published_or_final_version / abstract / Ecology and Biodiversity / Doctoral / Doctor of Philosophy

A major challenge in plant ecology is in understanding how species strategies mediate interactions between the environment and fitness. Variation in niche strategies that affect phenological, physiological, and reproductive traits will allow species to partition resources differently in space and time, allowing for coexistence of many species and strategies within a community. How species differentially respond to variable environments will ultimately influence their population dynamics and geographic distribution. This dissertation approaches this topic from two perspectives: (a) examining the interaction between biogeography and variable demographic strategies in desert annual plants, and (b) examining the costs and benefits of contrasting reproductive strategies in co-occurring selfing and outcrossing desert annuals. Firstly, I tested the abundant center model to determine the role of range position on plant population dynamics. I examined how the geographic and climatic position of 13 desert annuals found at a common location, the Desert Laboratory at Tumamoc Hill in Tucson, Arizona, related to their demography over a 25-year time span. I found that species for which the Desert Laboratory was close to the center of their geographic range have less variable long-term survival and fecundity compared to species for which the Desert Laboratory was further from the center of their range. Secondly, I studied how related species with contrasting mating systems respond to variable environments to affect plant performance. In a three-year field study I investigated how inter-annual variation in plant reproductive phenology affects synchrony with pollinators and herbivores. Since selfing species are guaranteed to reproduce in the absence of pollinators, seasonal and annual variation in phenology resulted in less variable plant reproductive success compared to outcrossing species. Greater variation in reproduction in outcrossing species resulted from asynchrony in some years between plants and pollinators. In a greenhouse study examining the interaction between mating system and drought, I found that the physiological functioning and survival of outcrossing species was more strongly negatively affected by drought conditions, suggesting that selfing species have an advantage in more arid environments. These studies demonstrate how plant reproductive and physiological strategies can play a critical role in influencing fitness, population dynamics and geographic distribution.

2008 – 2009 metais LAMMC SDI obelų pomologiniame sode atlikti introdukuotų obelų veislių su B. 396 poskiepiu tyrimai, tikslu – ištirti biologines ir ūkines savybes. Tyrimų metu nustatyta vaismedžių žydėjimo tarpsniai (žydėjimo pradžia, masinio žydėjimo pradžia ir pabaiga, žydėjimo pabaiga); vaismedžių augumas (medžio aukštis, vainiko skersmuo, kamieno skersmuo, vainiko projekcijos plotas); vaismedžių derlius, vaisių paskirstymas į klases pagal skersmenį, vaisių skynimo laikas, laikymosi pabaiga, vaisiaus masė, kokybė (išvaizda, patrauklumas, bendra kokybė balais), cheminė sudėtis (tirpios sausosios medžiagos, sausosios medžiagos, titruojamasis rūgštingumas), sulčių išeiga, odelės ir minkštimo tvirtumas. Įvertinus tirtų požymių visumą, nustatyta, kad Lietuvos sąlygomis geriausios kokybės vaisius su B.396 poskiepiu išaugina veislės ‘Aldas’, ‘Vanda’, ‘Rosana’ ir ‘Rubinola’ vaismedžiai. Darbo apimtis: puslapiai 39, lentelės 6, paveikslų 12, naudotos literatūros šaltiniai. / The research was carried out at the Institute of Horticulture, Lithuanian Research Centre for Agriculture and Forestry in 2008–2009. The aim of this study was to evaluate the biological and economical characteristics of the introduced apple cultivars on B.396 rootstock. The following characters of apple trees were established: blooming periods (opening of the first blossoms, beginning/end of the full bloom, the end of blooming), vigour of the trees (height, diameter of a crown, diameter of a trunk, area of a crown projection); apple yield, distribution of fruits to classes according to diameter, harvesting time, storage life, fruit size, quality (appearance, taste, general quality evaluation) and chemical composition (soluble solids, dry matter, titratable acidity), juice output, substance of skin and flesh. ‘Aldas’, ‘Vanda’, ‘Rosana’ and ‘Rubinola’ according biological and economical characteristics is the best among other investigated cultivars on B.396 rootstock in Lithuanian climatic conditions. Work size: pages 39, tables 6, pictures 12, references 44.

