Institut für Landschafts- und Pflanzenökologie

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Alien plant fitness is limited by functional trade‐offs rather than a long‐term increase in competitive effects of native communities
(2023) Brendel, Marco R.; Schurr, Frank M.; Sheppard, Christine S.
Alien plants experience novel abiotic conditions and interactions with native communities in the introduced area. Intra‐ and interspecific selection on functional traits in the new environment may lead to increased population growth with time since introduction (residence time). However, selection regimes might differ depending on the invaded habitat. Additionally, in high‐competition habitats, a build‐up of biotic resistance of native species due to accumulation of eco‐evolutionary experience to aliens over time may limit invasion success. We tested if the effect of functional traits and the population dynamics of aliens depends on interspecific competition with native plant communities. We conducted a multi‐species experiment with 40 annual Asteraceae that differ in residence time in Germany. We followed their population growth in monocultures and in interspecific competition with an experienced native community (varying co‐existence times between focals and community). To more robustly test our findings, we used a naïve community that never co‐existed with the focals. We found that high seed mass decreased population growth in monocultures but tended to increase population growth under high interspecific competition. We found no evidence for a build‐up of competition‐mediated biotic resistance by the experienced community over time. Instead, population growth of the focal species was similarly inhibited by the experienced and naïve community. By comparing the effect of experienced and naïve communities on population dynamics over 2 years across a large set of species with a high variation in functional traits and residence time, this study advances the understanding of the long‐term dynamics of plant invasions. In our study system, population growth of alien species was not limited by an increase of competitive effects by native communities (one aspect of biotic resistance) over time. Instead, invasion success of alien plants may be limited because initial spread in low‐competition habitats requires different traits than establishment in high‐competition habitats.
Constant hydraulic supply and ABA dynamics facilitate the trade-offs in water and carbon
(2023) Abdalla, Mohanned; Schweiger, Andreas H.; Berauer, Bernd J.; McAdam, Scott A. M.; Ahmed, Mutez Ali
Carbon-water trade-offs in plants are adjusted through stomatal regulation. Stomatal opening enables carbon uptake and plant growth, whereas plants circumvent drought by closing stomata. The specific effects of leaf position and age on stomatal behavior remain largely unknown, especially under edaphic and atmospheric drought. Here, we compared stomatal conductance (gs) across the canopy of tomato during soil drying. We measured gas exchange, foliage ABA level and soil-plant hydraulics under increasing vapor pressure deficit (VPD). Our results indicate a strong effect of canopy position on stomatal behavior, especially under hydrated soil conditions and relatively low VPD. In wet soil (soil water potential > -50 kPa), upper canopy leaves had the highest gs (0.727 ± 0.154 mol m-2 s-1) and assimilation rate (A; 23.4 ± 3.9 µmol m-2 s-1) compared to the leaves at a medium height of the canopy (gs: 0.159 ± 0.060 mol m2 s-1; A: 15.9 ± 3.8 µmol m-2 s-1). Under increasing VPD (from 1.8 to 2.6 kPa), gs, A and transpiration were initially impacted by leaf position rather than leaf age. However, under high VPD (2.6 kPa), age effect outweighed position effect. The soil-leaf hydraulic conductance was similar in all leaves. Foliage ABA levels increased with rising VPD in mature leaves at medium height (217.56 ± 85 ng g-1 FW) compared to upper canopy leaves (85.36 ± 34 ng g-1 FW). Under soil drought (< -50 kPa), stomata closed in all leaves resulting in no differences in gs across the canopy. We conclude that constant hydraulic supply and ABA dynamics facilitate preferential stomatal behavior and carbon-water trade-offs across the canopy. These findings are fundamental in understanding variations within the canopy, which helps in engineering future crops, especially in the face of climate change.
Density dependence of seed dispersal and fecundity profoundly alters the spread dynamics of plant populations
(2023) Zhu, Jinlei; Lukić, Nataša; Pagel, Jörn; Schurr, Frank M.
