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Myxomycetes or Myxogastria (supergroup Amoebozoa) are one of several Protistean groups dispersing via airborne spores. The model organism for the group, so far exclusively studied in a laboratory environment, is Physarum polycephalum. Here, molecular evolution, distribution and the ecology of spores dispersal was investigated for the non-model species Physarum albescens. This nivicolous myxomycete fruits with snow melt in most mountain ranges of the northern hemisphere and disperses via spherical, dark-colored and melanin-rich spores. Fruit body development and subsequent spore dispersal occurs within a short time window of a few days. At this time, the fruiting plasmodium is fully exposed to the harsh environment if the protecting snow melts away. The spores, with a diameter of 10–13 µm of the typical size for myxomycetes, can potentially reach all suitable habitats worldwide, which led to the assumption that not only Ph. albescens but most myxomycete species should be ubiquitously distributed over the world.
In the first part of this study (article 1), the question was, if spore dispersal can realize a gene flow sufficient to meet the above-mentioned assumption. A total of 324 accessions of Ph. albescens, collected all over the northern hemisphere, was sequenced for 1-3 genetic markers (SSU, EF1A, COI), and 98 specimens were further analyzed using the genotyping by sequencing technique. As a result, at least 18 reproductively isolated units, which can be seen as cryptic biological species, emerged as phylogroups in a three-gene phylogeny, but as well in a SNP-based phylogeny and were confirmed by a recombination analysis between the three markers. However, this evolutive radiation is not simply caused by geographic fragmentation due to low dispersal capability: within a certain region, multiple phylogroups coexisted next to each other, although some appeared to be regional endemics. Most likely, mutations in mating-type genes, as shown to exist for the cultivable Ph. polycephalum, are the main drivers of speciation. This challenges the hypothesis of ubiquitous distribution of Ph. albescens and corroborates the results of the few available studies for other myxomycete species. In addition, groups of clonal specimens, mostly but not always restricted to a certain slope or valley indicated that sexual and asexual reproduction coexists in the natural populations of Ph. albescens.
In the second part (articles 2), the fundamental niche for Ph. albescens was described using all available records for the species. The resulting set of 537 unique occurrence points was subjected to a correlative spatial approach using the software MaxEnt. In dependence on the predictor variables three species distribution models emerged which differed only in details. The first consisted of only 19 bioclimatic variables and an elevation map from the WorldClim dataset. The second was corrected for pseudo-absences resulting from missing survey activities, and the third was expanded with an additional categorical environment variable on snow cover. High mean AUC (area under the curve) values above 0.97 could be reached with all three models. Variables for snow cover, precipitation of the coldest quarter (of the year), and elevation correlated highly to predict the distribution of Ph. albescens. Only in mid-northern latitudes, elevation alone was a good predictor, but it would cause false-positive predictions in arid mountain ranges and failed to explain occurrence in lowland sites at higher latitudes. Mountains in humid climates showed the highest incidences, confirming recent studies that long-lasting snow covers combined with mild summers are crucial for the ecological guild of nivicolous myxomycetes, with Ph. albescens as a typical species.
Spore size is crucial for dispersal ability and should thus be a character under strong selection. In addition, spores carrying two nuclei with opposite mating types should have a colonization advantage. This was the hypothesis for the last part of this study (articles 3 and 4), which investigated this trait in a quantitative manner. This required a method to analyze thousands of spores automatically (article 3) and with high precision for size and the number of nuclei enclosed. Human errors should be excluded, to reveal even subtle differences in the resulting spore size distributions. Two challenges had to be met for this approach. First, a preparation technique was developed to reduce false segmentations due to overlaying spores by aligning spores on one common plane with a high-frequency vibration device. Second, the segmentation process was automated to allow separating spores that are densely packed in the respective images. A machine learning algorithm was set up and trained to reliable identify and measure dark-colored spores. The technique produced consistent results with high accuracy, and the large number of spores allowed to compile spore size distributions, to check for the constancy of this character, which is impossible with manual measurements limited to low numbers.
