Understanding dispersal and gene flow in human-modified landscapes is crucial for effective conservation. Seed dispersal governs colonization, recruitment, and distribution of plant species, whereas both pollen and seed dispersal determine gene flow among populations. This PhD thesis tests the effect of rotational shepherding on seed dispersal and gene flow in fragmented calcareous grasslands. Calcareous grasslands (Gentiano-Koelerietum pyramidatae vegetation) in Central Europe are semi-natural communities traditionally used for rotational grazing that experienced a decline of plant species during the 20th century due to abandonment of shepherding. This PhD profits from a management project started in 1989 in Bavaria, Germany to reconnect previously abandoned calcareous grasslands in three non-overlapping shepherding systems. Two vegetation surveys in 1989 and 2009 revealed colonizations in previously abandoned grasslands reconnected by shepherding. First, I propose a comprehensive approach to identify determinants of community-level patch colonization rates based on 48 habitat specialist plants by testing competing models of pre-dispersal and dispersal effects and accounting for post-dispersal effects. Mean source patch species occupancy in 1989, and structural elements in focal patches related to establishment explained community-level patch colonization rates. Secondly, by adapting the community analysis to all 31 individual species of the same community with sufficient data, I corroborate the role of shepherding to support dispersal for a range of species, even if they lack seed morphological traits related to zoochory. Thirdly, for the habitat specialist Dianthus carthusianorum, I genotyped 1,613 individuals from 64 populations at eleven microsatellites to test the effect of dispersal by sheep on spatial genetic structure at the landscape scale. Genetic distances between grazed patches of the same herding system were related to distance along herding routes, whereas ungrazed patches showed isolation by geographic distance. Lastly, within individual grassland patches, shepherding significantly decreases the degree of relatedness among neighboring individuals (kinship structure) and increases genetic diversity. My thesis contributes towards understanding the effects of zoochory on spatial dynamics in plant populations across scales.

Species distribution models must balance the need for model generality with that for precision and accuracy. This is critical when modelling range-expanding species such as invasive species. Given the increased use of species distribution models to study invasive species-landscape relationships, a better understanding of the effect of spatial scales, sampling biases, model transferability and discrepancies between different models’ future predictions is necessary. This dissertation addresses these knowledge gaps using mute swans (Cygnus olor) as a case study species. I specifically examine mute swan’s distributions in parts of their native range of Britain and their non-native range of Ontario, Canada. I first investigate which environmental variables at which spatial scales best explain mute swan’s distribution in its non-native range. Second, I perform a sample selection bias study to examine predictive accuracy when species distribution models are built using varying ranges of environmental variables and applied to broader spatial extents. Third, I examine the potential for, and limitations of model transferability between native and non-native regions. Finally, I use two different modelling approaches and three different climate change and land use change scenarios to predict future mute swan habitat suitability. The results indicate that (1) models with better predictive accuracy include environmental variables from multiple ecologically-meaningful scales and measured at spatial extents that include a broad range of environmental variable values; (2) models can exhibit asymmetrical transferability; (3) climate change will facilitate mute swan range expansion in the future more than land use change; and (4) mute swans are often found near urban waterbodies. When modelling invasive species distributions, I suggest that ecologists consider: (I) spatial scale of the underlying landscape processes and species’ use of the landscape; (II) variability and range of each environmental variable used for building models; and (III) stage of establishment of the invasive species.

The dynamics of ecological populations and communities are predominantly governed by three ecological processes, environmental filtering, species interactions and dispersal, and these processes may vary with heterogeneity of the environment. In my PhD research, I investigated how ecologists conceptualize landscape heterogeneity, and how these three ecological processes may vary with spatial and temporal environmental heterogeneity. I conducted my empirical work in Alliaria petiolata, a non-native invasive species in North America, at the Koffler Scientific Reserve at Joker’s Hill in Ontario, Canada. The thesis contains six chapters, where chapters 2 – 5 are structured as stand-alone manuscripts. In chapter 2, I conducted a quantitative review to link the metacommunity concept (which combines the above-mentioned three processes) with different conceptual models of landscape spatial heterogeneity. I found that 78% of metacommunity studies were not explicit about the underlying model of landscape heterogeneity, though there was a significant association between the implied model of landscape heterogeneity and the observed metacommunity model. In chapter 3, I quantified dispersal of Alliaria petiolata, assessed the spatial structure of rosette and adult density, and compared the effects of the different processes on rosette and adult density. Seed dispersal followed a lognormal distribution (μ = 0.01, σ = 0.65). Both adults and rosettes exhibited significant spatial structure up to 2 m. Propagule pressure and interactions among life stages were significant processes shaping rosette density, whereas propagule pressure was the only important process shaping adult density. In chapter 4, I investigated patterns, determinants and demographic consequences of herbivory in A. petiolata. I found that patterns, determinants and demographic consequences of herbivory may vary between life stages and habitat types. One striking finding was that herbivory incidence in A. petiolata may strongly depend on plant life stage, possibly due to a defense–fitness trade off. In chapter 5, I tested whether intra-specific interactions in A. petiolata shift with temporal environmental heterogeneity (seasonality). I found significant negative density-dependent survival in summer and positive density-dependent survival over winter. I suggested that predictions of the stress gradient hypothesis at the intra-specific level are applicable to seasonal variation in environmental stress.