Plant population spread has fundamental ecological and evolutionary importance. Both determinants of plant population spread, fecundity and dispersal, can be density‐dependent, which should cause feedback between population densities and spread dynamics. Yet it is poorly understood how density‐dependence affects key characteristics of spread: spread rate at which the location of the furthest forward individual moves, edge depth (the geographical area over which individuals contribute to spread) and population continuity (occupancy of the spreading population). We present a general modelling framework for analysing the effects of density‐dependent fecundity and dispersal on population spread and parameterize this framework with experimental data from a common‐garden experiment using five wind‐dispersed plant species grown at different densities. Our model shows that density‐dependent fecundity and dispersal strongly affect all three population spread characteristics for both exponential and lognormal dispersal kernels. Spread rate and edge depth are strongly correlated but show weaker correlations with population continuity. Positive density‐dependence of fecundity increases all three spread characteristics. Increasingly positive density‐dependence of dispersal increases spread rate and edge depth but generally decreases population continuity. Density‐dependent fecundity and dispersal are largely additive in their effect on spread characteristics. For population continuity, the joint effects of density‐dependent fecundity and dispersal are somewhat contingent on the dispersal kernel. The common‐garden experiment and the experimentally parameterized mechanistic dispersal model revealed density‐dependent fecundity and dispersal across study species. All study species exhibited negatively density‐dependent fecundity, but they differed qualitatively in the density‐dependence of dispersal distance and probability of long‐distance dispersal. The negative density‐dependence of fecundity and dispersal found for three species reinforced each other in reducing spread rate and edge depth. The positively density‐dependent dispersal found for two species markedly increased spread rate and edge depth. Population continuity was hardly affected by population density in all study species except Crepis sancta in which it was strongly reduced by negatively density‐dependent fecundity. Synthesis. Density‐dependent fecundity and seed dispersal profoundly alter population spread. In particular, positively density‐dependent dispersal should promote the spread and genetic diversity of plant populations migrating under climate change but also complicate the control of invasive species.
Differences in mucilage properties and stomatal sensitivity of locally adapted Zea mays in relation with precipitation seasonality and vapour pressure deficit regime of their native environment
(2023) Berauer, Bernd J.; Akale, Asegidew; Schweiger, Andreas H.; Knott, Mathilde; Diehl, Dörte; Wolf, Marc‐Philip; Sawers, Ruairidh J. H.; Ahmed, Mutez A.
With ongoing climate change and the increase in extreme weather events, especially droughts, the challenge of maintaining food security is becoming ever greater. Locally adapted landraces of crops represent a valuable source of adaptation to stressful environments. In the light of future droughts—both by altered soil water supply and increasing atmospheric water demand (vapor pressure deficit [VPD])—plants need to improve their water efficiency. To do so, plants can enhance their access to soil water by improving rhizosphere hydraulic conductivity via the exudation of mucilage. Furthermore, plants can reduce transpirational water loss via stomatal regulation. Although the role of mucilage and stomata regulation on plant water management have been extensively studied, little is known about a possible coordination between root mucilage properties and stomatal sensitivity as well as abiotic drivers shaping the development of drought resistant trait suits within landraces. Mucilage properties and stomatal sensitivity of eight Mexican landraces of Zea mays in contrast with one inbred line were first quantified under controlled conditions and second related to water demand and supply at their respective site of origin. Mucilage physical properties—namely, viscosity, contact angle, and surface tension—differed between the investigated maize varieties. We found strong influences of precipitation seasonality, thus plant water availability, on mucilage production (R2 = .88, p < .01) and mucilage viscosity (R2 = .93, p < .01). Further, stomatal sensitivity to increased atmospheric water demand was related to mucilage viscosity and contact angle, both of which are crucial in determining mucilage's water repellent, thus maladaptive, behavior upon soil drying. The identification of landraces with pre‐adapted suitable trait sets with regard to drought resistance is of utmost importance, for example, trait combinations such as exhibited in one of the here investigated landraces. Our results suggest a strong environmental selective force of seasonality in plant water availability on mucilage properties as well as regulatory stomatal effects to avoid mucilage's maladaptive potential upon drying and likely delay critical levels of hydraulic dysfunction. By this, landraces from highly seasonal climates may exhibit beneficial mucilage and stomatal traits to prolong plant functioning under edaphic drought. These findings may help breeders to efficiently screen for local landraces with pre‐adaptations to drought to ultimately increase crop yield resistance under future climatic variability.
Editorial: Seed behavior in response to extreme environments
(2023) Zhu, Jinlei; Wang, Lei
Effects of biotic interactions on plant fecundity depend on spatial and functional structure of communities and time since disturbance
(2022) Walter, Hanna E.; Pagel, Jörn; Cooksley, Huw; Neu, Alexander; Schleuning, Matthias; Schurr, Frank M.