The resulting spore size distributions, obtained from over 80 specimens (article 4), were mostly narrow, which is in accordance with our hypothesis. Spore size was as well fairly constant within fructifications from one colony. However, mean spore size within different accessions of Ph. albescens showed large variation (ca. 10%, a range often indicated to key out different morphospecies of myxomycetes), and this was explained only by a minor part with differences between biospecies. Not much smaller (8%) was the variation within a group of clonal specimens collected within 25 m distance. This points to a strong influence of environmental factors even at a micro spatial scale, perhaps caused by microclimatic differences and high phenotypic plasticity for spore size. The influence of large-scale covariates like altitude or latitude was negligible. However, spore size correlated with the variance in this trait, indicating that oversized spores may be caused by detrimental environmental conditions. Two aberrations in spore development were found: First, a few specimens showed a multimodal distribution for spore size with two or even three discernible spore populations. The estimated volumes of those populations correspond to a multiple of the first and most abundant conspicuous spore size population. Second, not all spores were uninucleate as to be expected for meiotic products. This was revealed by fluorescence signals from staining the same spores with DAPI, with a second machine learning algorithm trained to identify the nuclei in a spore. A few specimens showed a significant proportion of binucleated spores in the size range of normal-sized ones, and these specimens were not the ones with multimodal spore size distributions. This indicates that the negative impacts (inbreeding) of multinucleate spores should outweigh a possible colonization advantage and is in accordance with the high genetic diversity found in the worldwide population of Ph. albescens, indicating predominantly sexual reproduction in wild populations of myxomycetes.
Forests influence the climate of our Earth and provide habitat and food for many species and resources for human use. These valuable ecosystems are threatened by fast environmental changes caused by human-induced climte change. Negative growth responses and higher tree mortality rates were associated with increasing physiological stress induced by global warming. Especially boreal forests at high latitudes in the arctic region are threatened, a region predicted to undergo the highest increase in temperature during the next decades. Therefore, it is important to assess the adaptation potential in trees. For this purpose, I studied natural populations of white spruce (Picea glauca (Moench) Voss) in Alaska. In this thesis, I present three scientific papers in which my co-authors and I studied the phenotypic plasticity and genetic basis of tree growth, wood anatomy and drought tolerance as well as the genetic structure of white spruce populations in contrasting environments. We established three sites representing two cold-limited treelines and one drought-limited treeline with a paired plot design including one plot located at the treeline and one plot located in a closed-canopy forest, respectively. Additionally, the study design included one forest plot as reference. Within the entire project, in total 3,000 trees were measured, genotyped and dendrochronological data was obtained. I used several approaches to estimate the neutral and adaptive genetic diversity and phenotypic plasticity of white spruce as a model organism to explore the adaptation potential of trees to climate change.
In the first chapter, I combined neutral genetic markers with dendrochronological and climatic data to investigate population structure and individual growth of white spruce. Several individual-based dendrochronological approaches were applied to test the influence of genetic similarity and microenvironment on growth performance. The white spruce populations of the different sites showed high gene flow and high genetic diversity within and low genetic differentiation among populations, rather explained by geographic distance. The individual growth performances showed a high plasticity rather influenced by microenvironment than genetic similarity.
In the second chapter, I investigated the populations of the drought and cold-limited treeline sites to decipher the underlying genetic structure of drought tolerance using different genotype-phenotype association analyses. Based on tree-ring series and climatic data, growth declines caused by drought stress were identified and the individual reaction to the drought stress event was determined. A subset of 458 trees was genotyped, using SNPs in candidate genes and associated with the individual drought response. Most of the associations were revealed by an approach which took into account small-effect size SNPs and their interactions. Populations of the contrasting treelines responded differently to drought stress events. Populations further showed divergent genetic structures associated with drought responsive traits, most of them in the drought-limited site, indicating divergent selection pressure.