Either tropical niche divergence or tropical niche conservatism could drive the latitudinal diversity gradient. Greater niche divergence in the tropics could accelerate reproductive isolation leading to more rapid species formation. Alternatively, latitudinal asymmetry in niche conservatism, whereby tropical species are more conserved than high latitude species, could promote more dispersal in to than out of the tropics, leading to greater tropical richness. Here I test whether rates of climatic niche evolution vary across the latitudinal gradient for 164 closely related pairs of species. Using the evolutionary ages at which sister species diverge, and the niche divergence between them, I applied Brownian motion models to test whether rates of climatic niche evolution varied with latitude. My results indicate that climatic niche conservatism is strongest in the tropics. This suggests that the latitudinal diversity gradient is driven by the inability of tropical to adapt to temperate climates and colonize non-tropical latitudes.

The dynamics of ecological populations and communities are predominantly governed by three ecological processes, environmental filtering, species interactions and dispersal, and these processes may vary with heterogeneity of the environment. In my PhD research, I investigated how ecologists conceptualize landscape heterogeneity, and how these three ecological processes may vary with spatial and temporal environmental heterogeneity. I conducted my empirical work in Alliaria petiolata, a non-native invasive species in North America, at the Koffler Scientific Reserve at Joker’s Hill in Ontario, Canada. The thesis contains six chapters, where chapters 2 – 5 are structured as stand-alone manuscripts. In chapter 2, I conducted a quantitative review to link the metacommunity concept (which combines the above-mentioned three processes) with different conceptual models of landscape spatial heterogeneity. I found that 78% of metacommunity studies were not explicit about the underlying model of landscape heterogeneity, though there was a significant association between the implied model of landscape heterogeneity and the observed metacommunity model. In chapter 3, I quantified dispersal of Alliaria petiolata, assessed the spatial structure of rosette and adult density, and compared the effects of the different processes on rosette and adult density. Seed dispersal followed a lognormal distribution (μ = 0.01, σ = 0.65). Both adults and rosettes exhibited significant spatial structure up to 2 m. Propagule pressure and interactions among life stages were significant processes shaping rosette density, whereas propagule pressure was the only important process shaping adult density. In chapter 4, I investigated patterns, determinants and demographic consequences of herbivory in A. petiolata. I found that patterns, determinants and demographic consequences of herbivory may vary between life stages and habitat types. One striking finding was that herbivory incidence in A. petiolata may strongly depend on plant life stage, possibly due to a defense–fitness trade off. In chapter 5, I tested whether intra-specific interactions in A. petiolata shift with temporal environmental heterogeneity (seasonality). I found significant negative density-dependent survival in summer and positive density-dependent survival over winter. I suggested that predictions of the stress gradient hypothesis at the intra-specific level are applicable to seasonal variation in environmental stress.

Understanding dispersal and gene flow in human-modified landscapes is crucial for effective conservation. Seed dispersal governs colonization, recruitment, and distribution of plant species, whereas both pollen and seed dispersal determine gene flow among populations. This PhD thesis tests the effect of rotational shepherding on seed dispersal and gene flow in fragmented calcareous grasslands. Calcareous grasslands (Gentiano-Koelerietum pyramidatae vegetation) in Central Europe are semi-natural communities traditionally used for rotational grazing that experienced a decline of plant species during the 20th century due to abandonment of shepherding. This PhD profits from a management project started in 1989 in Bavaria, Germany to reconnect previously abandoned calcareous grasslands in three non-overlapping shepherding systems. Two vegetation surveys in 1989 and 2009 revealed colonizations in previously abandoned grasslands reconnected by shepherding. First, I propose a comprehensive approach to identify determinants of community-level patch colonization rates based on 48 habitat specialist plants by testing competing models of pre-dispersal and dispersal effects and accounting for post-dispersal effects. Mean source patch species occupancy in 1989, and structural elements in focal patches related to establishment explained community-level patch colonization rates. Secondly, by adapting the community analysis to all 31 individual species of the same community with sufficient data, I corroborate the role of shepherding to support dispersal for a range of species, even if they lack seed morphological traits related to zoochory. Thirdly, for the habitat specialist Dianthus carthusianorum, I genotyped 1,613 individuals from 64 populations at eleven microsatellites to test the effect of dispersal by sheep on spatial genetic structure at the landscape scale. Genetic distances between grazed patches of the same herding system were related to distance along herding routes, whereas ungrazed patches showed isolation by geographic distance. Lastly, within individual grassland patches, shepherding significantly decreases the degree of relatedness among neighboring individuals (kinship structure) and increases genetic diversity. My thesis contributes towards understanding the effects of zoochory on spatial dynamics in plant populations across scales.

