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Due to massive energetic investments in woody support structures, trees are subject to unique physiological, mechanical, and ecological pressures not experienced by herbaceous plants. Despite a wealth of studies exploring trait relationships across the entire plant kingdom, the dominant traits underpinning these unique aspects of tree form and function remain unclear. Here, by considering 18 functional traits, encompassing leaf, seed, bark, wood, crown, and root characteristics, we quantify the multidimensional relationships in tree trait expression. We find that nearly half of trait variation is captured by two axes: one reflecting leaf economics, the other reflecting tree size and competition for light. Yet these orthogonal axes reveal strong environmental convergence, exhibiting correlated responses to temperature, moisture, and elevation. By subsequently exploring multidimensional trait relationships, we show that the full dimensionality of trait space is captured by eight distinct clusters, each reflecting a unique aspect of tree form and function. Collectively, this work identifies a core set of traits needed to quantify global patterns in functional biodiversity, and it contributes to our fundamental understanding of the functioning of forests worldwide.
Questions: Both species turnover and intraspecific trait variation can affect plant assemblage dynamics along environmental gradients. Here, we asked how community assemblage patterns in relation to species turnover and intraspecific variation differ between endemic and non-endemic species. We hypothesized that endemic species show lower intraspecific variation than non-endemic species because they tend to have high rates of in situ speciation, whereas non-endemic species are expected to have a larger gene pool and higher phenotypic plasticity.
Location: La Palma, Canary Islands.
Methods: We established 44 sampling sites along a directional gradient of precipitation, heat load, soil nitrogen, phosphorus and pH. Along this gradient, we estimated species abundances and measured three traits (plant height, leaf area and leaf thickness) on perennial endemic and non-endemic plant species. In total, we recorded traits for 1,223 plant individuals of 43 species. Subsequently, we calculated community-weighted mean traits to measure the relative contribution of species turnover, intraspecific variation and their covariation along the analysed gradient.
Results: The contribution of intraspecific variation to total variation was similar in endemic and non-endemic assemblages. For plant height, intraspecific variation explained roughly as much variation as species turnover. For leaf area and leaf thickness, intraspecific variation explained almost no variation. Species turnover effects mainly drove trait responses along the environmental gradient, but intraspecific variation was important for responses in leaf area to precipitation.
Conclusions: Despite their distinct evolutionary history, endemic and non-endemic plant assemblages show similar patterns in species turnover and intraspecific variation. Our results indicate that species turnover is the main component of trait variation in the underlying study system. However, intraspecific variation can increase individual species’ fitness in response to precipitation. Overall, our study challenges the theory that intraspecific trait variation is more important for the establishment of non-endemic species compared with endemic species.
Positive plant–plant interactions are thought to drive vegetation patterns in harsh environments, such as semi-arid areas. According to the stress-gradient hypothesis (SGH), the role of positive interactions between species (facilitation) is expected to increase with harshness, predicting associated variation in species composition along environmental gradients. However, the relation between stress and facilitation along environmental gradients is debated. Furthermore, differentiating facilitative interactions from other underlying mechanisms, such as microtopographic heterogeneity, is not trivial. We analysed the spatial co-occurrence relationships of vascular plant species that form patchy vegetation in arid lapilli fields (tephra) from recent volcanic eruptions on La Palma, Canary Islands. Assuming a harshness gradient negatively correlated with elevation because the lower elevations are more arid and water availability is considered the most limiting resource, and that an outcome of facilitation is plants co-occurring in the same patch, from the SGH we expected a greater degree of co-occurrence at lower elevation. We tested this at both the species and the individual plant level. We analysed the species composition of 1277 shrubby vegetation patches at 64 different sampling points, ranging from the coast to around 700 m a.s.l. Patch morphology and microtopographic heterogeneity variables were also measured, to account for their potential effects on the species composition of patches. We used generalized linear models and generalized mixed-effects models to analyse species richness, number of individuals in patches and percentage of patches with positive co-occurrences, and a pairwise co-occurrence analysis combined with a graphical network analysis to reveal positive links between 13 of the species. We found that the percentage of patches with positive co-occurrences increased at higher elevations, in contrast to the predictions of the SGH, but in accordance with a refined stress-gradient hypothesis for arid sites, in which characteristics of the interacting species are incorporated.