In the year 2000, croplands covered about 12% of the Earth’s ice-free land surface. Through cropland management, humankind momentarily appropriates about 25% of terrestrial ecosystem productivity. Not only are croplands a key element of human food supply, but also bear potential in increased carbon (C) uptake when best-practice land management approaches are adopted. A detailed assessment of the impact of land use on terrestrial ecosystems can be achieved by modelling, but the simulation of crop C cycling itself is a relatively new discipline. Observational data on crop net ecosystem exchange (NEE) are available only recently, and constitute an important tool for model development, diagnosis, and validation. Before crop functional types (CFT) had been introduced, however, large-scale biogeochemical models (BGCM) lacked crop-specific patterns of phenology, C allocation, and land management. As a consequence, the influence of cropland C cycling on biosphere-atmosphere C exchange seasonality and magnitude is currently poorly known. To date, no regional assessment of crop C cycling and yield formation exists that specifically accounts for spatially and temporally varying patterns of sowing dates within models. In this thesis, I present such an assessment for the first time. In the first step (chapter 2), I built a crop C mass balance model (SPAc) that models crop development and C allocation as a response to ambient meteorological conditions. I compared model outputs against C flux and stock observations of six different sites in Europe, and found a high degree of agreement between simulated and measured fluxes (R2 = 0.83). However, the model tended to overestimate leaf area index (LAI), and underestimate final yield. In a model comparison study (chapter 3), I found in cooperation with further researchers that SPAc best reproduces observed fluxes of C and water (owed to the model’s high temporal and process resolution), but is deficient due to a lack in simulating full crop rotations. I then conducted a detailed diagnosis of SPAc through the assimilation of C fluxes and biometry with the Ensemble Kalman Filter (EnKF, chapter 4), and identified potential model weaknesses in C allocation fractions and plant hydraulics. Further, an overestimation of plant respiration and seasonal leaf thickness variability were evident. Temporal parameter variability as a response to C flux data assimilation (DA) is indicative of ecosystem processes that are resolved in NEE data but are not captured by a model’s structure. Through DA, I gained important insights into model shortcomings in a quantitative way, and highlighted further needs for model improvement and future field studies. Finally, I developed a framework allowing for spatio-temporally resolved simulation of cropland C fluxes under observational constraints on land management and canopy greenness (chapter 5). MODIS (Moderate Resolution Imaging Spectroradiometer) data were assimilated both variationally (for sowing date estimation) and sequentially (for improved model state estimation, using the EnKF) into SPAc. In doing so, I was able to accurately quantify the multiannual (2000-2006) regional C flux and biometry seasonality of maize-soybean crop rotations surrounding the Bondville Ameriflux eddy covariance (EC) site, averaged over 104 pixel locations within the wider area. Results show that MODIS-derived sowing dates and the assimilation of LAI data allow for highly accurate simulations of growing season C cycling at locations for which groundtruth sowing dates are not available. Through quantification of the spatial variability in biometry, NEE, and net biome productivity (NBP), I found that regional patterns of land management are important drivers of agricultural C cycling and major sources of uncertainty if not appropriately accounted for. Observing C cycling at one single field with its individual sowing pattern is not sufficient to constrain large-scale agroecosystem behaviour. Here, I developed a framework that enables modellers to accurately simulate current (i.e. last 10 years) C cycling of major agricultural regions and their contribution to atmospheric CO2 variability. Follow-up studies can provide crucial insights into testing and validating large-scale applications of biogeochemical models.