Biotic interactions in plant communities affect individual fitness and community dynamics. Interactions between plants vary in space, over time and with organisational complexity. Yet it is challenging to quantify temporal, spatial and functional determinants of different types of interactions between long‐lived perennial plant species and their effect on lifetime fecundity. We studied how plant–plant, pollinator‐ and seed predator‐mediated interactions affect year‐to‐year variation in three fecundity components (cone production, seed set and seed survival) during post‐fire recovery. Age‐stratified data on the three fecundity components were collected in 19 even‐aged communities comprising 20 serotinous Protea shrub species in the South African Fynbos. We analyse data on these fecundity components with neighbourhood models to infer the sign and strength of interactions throughout post‐disturbance recovery, the neighbour plant traits that shape them and the spatial scale at which interactions take place. For each fecundity component, these models describe how neighbourhood effects change over time and with spatial distance between plants. We then predicted neighbourhood effects on individual fecundity components and cumulative reproductive output at different post‐fire stand ages for each focal plant. Competitive effects on cone production and seed set increased with post‐fire stand age as biomass and floral resources for pollinators build up. In contrast, neighbourhood effects on seed survival were weak throughout post‐disturbance recovery. Plant–plant interactions were shaped by neighbour traits related to resource acquisition, whereas animal‐mediated interactions depended on neighbour traits related to resource availability for pollinators and seed predators. The spatial scale of the interactions increased from plant–plant over predator‐mediated to pollinator‐mediated interactions. The joint effect of these interactions on cumulative reproductive output caused the proportion of focal plants experiencing competition to increase with time since fire. Synthesis. We show that temporal changes in biotic interactions throughout post‐disturbance recovery of perennial plant communities depend on functional traits and can be integrated into neighbourhood effects on lifetime fecundity. Studying the temporal, spatial and functional determinants of neighbourhood effects on lifetime fecundity is important for predicting not only individual plant fitness but also population and community dynamics in changing environments.
From alien to native Asteraceae : how effects of climate, functional traits, and biotic interactions on population growth change with residence time
(2023) Brendel, Marco; Sheppard, Christine
Biological invasions pose a major threat to native biodiversity and even drive native species to extinction. It is thus of utmost importance to gain a better understanding of limits to population growth and spread of invasive plants. Invasion success in the introduced area is determined by the combined effects of climatic mismatches between the area of origin and the introduced area as well as biotic resistance of resident native communities. Alien plants can respond to environmental selection via changes in functional traits and thereby adapt to novel abiotic conditions. Native community species are expected to adapt to the presence of the invader by gaining eco-evolutionary experience and build-up biotic resistance over time. The aim of this thesis is to investigate interactions of alien plants with the novel abiotic and biotic environment in their introduced range over eco-evolutionary timescales. To this end, I conducted common garden experiments based on an alien-native species continuum to cover a broad range of residence times in Germany (7 to 12,000 years before present). I followed the population growth of 47 annual Asteraceae (including neophytes, archaeophytes, and natives) over two years and measured their performance in intra- and interspecific competition to answer the following questions: 1) How are effects of climatic distances between the area of origin and the introduced area as well as functional traits on population dynamics of alien plants determined by residence time? 2) How is biotic resistance of native communities towards alien plants related to residence time? 3) How are competitive outcomes between single alien and native plants shaped by residence time and serve as a predictor of range sizes? For the first question, I followed population growth of the Asteraceae in monocultures. I calculated climatic distances between the area of origin and the introduced area and measured functional traits in terms of seed mass, maximum height, and specific leaf area. Firstly, I tested whether negative effects of climatic distances on population growth weaken with residence time. Secondly, I investigated trait-demography relationships and tested if functional traits converge towards values that increase population growth. I found a strong effect of seed mass and no effects climatic distances on population growth. A strong negative relationship between seed mass and population growth resulted in directional selection of seed mass towards low values with increasing residence time. For the second question, I measured population growth of the Asteraceae in a Central European grassland community. I tested if competitive effects of the community on the Asteraceae increase with residence time (i.e. co-existence time with the native community). I used a second community native to North American grasslands that never co-existed with the Asteraceae to disentangle competitive effects related to eco-evolutionary experience of the native community from inherent competitive abilities of the Asteraceae. I compared trait-demography relationships in both community types with monocultures and found very similar competitive effects of both communities on the Asteraceae and thus no evidence for a build-up of competition-induced biotic resistance over time. Instead, invasion success was determined by a strong seed-mass-mediated trade-off between population growth in low- vs. high-competition. For the third question, I tested if the response of biomass and seed production of native targets to competition with alien and native neighbours depends on residence time. I tested if competitive effects differ between invasion status groups and explain species’ range sizes in Germany. I generally did not find a higher tolerance of native Asteraceae to competition of neighbouring aliens and natives with increasing residence time. Both established neophytes and natives showed similar competitive abilities and species’ range sizes were not influenced by competitive effects. The detected trait-demography relationships and related directional selection as a mechanism of adaptation to novel abiotic conditions improve the understanding of constraints on population growth and spread of invaders. The lack of interspecific competitive superiority as a determinant of range sizes might indicate that other mechanisms are more important for invasion success. The functional trade-off between population growth in low vs. high competition reveals that invaders that are likely to escape this trade-off should be of highest management concern. By the combination of experimental macroecology with approaches of functional and community ecology used in my study, I strongly advanced the understanding of mechanisms of limits to population growth and spread of alien plants and provide a fundamental basis for future research in invasion ecology.
