pratensis. Octoploid populations occurred more often in shaded and nongrazed habitats than tetraploids. Octoploid plants were larger and had fewer but larger flowers than tetraploids. Butterfly attack rates were higher in tetraploid than in octoploid populations, whereas the gall midge attacked only octoploids. These differences were associated with higher abundance of butterflies in sunny habitats and gall midges in shaded habitats. In contrast to the pattern at the population level, octoploid flower shoots were more likely to be attacked by the butterfly in sympatric populations. Also trait selection differed between ploidy levels, both in the absence and in the presence of herbivores. In a field experiment, butterfly preference did not alter the trait selection in tetraploids. In octoploids, the two herbivores did not change selection considered separately. However, their joint effect resulted in significant selection for smaller flower shoots and reduced selection on number of flowers. This thesis demonstrates that differences in habitat preference and phenotypic plant traits between polyploid cytotypes can lead to altered interactions with herbivores. Such differences in interactions with animals may alter not only the relative fitness of cytotypes but also trait selection within the respective ploidy type.
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| 7. |
- Boalt, Elin, 1976-
(författare)
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Ecology and evolution of tolerance in two cruciferous species
- 2008
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- Tolerance to herbivory is the ability of plants to maintain fitness in spite of damage. The goal of this thesis is to investigate the genetic variation and expression of tolerance within species, determine whether and in what conditions tolerance has negative side-effects, and how tolerance is affected by different ecological factors. Tolerance is investigated with special focus on the effects of different damage types, competitive regimes, history of herbivory, and polyploidization in plants. Studies are conducted as a literature review and three experiments on two cruciferous species Raphanus raphanistrum and Cardamine pratensis.In the tolerance experiments, plants are subjected to artificial damage solely, or in a combination with natural damage. A literature review was conducted in order to investigate the effects of damage method. We found that traits related to tolerance, such as growth and fitness were not as sensitive in regard to damage method as measures of induced chemical traits, or measures of secondary herbivory.Genetic variation of tolerance was demonstrated within populations of R. raphanistrum and between subspecies of C. pratensis. In R. raphanistrum, traits involved in floral display and male fitness were positively associated with plant tolerance to herbivore damage. A potential cost of tolerance was demonstrated as a negative correlation between levels of tolerance in high and low competitive regimes. I found no evidence of other proposed costs of tolerance in terms of highly tolerant plants suffering of reduced fitness in the absence of herbivores or trade-offs in terms of a negative association between tolerance to apical and leaf damage, or between tolerance and competitive ability. In C. pratensis, higher ploidy level in plants involved higher levels of tolerance measured as clonal reproduction. Furthermore, populations exposed to higher levels of herbivory had better tolerance than populations exposed to lower levels of herbivory. In this thesis, I demonstrate evidence of different components for the evolution of tolerance in plants: genotypic variation, selective factors in terms of costs and ploidization, and selective agents in terms of changing environment or herbivore pressure.
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| 8. |
- Bolinder, Kristina, 1987-
(författare)
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Pollen and pollination in Ephedra (Gnetales)
- 2017
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- Ephedra (Gnetales) is a gymnosperm genus with a long evolutionary history; the first dispersed pollen grains with affinity to the group are known already from the Permian. This thesis focuses on the evolutionary history of the group and different aspects of its pollination mechanisms. Despite the limited number of extant species of the genus (50-60), and a low morphological and genetic divergence among species, there is variation in pollination syndrome in the genus. The prevailing state in Ephedra, and most gymnosperms, is wind pollination. It is therefore surprising that one species, E. foeminea, is insect-pollinated. Together with co-workers I documented the pollination syndromes of E. foeminea and a sympatric species, E. distachya, based on long term field experiments in north-eastern Greece and aerodynamic investigations and calculations. Placing the results into an evolutionary framework reveals that the insect-pollinated species E. foeminea is sister to the remaining (mostly wind-pollinated) genus, and indicates that insect pollination is the ancestral state in the Gnetales. During the course of evolution of the group there has been a shift to wind pollination, which may have played a crucial role for the diversification of the crown group in the Paleogene. Pollination biology is often correlated with the morphology of the pollen such that pollen grains of anemophilous plants are small with a smooth surface, whereas pollen grains of entomophilous plants are larger with an ornamented surface and a covering of pollenkitt. The pollen morphology of Ephedra can be broadly divided into two types: an ancestral type with an unbranched pseudosulcus between each pair of plicae, and a derived type with a branched pseudosulcus between each pair of plicae. Further, the pollen morphology and ultrastructure of the pollen wall in Ephedra are to some degree correlated with the pollination syndrome and capability of long distance dispersal. Pollen of E. foeminea has a denser ultrastructure, as a result a higher settling velocity and is therefore capable of flying shorter distances than does pollen of the anemophilous E. distachya, and other investigated anemophilous species that show a more spacious ultrastructure of the pollen grain. These results can be useful in the reconstruction of the pollination mechanism of extinct taxa of the Ephedra-lineage in the future.