Over the last millennium peatlands have accumulated significant carbon stores. Drainage for agricultural use has been widespread and has altered the functioning of these mires: shifting them towards carbon release. Recently, in recognition of the range of ecosystem services derived from these landscapes peatland restoration projects have been initiated. Carbon storage is often cited amongst the aims of these projects, especially since the inclusion of rewetting wetlands in the Kyoto Protocol. However, little is known about the effects of ditch blocking on CO2 fluxes, particularly in Molinia caerulea dominated peatlands, a species common on degraded peatlands which tolerates a range of water table depths. This thesis aims firstly to quantify CO2 fluxes from a drained Molinia caerulea dominated blanket bog and to improve understanding of the temporal and spatial controls on these fluxes and secondly, to quantify the immediate effects of ditch blocking. Closed chamber measurements of net ecosystem exchange and partitioned below-ground respiration from control-restored paired sites were collected over the growing seasons immediately pre- (2012) and post-restoration (2013/2014). These flux data were coupled with remotely sensed data quantifying vegetation phenology and structure with a fine resolution (daily/cm) over large extents (annual/catchment). Although temporal variation in water table depth was not related to CO2 fluxes, the seasonal average related to vegetation composition suggesting raising water tables may promote a change in vegetation composition within these species-poor ecosystems. The distribution of water table depths, vegetation composition and CO2 fluxes did not vary with proximity to drainage ditches despite their prominence. An empirical model suggests in a drained state these peatlands are CO2 sources, indicating carbon previously accumulated is gradually being lost. Data suggest restoration does not always significantly affect water tables and consequently CO2 fluxes in the short-term. Where shallower water tables were maintained during dry conditions photosynthesis decreased and heterotrophic respiration increased: enhancing carbon release. Research undertaken during atypical weather has been unable to determine if restoration will be able to raise water tables sufficiently to protect the existing peat store and promote the vegetation change required to reinstate CO2 sequestration in the longer-term.

This study examined aspects of local reproductive variation in the flowering dogwood (Cornus florida L.) coincident with recent differences in primary canopy architecture. The dogwood trees in this study were impacted by a hurricane that created numerous treefall gaps, which created fine scale heterogeneity in the primary canopy. Fine scale disturbances in a forest can result in changes for multiple members of the forest community, including the reproductive responses of the trees and interspecific pollination mutualisms. Previously determined differences in offspring genetic structure suggested that pollen movement among genetically unstructured maternal individuals was significantly impacted between open, or disturbed, canopy and closed, or undisturbed, types. To further understand mechanisms by which this nonrandom mating occurred, this study examined both intrinsic and extrinsic reproductive factors for C. florida. The first chapter examines several parameters of the reproductive phenology of sample groups of dogwood between the canopy types as intrinsic factors. The parameters observed included initiation, course and termination of flowering, seed set and seed germination ratios. The results showed some significant differences between the sampling areas in flowering intensity, although the majority of the variation was between maternal individuals throughout the entire sample. The second chapter examines insect community composition across areas of differential canopy disturbance. Community analysis showed that the open canopy contained a larger and more diverse assemblage of insects than the closed canopy or the field, which represented an area of complete canopy removal. This shift in insect community composition may have created functional differences in the local pattern of pollen flow by altering the functional composition of local potential pollinator assemblages. The major finding of this study was that the impact of intermediate natural disturbance on the mating systems of understory plant populations may be more indirect than direct. In conclusion, the observed differences in insect community structure within these habitat types in this study were consistent with predetermined patterns of pollen flow; this structure can explain some of the previously observed genetic structure within locally proximate understory tree populations.