Questions: Habitat islands are often characterized by the presence of more or less sharp boundaries to adjacent matrix habitats. However, knowledge on boundaries of natural habitat islands is scarce, especially regarding patterns of beta diversity and its two underlying components: species turnover and nestedness. We therefore aim to quantify the effects of fine-scaled and sharp boundaries of quartz islands (quartz gravel-covered soils) on the different components of plant beta diversity and how they are linked to different soil environmental drivers. Location: Knersvlakte, Western Cape, South Africa. Methods: We sampled plant species richness in 56 fine-scale transects of 6 m × 1 m plots across eight different boundary types (four quartz island to matrix, four between habitats on quartz islands). Soil depth and chemistry (pH, electrical conductivity) were analyzed for each 1 m2 plot. Differences in the two beta diversity components (turnover and nestedness) for each boundary type were tested by t tests. We used linear models to test relationships between species and environmental dissimilarity. Results: All boundary types showed high beta diversity. Species turnover was the prevailing component for six boundary types, the nestedness component was only important for two boundary types. We found a significant linear increase of species dissimilarity with increasing dissimilarity in soil pH and distinct plant communities for the habitat types, but no significant increase for electrical conductivity or soil depth. Conclusions: The spatial distinctiveness of the quartz islands leads to sharp boundaries, which result in high beta diversity, mainly through species turnover. This reflects the high levels of diversification and adaptation of the local plant communities. Nestedness occurred at two boundaries to the matrix, indicating that the latter does not necessarily represent an impermeable boundary for all species of the respective ecosystem. Studying diversity patterns across boundaries contributes to the question of applicability of island biogeography theory to habitat islands.
One of the most important events in human history occurred during the Early Pleistocene: the dispersal of early hominins out of Africa and into Europe and Asia. In Western Europe, the earliest evidences of the genus Homo have been found in the Baza Basin, at the sites of Orce in the SE of the Iberian Peninsula. These sites contain fossils and lithic industry dated approximately as 1.4–1.3 Ma.While hominin remains and artifacts at Orce, as well as the accompanying fauna, have been extensively studied, the properties and evolution of the Early Pleistocene vegetation in the basin remain unknown. The general effect of climate change on the expansion of early hominins from Africa into Eurasia still remains unclear. It is not known if the Early Pleistocene climate changes and the development of glacials periods led to the extirpation of European communities, or if those communities were able to endure and persist through such adverse climatic periods. This open question highlights the need for climate and environmental analyses for the time before, during and after the first presence of Homo in Europe. This PhD thesis contributes to that need by the presentation of the first long pollen record of the Baza Basin, where the oldest hominin sites in Western Europe are found.
Climate controls the broad-scale distribution of vegetation and change in climate will alter the vegetation distribution, biome boundaries, biodiversity, phenology and supply of ecosystem services. A better understanding of the consequences of climate change is required, particularly in under-investigated regions such as tropical Asia, i.e., South and South-east Asia, which is a host to 7 of the 36 global biodiversity hotspots. Conservation strategies would also require an in-depth understanding of the response of vegetation to climate change. Therefore, the main objective of this thesis was to investigate the impact of climate change and rising CO2 vegetation in tropical Asia. Dynamic global vegetation model (DGVMs) are the well-known tools to investigate vegetation-climate interactions and climate change impacts on ecosystems. In this thesis, I used a complex trait-based DGVM called adaptive dynamic vegetation model version 2 (aDGVM2).
In Chapter 1, I presented a brief background of the phytogeography and discussed the exiting knowledge gap on vegetation-climate interactions in the region. One major disadvantage for available DGVMs studies for the tropical Asia is that most of them have used fixed plant functional types (PFTs) and do not explicitly represent the distinct varieties of vegetation type of the region such as Asian savannas. In Chapter 2, I discussed at great length to improve DGVMs for South Asia and discussed ways to include them in the model for better representation of region vegetation-climate interaction.