In the third chapter, my co-authors and I studied xylem anatomical traits at one of the cold-limited treeline sites to investigate whether genetic or spatial grouping affected the anatomy and growth of white spruce. Annual growth and xylem anatomy were compared between spatial groups and between genetic groups and individuals. Overall, wood traits were rather influenced by spatial than genetic grouping. Genetic effects were only found in earlywood hydraulic diameter and latewood density. Environmental conditions indirectly influenced traits related to water transport.
In conclusion, white spruce showed a high genetic diversity within and a low genetic differentiation among populations influenced by high gene flow rates. Genetic differences among populations are rather caused by geographical distance and therefore genetic drift. Differing selection pressure at the treeline ecotones presumably lead to divergent genetic structures underlying drought-tolerant phenotypes among the populations. Thus, adaptation to drought most likely acts on a local scale and involves small frequency shifts in several interacting genes. The identified genes with adaptive growth traits can be used to further exlore local adaptation in white spruce. Tree growth and wood anatomical traits are rather influenced by the environment than genetics and showed a high phentoypic plasticity. The high genetic diverstiy and phenotypic plasticity of white spruce may help the species to cope with rapid environmental changes. Still, additional work is needed to further explore adaptation processes to estimate how tree species reacted to rapid climate change. The presented thesis shed some light on the adaptation potential of trees by the example of white spruce using several approaches.
A massive shift in agricultural practices over the past decades, to support exceptionally high yields and productivities involving intensive agriculture, have led to unsustainable agriculture practices across the globe. Sustenance of such high yields and productivities demand high use of organic and industrial fertilizers. This acts as a negative pressure on the environment. Excessive use of fertilizers leads to nutrient surplus in the fields, which, as a part of catchment runoff, flows into the water bodies as diffuse pollution. These nutrients through rivers are eventually passed into seas. High nutrients ending up into water bodies cause eutrophication. The situation is worsened when such unsustainable agricultural activities are carried out on drained peatlands. As a result, the nutrients that were not part of the nutrient cycle in the landscape for years begin to leach out due to mineralization of peatlands, thereby putting an additional load of nutrients on the environment, that was already under the negative impact of nutrient surplus. In view of the above, a small lowland catchment of the Ryck river in northeast Germany was assessed for its nitrogen losses from agricultural lands through empirical modelling. Initial empirical modelling resulted in an average annual total nitrogen loss of 14.7 kg ha−1 year−1. After a comparative analysis of these results with procured data, the empirical equation was modified to suit the catchment, yielding more accurate results. The study showed that 75.6% of peatlands in the catchment are under agricultural use. Subsequently, a proposal was made for potential wetland buffer zones in the Ryck catchment. Altogether, 13 peatland sites across 8 sub-catchments were recommended for mitigation of high nutrient runoff. In the end, nutrient efficiency of proposed WBZs in one of the sub-catchments of Ryck has been discussed. The results show that (i) the modified empirical equation can act as a key tool in application-based future strategies for nitrogen reduction in the Ryck catchment, (ii) restoration of peatlands and introduction of WBZs can help in mitigating the nutrient runoff for improved water quality of Ryck, and subsequently (ii) contribute to efficient reduction of riverine loads of nutrients into the Baltic Sea.
Samples of two duckweed species, Spirodela polyrhiza and Lemna minor, were collected around small ponds and investigated concerning the question of whether natural populations of duckweeds constitute a single clone, or whether clonal diversity exists. Amplified fragment length polymorphism was used as a molecular method to distinguish clones of the same species. Possible intraspecific diversity was evaluated by average-linkage clustering. The main criterion to distinguish one clone from another was the 95% significance level of the Jaccard dissimilarity index for replicated samples. Within natural populations of L. minor, significant intraspecific genetic differences were detected. In each of the three small ponds harbouring populations of L. minor, based on twelve samples, between four and nine distinct clones were detected. Natural populations of L. minor consist of a mixture of several clones representing intraspecific biodiversity in an aquatic ecosystem. Moreover, identical distinct clones were discovered in more than one pond, located at a distance of 1 km and 2.4 km from each other. Evidently, fronds of L. minor were transported between these different ponds. The genetic differences for S. polyrhiza, however, were below the error-threshold of the method within a pond to detect distinct clones, but were pronounced between samples of two different ponds.