Afin d’appréhender la réponse des forêts au changement climatique, la phénologie de six espèces d’arbres a été étudiée de 2005 à 2007 à la fois le long d’un gradient altitudinal (fournissant un gradient thermique d’environ 7°C), et en tests de provenances disposés à différentes altitudes. L’objectif était (i) de quantifier les variations phénotypiques des dates de débourrement et de sénescence in situ, (ii) de déterminer les variables climatiques à l’origine de ces patrons, notamment à l’aide de modèles phénologiques, et (iii) d’évaluer la variabilité génétique et la plasticité phénotypique de ces deux événements phénologiques. Nos résultats montrent que la température printanière affecte différentiellement les dates de débourrement des six espèces (de -1.9 jours /°C à -6.5 jours /°C respectivement pour le hêtre et le chêne) mais pas entre les populations d’une espèce. Concernant les dates de sénescence, nous avons mis en évidence que la température induit un fort décalage de cet événement chez le chêne et le hêtre (> 5 jours /°C), alors qu’aucun cline n’est détecté chez l’érable et le frêne. L’allongement de la saison de végétation en réponse à une augmentation de la température est ainsi principalement la conséquence d’une avance des dates de débourrement pour toutes les espèces, à l’exception du hêtre qui présente une plus forte sensibilité pour la sénescence. Les modèles phénologiques utilisés soulignent l’importance des températures printanières affectant les bourgeons en phase de quiescence, tandis que les températures froides hivernales susceptibles de lever la dormance des bourgeons ne semblent pas significativement influencer l’occurrence du débourrement. Concernant la sénescence, les modèles ont mis en évidence le rôle prépondérant de la température pour le chêne sessile et le hêtre, tandis que la photopériode et d’autres facteurs pourraient être impliqués chez le frêne et l’érable. Ainsi, les modèles prédisent que la durée de saison de croissance du chêne va augmenter plus rapidement que celle du hêtre dans les prochaines décennies, et que l’équilibre compétitif entre ces deux espèces en terme phénologique est susceptible d’évoluer vers des altitudes plus élevées. Enfin, nous avons mis en évidence que les différentes populations échantillonnées présentaient de fortes adaptations pour la phénologie et la croissance malgré leur proximité géographique. De plus, les normes de réaction obtenues démontrent, d’une part, l’existence d’une forte plasticité phénologique des espèces, et d’autre part que cette plasticité semble être une caractéristique intrinsèque de l’espèce. Ces résultats révèlent que les arbres ont des capacités adaptatives importantes concernant les traits phénologiques qui pourraient leur permettre, dans une certaine mesure, de faire face au réchauffement du climat. / To assess the response of forests to climate change, the phenology of six tree species was monitored from 2005 to 2007 both along an altitudinal gradient (providing a thermal gradient of about 7 ° C), and in provenance trials at various altitudes. The aim was (i) to characterize phenological patterns of leaf unfolding and leaf senescence timings in situ, (ii) to determine climatic variables responsible for these patterns, in particular using phenological models, and (iii) to assess the genetic variability and phenotypic plasticity of these phenological events. Our results showed that spring temperature differentially affected the leaf unfolding dates of the six species, with significant disparity in responses among species (from -1.9 days / ° C to -6.5 days / ° C for beech and oak, respectively) but not between the populations of a given species. Regarding the dates of senescence, we highlighted that temperature induced a strong shift of this event for oak and beech (> 5 days / ° C), while no cline was detected for sycamore and ash. The lengthening the growing season in response to an increase in temperature is thus mainly the result of an advance in flushing dates for all species except beech, whose growing season length changes were greatly resulting from shifts in senescence. Phenological models stressed the importance of forcing temperatures (effective during bud quiescence period), while the chilling temperatures (effective during dormancy) did not appear to significantly influence the occurrence of leaf unfolding. Concerning senescence variations , the models highlighted the role of temperature for sessile oak and beech, while the photoperiod and other factors could be involved in the ash and maple. Thus, for oak, the models predicted that the length of growing season will extend faster than the one for beech in the coming decades, and that the phenological competitive balance between these two species will likely to evolve towards higher altitudes. Finally, we showed that the sampled populations showed strong adaptations in phenology and growth in spite of their geographical proximity. Moreover, the reaction norms indicated, first, the existence of a strong phenological plasticity of species, and second, that this plasticity seemed to be an intrinsic characteristic of the species. These results underline that trees have consequent inherent adaptive capacities in phenological traits which may enable them to cope with global warming. However, the differences in phenological sensitivities among species suggest that global warming will significantly affect the competitive balance of species.