Functional traits shape plant–plant interactions and recruitment in a hotspot of woody plant diversity
(2023) Cooksley, Huw; Dreyling, Lukas; Esler, Karen J.; Griebenow, Stian; Neumann, Günter; Valentine, Alex; Schleuning, Matthias; Schurr, Frank M.
Understanding and predicting recruitment in species‐rich plant communities requires identifying functional determinants of both density‐independent performance and interactions. In a common‐garden field experiment with 25 species of the woody plant genus Protea, we varied the initial spatial and taxonomic arrangement of seedlings and followed their survival and growth during recruitment. Neighbourhood models quantified how six key functional traits affect density‐independent performance, interaction effects and responses. Trait‐based neighbourhood models accurately predicted individual survival and growth from the initial spatial and functional composition of species‐rich experimental communities. Functional variation among species caused substantial variation in density‐independent survival and growth that was not correlated with interaction effects and responses. Interactions were spatially restricted but had important, predominantly competitive, effects on recruitment. Traits increasing the acquisition of limiting resources (water for survival and soil P for growth) mediated trade‐offs between interaction effects and responses. Moreover, resprouting species had higher survival but reduced growth, likely reinforcing the survival–growth trade‐off in adult plants. Resource acquisition of juvenile plants shapes Protea community dynamics with acquisitive species with strong competitive effects suffering more from competition. Together with functional determinants of density‐independent performance, this makes recruitment remarkably predictable, which is critical for efficient restoration and near‐term ecological forecasts of species‐rich communities.
Improving measurements of the falling trajectory and terminal velocity of wind‐dispersed seeds
(2022) Zhu, Jinlei; Buchmann, Carsten M.; Schurr, Frank M.
Seed dispersal by wind is one of the most important dispersal mechanisms in plants. The key seed trait affecting seed dispersal by wind is the effective terminal velocity (hereafter “terminal velocity”, Vt), the maximum falling speed of a seed in still air. Accurate estimates of Vt are crucial for predicting intra‐ and interspecific variation in seed dispersal ability. However, existing methods produce biased estimates of Vt for slow‐ or fast‐falling seeds, fragile seeds, and seeds with complex falling trajectories. We present a new video‐based method that estimates the falling trajectory and Vt of wind‐dispersed seeds. The design involves a mirror that enables a camera to simultaneously record a falling seed from two perspectives. Automated image analysis then determines three‐dimensional seed trajectories at high temporal resolution. To these trajectories, we fit a physical model of free fall with air resistance to estimate Vt. We validated this method by comparing the estimated Vt of spheres of different diameters and materials to theoretical expectations and by comparing the estimated Vt of seeds to measurements in a vertical wind tunnel. Vt estimates closely match theoretical expectations for spheres and vertical wind tunnel measurements for seeds. However, our Vt estimates for fast‐falling seeds are markedly higher than those in an existing trait database. This discrepancy seems to arise because previous estimates inadequately accounted for seed acceleration. The presented method yields accurate, efficient, and affordable estimates of the three‐dimensional falling trajectory and terminal velocity for a wide range of seed types. The method should thus advance the understanding and prediction of wind‐driven seed dispersal.