Logging and sawing of timber using conventional tools by unskilled workers causes enormous damage to the valuable timber, residual stand, regeneration, and forest soil in Nepal. The purpose of this study was to find out the volume reduction factor and identify major strategies to reduce timber losses in the tree harvesting process in the Terai Shorea robusta forest of Nepal. Field measurements and product flow analysis of 51 felled trees from felling coupes and randomly selected 167 sawed logs were examined to study harvesting losses. Responses from 116 forest experts were analyzed to explore strategies for reducing harvesting and processing losses. The results showed that timber losses in the felling and bucking stage with and without stem rot were 23% and 22%, respectively. Similarly, timber losses in the sawing stage with and without stem rot were 31% and 30%, respectively. Paired t-test at 5% level of significance revealed that there was significant loss in both tree felling and log sawing stages with present harvesting practice. The most leading factor contributing to timber loss in all of the three stages was the use of inappropriate equipment during tree harvesting. Use of synthetic ropes for directional felling and skidding as well as flexible and portable sawing machine with size adjustment options during sawing were mainly recommended as strategies to reduce timber losses. This study serves as a baseline study to identify and quantify timber losses in different stages of tree conversion and also formulate their reduction strategies in Nepal.
AbstractArchetype analysis is a promising approach in sustainability science to identify patterns and explain mechanisms shaping the sustainability of social-ecological systems. Although considerable efforts have been devoted to developing quality standards and methodological advances for archetype analysis, archetype validation remains a major challenge. Drawing on the insights from two international workshops on archetype analysis and on broader literature on validity, we propose a framework that identifies and describes six dimensions of validity: conceptual; construct; internal; external; empirical; and application validity. We first discuss the six dimensions in relation to different methodological approaches and purposes of archetype analysis. We then present an operational use of the framework for researchers to assess the validity of archetype analysis and to support sound archetype identification and policy-relevant applications. Finally, we apply our assessment to 18 published archetype analyses, which we use to describe the challenges and insights in validating the different dimensions and suggest ways to holistically improve the validity of identified archetypes. With this, we contribute to more rigorous archetype analyses, helping to develop the potential of the approach for guiding sustainability solutions.
Agriculture in the populated islands of the Galapagos Archipelago, a protected area due to its unique biodiversity, has been detrimental to its conservation but highly required to meet food necessities. A potential solution to make agricultural farming more sustainable is adopting water-saving technologies (WSTs). Therefore, this study aimed to test the effectiveness of using WSTs such as Groasis Waterboxx® in three of the most valuable crops in the islands through participatory research with the involvement of a group of farmers from the Floreana and Santa Cruz islands and explore a possible transition to more sustainable agricultural practices. Capsicum annuum, Cucumis sativus and Solanum lycopersicum were cultivated using Groasis Waterboxx® and compared to conventional irrigation practices (drip-irrigated controls) to assess the variability of productivity, the number of fruits and individual fruit weight (IFW). In addition, differences in plant traits were analyzed by crop, and island. Results suggested that WSTs such as Groasis Waterboxx® may provide on-farm benefits regarding the yields of the studied traits. From this study, it is difficult to determine whether participation in such a research study will permanently change irrigation practices. However, the participant’s responses to the study suggest an increase in their understanding of the use and benefits of WST.