Master of Arts / Department of Geography / J. M. Shawn Hutchinson / Time series of normalized difference vegetation index (NDVI) data from satellite spectral measurements can be used to characterize and quantify changes in vegetation phenology and explore the role of natural and anthropogenic activities in causing those changes. Several programs and methods exist to process phenometric data from remotely-sensed imagery, including TIMESAT, which extracts seasonality parameters from time-series image data by fitting a smooth function to the series. This smoothing function, however, is dependent upon user-defined input parameter settings which have an unknown amount of influence in shaping the final phenometric estimates. To test this, a sensitivity analysis was conducted using MODIS maximum value composite NDVI time-series data acquired for Fort Riley, Kansas during the period 2001-2012. The phenometric data generated from the different input setting files were compared against that from a base scenario using Pearson and Lin’s Concordance Correlation Analyses. Findings show that small changes to parameter settings results in insignificant differences in phenometric estimates, with the exception of end of season data and growing season length. Next, a time-series analysis of the same MODIS NDVI data for Fort Riley and nearby Konza Prairie Biological Station (KPBS) was conducted to determine if significant differences existed in selected vegetation phenometrics. Phenometrics of interest were estimated using TIMESAT and based on a Savitzky-Golay filter with parameter settings found optimal in the previous study. The phenometrics start of season, end of season, length of season, maximum value, and small seasonal integral were compared using Kolmogorov-Smirnov (K-S) and showed significant differences existed for all phenometrics in the comparison of Fort Riley training areas and KPBS, as well as low- versus high-training intensity areas within Fort Riley. Fort Riley and high-intensity training areas have earlier dates for the start and end of the growing season, shorter growing season lengths, lower maximum NDVI values, and lower small seasonal integrals compared to KPBS and low-intensity training areas, respectively. Evidence was found that establishes a link between military land uses and/or land management practices and observed phenometric differences.

The release of the seed eating beetle Sulcobruchus subsuturalis for biological control of Caesalpinia decapetala in South Africa has been ongoing since 2000. This is the first post release evaluation of the efficacy of the agent against the weed. The study assessed the phenology of C. decapetala and S. subsuturalis at two sites and included determining the establishment of the beetle on the target weed at 25 study sites. In addition, the effects of S. subsuturalis on seed densities as well as seed germination and seedling recruitment were examined. Furthermore predation by native ants and attacks by native parasitoids were also examined at two release sites. Mature filled pods were available on the tree from September (spring) to March (summer) when the beetle is expected to be reproductively active. The majority of beetle releases by Working for water took place in summer. However the proportion of beetle infested seeds, which were only recovered inside pods in the tree canopy at study sites, was low (0 to 15.5%). Consequently high seed densities and seedling recruitment were observed in the field. In the laboratory, S. subsuturalis did not lay eggs on buried seeds, however seed infestation levels were generally high. Only 8.3% of the seeds containing adults germinated and 6.3% emerged into seedlings. Of the seeds containing larvae, only 14.6% germinated and 2.1% emerged into seedlings. Thus far, S. subsuturalis has failed to maintain high populations on the target weed, possibly due to egg predation by native ants (Crematogaster species, Pheidole megacephala, Messor natalensis and Tetramorium avium) and attacks by native parasitoids. Egg predation reached 100% within 10 days. Egg parasitism by unknown parasitoids ranged between 80 and 93.1% in Limpopo. The parasitic wasp Dinarmus altifrons was recovered once developing inside a C. decapetala seed infested with S. subsuturalis. Further investigations regarding attacks by indigenous parasitoids on S. subsuturalis are required to determine the extent of this problem. Meanwhile releases of S. subsuturalis against C. decapetala should continue. However release efforts need to be improved by (a) using inundative rather than inoculative releases (b) Releasing every year between September and March (summer) and (c) involving land owners, farmers and nature reserve authorities in release activities. Ultimately an additional agent should be sought for release against C. decapetala as the weed may not be the preferred host for S. subsuturalis.

The thesis is aimed at climatological, agroclimatological and phenological conditions at Tušimice observatory during 45 years of observations (1968-2012) on basis of meteorological and phenological data by using statistical methods, method of cumulative series and geographical information system. Basic climatic elements (air temperature, precipitation total, snow cover, cloudiness, wind, humidity, sunshine duration, fog and thunderstorm), agroclimatic characteristics according to the WMO recommendation, selected phenological phases (mainly beginning of flowering 10 %) of herb and wooden species were processed in the study. The literature overview was made on climatic and phenological conditions at Tušimice station, in the Czech Republic and other countries. Apart from statistical analysis, the elements were also evaluated in the interaction with the changing air circulation in Central Europe. In addition, results were compared to Quitt's classification based on long-term averages of meteorological data (1961-1990 versus 1981-2010). The drought occurrence was evaluated in 45 years of observations period. The thesis is divided into three main parts - Climatic conditions of Tušimice observatory (part 5.1), Agroclimatic conditions of Tušimice observatory (part 5.2) and Phenological conditions of...