Die Makrophyten-Vegetation ausgesuchter Fließgewässer des Vorarlberger Rheintals als Grundlage für die Bewertung des ökologischen Zustands
(2005) Jäger, Dietmar; Kohler, Alexander
During the 2002 and 2003 growing seasons, total macrophyte growth was thoroughly investigated in 24 selected stretches of running water in the Vorarlberg Rhine valley over a near-continuous distance of 75 km. Quantities of each species were recorded using the KOHLER evaluation method (KOHLER 1978; KOHLER & JANAUER 1995) in a total of 461 sections of running water and quantitative parameters for macrophyte vegetation were determined using the method defined by the authors JANAUER et al. (1993), KOHLER & JANAUER (1995) und PALL & JANAUER (1995). objectives of this study: 1. Recording of the macrophyte population in selected stretches of running water in the Vorarlberg Rhine valley, and the representation of distribution models for each stretch of water investigated 2. Observation and evaluation of relationships between the occurrence of individual species and relevant site factors in the area investigated, based upon data from the literature 3. Investigation of the indicator potential of macrophytes in the area investigated for specific pollution sources 4. Characteristic definition of running waters on the basis of floristic and ecological factors 5. Proposed regional model species for water restoration 6. Evaluation of the regional threat to individual species
Plant stress memory within and across generations
(2022) Lukić, Nataša; Schurr, Frank
Im Zuge des Klimawandels werden Trockenheit und Staunässe in vielen Teilen der Welt immer häufiger und intensiver auftreten. Diese Stressfaktoren beeinträchtigen das Pflanzenwachstum, indem sie die Photosynthese hemmen und oxidative Zellschäden verursachen. Eine vielversprechende Strategie von Pflanzen, um mit solchen schädlichen Bedingungen fertig zu werden, ist das pflanzliche Stressgedächtnis. Dies ist die Fähigkeit von Pflanzen, frühere Stressinformationen zu speichern, um als Reaktion auf wiederholten Stress in derselben Generation (Gedächtnis innerhalb einer Generation) oder in der nächsten Generation (transgenerationales Gedächtnis) eine Verbesserung der Pflanzenleistung auszulösen. Pflanzen verfügen auch über ein Cross-Stress-Gedächtnis; die Konfrontation mit einer Art von Stress, erhöht die Toleranz gegenüber eine anderen Stressart. Darüber hinaus könnte das Stressgedächtnis von Pflanzen bei Arten und Individuen mit eingeschränkter Ausbreitung stärker ausgeprägt sein, da eine eingeschränkte Ausbreitung die Kongruenz zwischen der Umgebung der Mutter und ihrer Nachkommen vergrößern könnte. Die Mechanismen, die dem Gedächtnis für Trockenheit und Staunässe und dem möglichen Cross-Stress-Gedächtnis innerhalb einzelner Generationen oder bei Nachkommen zugrunde liegen, sind derzeit unklar. Außerdem gibt es nur wenige Studien, die die Stärke des Stressgedächtnisses von Pflanzen untersuchen. In dieser Doktorarbeit stellte ich die Hypothese auf, dass Trockenheit und Staunässe zur Bildung eines Stressgedächtnisses führen, das die Leistung der Pflanzen verbessert und Veränderungen der morphologischen, photosynthetischen und antioxidativen Parameter bei wiederholter Trockenheit innerhalb derselben Generation bewirkt. Um diese Hypothese zu testen, habe ich Alopecurus pratensis über zwei Jahre hinweg wiederholt Staunässe und Trockenheit ausgesetzt. Im dritten Jahr wurden die Pflanzen ann zwei Wochen lang Trockenstress ausgesetzt. Meine Ergebnisse bestätigten die erste Hypothese, dass Pflanzen, die zuvor Trockenheit ausgesetzt waren, nach wiederholtem Trockenstress in derselben Generation weniger Gewebeschäden und einen höheren Rubisco-Gehalt, und Gehalt an antioxidativen Enzymen (POX und SOD) und Chlorophyll b aufwiesen. Gräser weisen ein langfristiges Trockenstressgedächtnis über mehrere Wochen hinweg auf und dies hängt mit dem anti-oxidativen System zusammen. Darüber hinaus stellte ich die Hypothese auf, dass Nachkommen, die Stress erfahren, besser abschneiden, wenn die Mütter zuvor Stress erfahren haben, unabhängig von der Art des Stresses. Um dies zu testen, führte ich ein voll-faktorielles Topfexperiment über zwei xvi Generationen durch, bei dem Mutter- und Nachkommenpflanzen Staunässe und Trockenheit ausgesetzt wurden. Im Einklang mit meiner zweiten Hypothese erhöhte die Erfahrung der Mutter mit Staunässe und Trockenheit bei vier mehrjährigen Arten die Biomasse und die Reproduktionsleistung der Nachkommen, die denselben Stressbedingungen ausgesetzt waren wie ihre Mütter. Dieses transgenerationale Gedächtnis wurde