Pollen productivity estimates (PPEs) are a key parameter for quantitative land-cover reconstructions from pollen data. PPEs are commonly estimated using modern pollen-vegetation data sets and the extended R-value (ERV) model. Prominent discrepancies in the existing studies question the reliability of the approach. We here propose an implementation of the ERV model in the R environment for statistical computing, which allows for simplified application and testing. Using simulated pollen-vegetation data sets, we explore sensitivity of ERV application to (1) number of sites, (2) vegetation structure, (3) basin size, (4) noise in the data, and (5) dispersal model selection. The simulations show that noise in the (pollen) data and dispersal model selection are critical factors in ERV application. Pollen count errors imply prominent PPE errors mainly for taxa with low counts, usually low pollen producers. Applied with an unsuited dispersal model, ERV tends to produce wrong PPEs for additional taxa. In a comparison of the still widely applied Prentice model and a Lagrangian stochastic model (LSM), errors are highest for taxa with high and low fall speed of pollen. The errors reflect the too high influence of fall speed in the Prentice model. ERV studies often use local scale pollen data from for example, moss polsters. Describing pollen dispersal on his local scale is particularly complex due to a range of disturbing factors, including differential release height. Considering the importance of the dispersal model in the approach, and the very large uncertainties in dispersal on short distance, we advise to carry out ERV studies with pollen data from open areas or basins that lack local pollen deposition of the taxa of interest.
Changing climate can strongly affect tree growth and forest productivity. The dendrochronological approach to assessing the impact of climate change on tree growth is possible through climate–growth correlation analysis. This study uses an individual tree-based approach to model Pinus wallichiana (P. wallichiana) radial growth response to climate across the physiographic gradients in the lower distributional range of Nepal. This study sampled six sites across the Makwanpur district of central Nepal that varied in elevation and aspect, obtaining 180 tree-ring series. Climate data series were obtained from Climate Research Unit (CRU 4.0). The pair correlation approach was used to assess P. wallichiana growth response to climate and site-level physiographic variables such as site-level environmental stress. The study also determined long-term growth trends across the elevation and aspect gradients. Trees at sites with higher elevation and northeast aspect (NEA) were more responsive to winter and spring precipitation, whereas trees with lower elevation and northwest aspect (NWA) were more responsive to winter and spring precipitation. Basal area increment (BAI) analysis showed the variation of growth at site-level environmental stress, suggesting that the sensitivity of forest ecosystems to changing climate will vary across the lower growth limit of P. wallichiana due to differences in local physiographic conditions.
We studied a pristine, prominently patterned raised bog in Tierra del Fuego, Argentina, to disentangle the complex interactions among plants and water and peat. The studied bog lacks complicating features often posed by other bogs. It is completely dominated by Sphagnum magellanicum, which covers all niches and growth forms, and is joined by only a dozen higher plant species; it is entirely ombrotrophic with very sharp borders to the surrounding fen; it has only one type of peat that shows an only limited range in degree of decomposition; and it is situated in a very even climate with minimal differences in rainfall and temperature over the year. We present detailed measurements along a 498-m-long transect crossing the bog, including water table measurements (n = 498), contiguous vegetation relevés (n = 248), hydraulic conductivity just below the water table (n = 246), and hydraulic conductivity in 11 depth profiles (n = 291); degree of humification of the corresponding peat was assessed in conjunction with the hydraulic conductivity measurements (n = 537). Sphagnum magellanicum moss samples were collected every 2 m along this transect as well and genotyped (n = 242). In addition, along short, 26-m-long transects crossing strings and flarks water table and hydraulic conductivity just below the water table were measured every meter. Sphagnum growth forms were assessed, and the vegetation of the entire bog was mapped in 10 × 10-m relevés (n = 3322). A simulation model was applied to a generalized shape of the bog and produced surface patterns that well matched those seen in the field. The results were integrated with information from the literature and discussed in the framework of a self-regulating and self-organizing raised bog. We identified 19 hydrological feedback mechanisms. We found that the various mechanisms overlap in both space and time, which means there is redundancy in the self-regulation of the system. Raised bogs, when in a natural state, are among the most resilient ecosystems known; resilience that is provided by feedbacks and backup systems to these feedbacks.