I upgraded the current version of aDGVM2 and added a new vegetation type i.e., C3 grasses, and modified the sub-module to simulate photosynthesis for each individual plants to aDGVM2. In chapter 3, I used this updated version of aDGVM2 to simulate the current and future vegetation distribution in South Asia under RCP4.5 and RCP8.5 (RCP: representative concentration pathway). The model predicted an increase in biomass, canopy cover, and tree height under the presence of CO2 fertilization, which triggered transitions towards tree-dominated biomes by the end of the 21st century under both RCPs. I found that vegetation along the Western Ghats and the Himalayas are more susceptible to change due to climate change and open biomes such as grassland and savanna are prone to woody encroachment.
In Chapter 4, the study domain was extended to include South-east Asia to verify if the model configuration used in Chapter 3 can also simulate vegetation patterns in tropical Asia. The aDGVM2 simulations showed a robust trend of increasing vegetation biomass and transitions from small deciduous vegetation to taller evergreen vegetation across most of tropical Asia. Shifts in plant phenology also affect ecosystem carbon cycles and ecosystem feedback to climate, yet the quantification of such impacts remains challenging. The study showed increased biomass due to CO2 fertilization, indicates that the region can remain a carbon sink given there is no other resource limitation. However, nutrient limitations on CO2 fertilization effects were not included in the study, and carbon sink potential has to be seen with caution.
In Chapter 5, I focused on Asian savannas, which have been mismanaged since the colonial era due to misinterpretation as a degraded forest. I proposed a biome classification scheme to distinguish between degraded forest or woodland and savanna based on the abundance of grass biomass and canopy cover. I found that considering vegetation systems as woodland or degraded forest could easily be mistaken as a potential for forest restoration within a tree-centric perspective. This would put approximately 35% to 40% of a unique savanna biome at risk. Although projected woody encroachments may imply a transition toward the forest that benefits climate mitigation. This raises potential conflicts of interest between biodiversity conservation in open ecosystems, i.e., savanna and active afforestation, to enhance carbon sequestration. Proper management strategies should be taken into account to maintain a balance for both objective
In conclusion, the model predicted that vegetation in South and South-East Asia would significantly shift towards tree-dominated biomes due to CO2-induced fertilization of C3-photosynthesis. The simulation under fixed CO2 and rising CO2 scenarios clearly showed that rising level of atmospheric CO2 is responsible for most of the predicted change in biome properties. This study is an important step towards understanding ecosystems of South and Southeast Asia, specifically savannas. The aDGVM2 can serve as tools to inform decision making for climate adaptation and mitigation for savanna. The thesis, thus contributes to our ability to improve conservation strategies to mitigate the consequences of climate change.
Aim: Biological invasions are likely determined by species dispersal strategies as well as environmental characteristics of a recipient region, especially climate and human impact. However, the contribution of climatic factors, human impact, and dispersal strategies in driving invasion processes is still controversial and not well embedded in the existing theoretical considerations. Here, we study how climate, species dispersal strategies, and human impact determine plant invasion processes on islands distributed in all major oceans in the context of directional ecological filtering.
Location: Six mountainous, tropical, and subtropical islands in three major oceans: Island of Hawai'i and Maui (Pacific), Tenerife and La Palma (Atlantic), and La Réunion and Socotra (Indian Ocean).
Taxon: Vascular Plants.
Methods: We recorded 360 non-native species in 218 plots along roadside elevational transects covering the major temperature, precipitation and human impact (i.e., road density) gradients of the islands. We collected dispersal strategies for a majority of the recorded species and calculated the environmental niche per species using a hypervolume approach.
Results: Non-native species’ generalism (i.e., mean community niche width) increased with precipitation, elevation and human impact but showed no relationship with temperature. Increasing precipitation led to environmental filtering of non-native species resulting in more generalist species under high precipitation conditions. We found no directional filtering for temperature but an optimum range of most species between 10 and 20°C. Niche widths of non-native species increased with the prevalence of certain dispersal strategies, particularly anemochory and anthropochory.
Main conclusions: Plant invasion on tropical and subtropical islands seems to be mainly driven by precipitation and human impact, while temperature seems to be of little importance. Furthermore, anemochory and anthropochory are dispersal strategies associated with large niche widths of non-native species. Our study allows a more detailed look at the mechanisms behind directional ecological filtering of non-native plant species in non-temperature-limited ecosystems.