mit Veränderungen im antioxidativen System einer Art in Verbindung gebracht, die oxidative Schäden durch die Hochregulierung von schützenden Enzymen bei den Nachkommen verringerten, die denselben Bedingungen ausgesetzt waren wie ihre Mütter. Allerdings konnte ich in dem Experiment nur eine Erinnerung an die gleiche Belastung, aber keine Erinnerung an eine andere Belastung feststellen (keine Cross-Stress-Toleranz). Schließlich stellte ich die Hypothese auf, dass die Stärke der adaptiven mütterlichen Effekte bei Arten oder einzelnen Samen mit geringerer Fähigkeit zur Samenverbreitung zunimmt. Um die dritte Hypothese zu prüfen, wurden von den unteren und oberen Teilen jeder Mutterpflanze Samenköpfe gesammelt, um die Stärke des transgenerationalen Gedächtnisses zu testen, und für die Berechnung der Ausbreitungsdistanz verwendet. Die Nachkommen der unteren und oberen Teile der Mutterpflanzen wurden denselben Wasserbehandlungen ausgesetzt wie die Mütter. Im dritten Experiment konnten die Mutterpflanzen ihre Nachkommen nicht auf den bevorstehenden Wasserstress vorbereiten. Auch eine mütterliche Stresserfahrung als solche konditionierte die Nachkommen nicht auf andere Arten von Stress. Außerdem zeigten Samen mit einer längeren erwarteten Ausbreitungsdistanz entgegen meinen Erwartungen einen stärkeren mütterlichen Anpassungseffekt, wenn sie demselben Wasserstress ausgesetzt waren wie ihre Mütter. Meine Forschung liefert Beweise für ein Trockenheitsgedächtnis innerhalb von Generationen und über Generationen hinweg und stellt eine Verbindung zu den zugrunde liegenden photosynthetischen und redoxbezogenen Mechanismen her. Darüber hinaus wurde die transgenerationale Staunässe mit einem ähnlichen Mechanismus in Verbindung gebracht. Dies könnte zu den schnellen Reaktionen der Pflanzen auf Umweltveränderungen beitragen. Wie ich jedoch in dieser Arbeit gezeigt habe, ist das pflanzliche Stressgedächtnis nicht bei allen Pflanzenarten vorhanden. In Zukunft wird es wichtig sein, die Ursache für die ökologisch bedeutsamen Diskrepanzen bei den mütterlichen Effekten zwischen den Arten zu untersuchen. Unterschiedliche mütterliche Effekte zwischen den Arten könnten die Artenvielfalt erhöhen. Ein umfassenderes Verständnis des transgenerationalen Gedächtnisses könnte dazu beitragen, die Strategien zur Verbesserung von Pflanzen für wirtschaftlich und ökologisch wichtige Arten anzupassen.
Seed dispersal by wind decreases when plants are water‐stressed, potentially counteracting species coexistence and niche evolution
(2021) Zhu, Jinlei; Lukić, Nataša; Rajtschan, Verena; Walter, Julia; Schurr, Frank M.
Hydrology is a major environmental factor determining plant fitness, and hydrological niche segregation (HNS) has been widely used to explain species coexistence. Nevertheless, the distribution of plant species along hydrological gradients does not only depend on their hydrological niches but also depend on their seed dispersal, with dispersal either weakening or reinforcing the effects of HNS on coexistence. However, it is poorly understood how seed dispersal responds to hydrological conditions. To close this gap, we conducted a common‐garden experiment exposing five wind‐dispersed plant species (Bellis perennis, Chenopodium album, Crepis sancta, Hypochaeris glabra, and Hypochaeris radicata) to different hydrological conditions. We quantified the effects of hydrological conditions on seed production and dispersal traits, and simulated seed dispersal distances with a mechanistic dispersal model. We found species‐specific responses of seed production, seed dispersal traits, and predicted dispersal distances to hydrological conditions. Despite these species‐specific responses, there was a general positive relationship between seed production and dispersal distance: Plants growing in favorable hydrological conditions not only produce more seeds but also disperse them over longer distances. This arises mostly because plants growing in favorable environments grow taller and thus disperse their seeds over longer distances. We postulate that the positive relationship between seed production and dispersal may reduce the concentration of each species to the environments favorable for it, thus counteracting species coexistence. Moreover, the resulting asymmetrical gene flow from favorable to stressful habitats may slow down the microevolution of hydrological niches, causing evolutionary niche conservatism. Accounting for context‐dependent seed dispersal should thus improve ecological and evolutionary models for the spatial dynamics of plant populations and communities.