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| 9. |
- Christiansen, Ditte Marie, 1990-
(författare)
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Responses of boreal forest understory plant communities to climate and forestry
- 2022
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- A warming climate is altering species distributions and community compositions. To understand and predict changes in species distributions to climate change, we often use species occurrences together with large-scale regional climate data. This can be problematic for several reasons. Species living near the ground experience small-scale spatial variation in temperatures, i.e., microclimate, that are influenced by topography and vegetation and can therefore deviate a lot from regional temperatures. Further, climate often affects species indirectly via species interactions, and such interactions can also change with climate. And last, species may respond slower than climate changes. Ignoring these aspects can complicate our understanding of species-climate relationships.In this thesis, I examined how microclimate and changes in microclimate due to forest management impact performances, interactions, and distributions of plant species in boreal forest understory communities. First, I quantified the importance of microclimate for species performances and distributions. Specifically, I compared the effects of spring temperatures measured on local and regional scales on the population dynamics of a southern forest herb (I). I also tested how small-scale spatial microclimate variation contributed to the regional co-existence of northern and southern understory plant species (II). Second, I examined the role of species interactions in driving abundance patterns of two moss species with different temperature niches across their Swedish ranges by transplanting them separately and together across a climate gradient (III). Lastly, I investigated how understory plant communities respond to changes in microclimate caused by forest management (IV), and how past microclimates influence current patterns of species occurrence, abundance, and reproduction (II).I found that local spring temperatures had a significant effect on the population dynamics of the southern forest herb that could not be detected using regional spring temperatures (I). Spatial variation in microclimate explained the regional co-existence of two northern and two southern species, where the northern species were favoured by cold microclimates and the southern species by warm microclimates (II). In the transplant experiment (III), I found that climate-mediated competition can override the direct effects of climate and limit abundances across ranges. Lastly, I found that microclimate changes caused by forest management activities had a large effect on understory communities (IV), and that current abundances of northern and southern species were partly explained by past microclimate (II).Overall, I demonstrated that, to understand how species (particularly understory plants) respond to climate, we need to replace the standard use of regional climate data with locally measured climate data or down-scaled gridded climate data that account for variation in topography as well as vegetation. To predict how species will respond to climate change, we also need to include species interactions and how these interactions change with a changing climate. Finally, changes in microclimate following changes in forest structure have large effects on understory species. The last finding is important to consider when studying changes in understory communities in a climate context and could be used to mitigate climate effects on forest biodiversity.
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| 10. |
- Greiser, Caroline, 1987-
(författare)
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Microclimate at range margins : Consequences for boreal forest understory species
- 2020
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- A warmer climate will shift species distributional range margins poleward, but near-ground microclimates may modify these shifts. Cold-adapted northern species at their rear edge may survive locally in microrefugia with a colder microclimate, and warm-adapted southern species at their leading edge may colonize stepping stone habitats with a warmer microclimate. However, we do not always know if species ranges are limited by climate and which role microclimate variation plays in modifying range margins. This is especially true for lowland forests, where forest structure and composition have relatively large influences on near-ground microclimates.In this thesis, I explored patterns and drivers of forest microclimate at the southern margin of the boreal zone in central Sweden, where many northern and southern species meet. First, I measured, modelled and mapped near-ground temperatures across ca. 20 000 km2 of forested land (Paper I). Second, I tested if cold and warm microclimates favour northern and southern understory species, respectively. To answer this, I investigated the occurrence and performance patterns of understory vascular plants, bryophytes and lichens across microclimate gradients at the species’ northern or southern range margins (Paper II-IV). I performed both correlational analyses on natural populations and experimental testing with transplanted populations. Third, I derived recommendations and tools for biodiversity conservation and forest management (Paper I-IV).I found high spatial and temporal variation of forest microclimate, which was in the summer mainly linked to differences in forest density and in the cold season to terrain effects (Paper I). Cold and warm microclimates were occupied by natural edge populations of northern and southern species, respectively (Paper II and IV). However, in the transplant experiments with removed competition other factors were more important for the species performance. The southern herb appeared to cope well with the range of microclimate at its current northern range margin and instead seems to be limited by soil and light in northern conifer-dominated forests (Paper IV). The northern transplanted bryophytes and lichens showed no or a positive response to warmer temperature, but also to higher moisture, to more conifers in the overstory and to less gastropod grazing (Paper III). The results indicate that competition with southern species, herbivory, leaf litter and water scarcity might be more important than temperature as direct limiting factors at the species’ current southern range margin. To conclude, microclimate influences the occurrence and performance of range edge populations, but it likely does so indirectly via effects on water availability and biotic interactions.Forest management heavily modifies near-ground temperature and humidity and hence likely impacts the climate-driven range shifts of understory species. I call for considering these effects in conservation and management actions, e.g. by protecting valuable microclimates, moving from clear-cutting to selective logging, reducing forest fragmentation and drainage and favouring either broad-leaved or coniferous trees in the overstory - depending on the local conservation target (Paper I-IV). Climate-change induced biodiversity loss may thus be slowed down by responsible forest management that provides stepping stone habitats for advancing southern species as well as microrefugia for retreating northern species.
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