Spatial and functional determinants of long-term fecundity in serotinous shrub communities
(2016) Nottebrock, Henning; Schurr, Frank
Understanding the dynamics of biological communities is a central aim of ecological research. Contemporary environmental change reinforced this interest: in order to predict how communities will react to environmental change, we have to understand the processes driving their dynamics. Ultimately, the dynamics of a community depends on the reproduction, mortality and dispersal of its component individuals, and on how these demographic processes are altered by environmental factors and biotic interactions. A general understanding of biological communities is unlikely to arise from a species-specific approach that attempts to quantify all pairwise interactions between species. Instead, it seems promising to pursue a trait-based research program that quantifies how variation in the performance of species and individuals is shaped by the interplay of functional traits, biotic interactions and environmental factors. In this thesis, I investigated how functional plant traits determine plant-plant, plant-pollinator and plant-herbivore interactions in space and time, and how these spatiotemporal interactions affect the long-term fecundity of plants. In the South African Fynbos biome (a global biodiversity hotspot), I studied a species-rich, ecologically and economically important group of woody plants (genus Protea) and its interactions with pollinators and seed predators. The objectives of this thesis were: (1) to combine plant traits and high-resolution maps of Protea communities in order to quantify the landscapes of nectar sugar and seed crops that plant communities provide for pollinators and seed predators, (2) to examine how sugar landscapes shape pollinator behaviour, and how pollinator behaviour and pollinator-mediated interactions between plants affect the reproduction of Protea individuals, (3) to study how the spatial structure of plant communities and seed crop landscapes determine direct and predator-mediated interactions between plants, and (4) to understand how the interplay of these biotic interactions shapes the dynamics of plant communities. I addressed these objectives by analysing spatially-explicit data and high-resolution maps from 27 sites of 4 ha each that contained 129,750 plants of 22 Protea species. The results show that Protea plants and their pollinators interact on several spatial and temporal scales, and that these interactions are shaped by sugar landscapes. Within plants, inflorescences compete for pollination. At a neighbourhood scale, Protea reproduction benefits from nectar sugar of conspecific neighbours but not from heterospecific neighbour sugar. Seed set also increases with the amount of nectar sugar at the scale of entire study sites. This corresponds with the finding that the abundance and the visitation rates of key bird pollinators strongly depend on phenological variation of site-scale sugar amounts. Nectar sugar furthermore influences the strength of interactions between Protea species and bird pollinators: Protea species that provide nectar of high sugar concentration depend more strongly on bird pollinators to reproduce. When foraging in sugar landscapes, these bird pollinators show both temporal specialization on single plant species and a preference for common plant species. In addition to these pollinator-mediated interactions, the long-term fecundity of Protea individuals is reduced through both competition and apparent competition mediated by seed predators. Competition is stronger between conspecifics than between heterospecifics, whereas apparent competition shows no such differentiation. The intensity of competition between plants depends on their size and the intensity of apparent competition between plants depends on their seed crops. Moreover, competition has a stronger effect on plant fecundity than apparent competition. These findings have interesting implications for understanding the dynamics of Protea communities and the maintenance of plant diversity in the Fynbos biome. The positive interspecific density-dependence resulting from pollinator-mediated interactions causes community-level Allee effects that may lead to extinction cascades. My analyses also imply that competition stabilizes the coexistence of Protea species (because intraspecific competition is stronger than interspecific competition), whereas apparent competition via seed predators does not have such a stabilizing effect. In summary, this study highlights the benefits of ‘community demography’, the demographic study of multiple interacting species. Community demographic studies have the potential to identify general determinants of biotic interactions that act across species and communities. In this thesis, I identified nectar sugar and seed crops as interaction currencies that determine how multiple plant species interact through shared pollinators and seed predators. In megadiverse systems such as Fynbos, such generalizations are urgently needed to understand and forecast community dynamics. The analysis of community dynamics with respect to such interaction currencies provides an alternative to the classical species-specific approach in community ecology.

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