Species distribution models must balance the need for model generality with that for precision and accuracy. This is critical when modelling range-expanding species such as invasive species. Given the increased use of species distribution models to study invasive species-landscape relationships, a better understanding of the effect of spatial scales, sampling biases, model transferability and discrepancies between different models’ future predictions is necessary. This dissertation addresses these knowledge gaps using mute swans (Cygnus olor) as a case study species. I specifically examine mute swan’s distributions in parts of their native range of Britain and their non-native range of Ontario, Canada. I first investigate which environmental variables at which spatial scales best explain mute swan’s distribution in its non-native range. Second, I perform a sample selection bias study to examine predictive accuracy when species distribution models are built using varying ranges of environmental variables and applied to broader spatial extents. Third, I examine the potential for, and limitations of model transferability between native and non-native regions. Finally, I use two different modelling approaches and three different climate change and land use change scenarios to predict future mute swan habitat suitability. The results indicate that (1) models with better predictive accuracy include environmental variables from multiple ecologically-meaningful scales and measured at spatial extents that include a broad range of environmental variable values; (2) models can exhibit asymmetrical transferability; (3) climate change will facilitate mute swan range expansion in the future more than land use change; and (4) mute swans are often found near urban waterbodies. When modelling invasive species distributions, I suggest that ecologists consider: (I) spatial scale of the underlying landscape processes and species’ use of the landscape; (II) variability and range of each environmental variable used for building models; and (III) stage of establishment of the invasive species.

In order to predict future range shifts for invasive species it is important to explore their ability to acclimate to the new environment and understand physiological and reproductive constraints controlling their distribution. My dissertation studied mechanisms by which temperature may affect the distribution of two of the most aggressive plant invaders in North America, Bromus tectorum and Bromus rubens. While Bromus tectorum is dominant in the “cold desert” steppes of the Intermountain region of western North America, B. rubens is one of the severe grass invaders in the “hot deserts” of southwestern North America. I first evaluated whether winter freezing tolerance is the mechanism responsible for the distinct northern range limits of Bromus species. Bromus rubens has a slower rate of freezing acclimation that leads to intolerance of sudden, late-autumn reductions in temperature below -12°C, Bromus tectorum, by contrast, cold hardens rapidly and is not impacted by the sudden severe late-autumn cold. Photosynthetic response to temperature does not explain their current range separation. Bromus species differ little in their photosynthetic temperature responses and the acclimation pattern of photosynthesis. Both species acclimated to a broad range of temperature through the amelioration of Pi regeneration limitation at sub-optimal temperatures and improved carboxylation capacity above the thermal optimum which probably resulted from increased thermostability of Rubisco activase. The effect of elevated temperatures during flowering on the seed yield of Bromus species demonstrates that neither species produces seed at 36°C and above. These thresholds are close to temperatures encountered during flowering in their natural environment. In summary, climatic changes will cause northward range expansion of Bromus species due to less severe autumn and winter, while reproductive failure could cause range contraction at their southern margins.

Saproxylic insects rely on dead or dying wood at some point in their development and appear to be sensitive to forest management. In 2005 and 2006, I sampled saproxylic insects in mixedwood boreal forests in northeastern Ontario to assess effects of: 1) different logging practices, 2) variations in woody debris (WD) qualities, and 3) different sampling methods. I also compared insect communities between heavily decayed WD and soil. Although I collected data on all arthropod orders, I focused on the families of the Diptera and parasitic Hymenoptera and the morphospecies of the Scelionidae and Diapriidae. These insects showed clear responses to the WD quality treatments, but not to the different logging treatments. A few families were affected by ex situ sampling. I also found that the faunas of soil and WD are rather distinct from each other. This study shows that WD is a source of forest biodiversity. Consequently, sustainable forest management will rely on studies like this to conserve and maintain the biodiversity of Canada's largest group of eco-zones.

Tropical forests play a major role in global carbon (C) dynamics and maintain some of the highest biological complexity on Earth; however, little is known about how variation in wood chemical traits contributes to tropical forest structure and function. This research examines inter- and intraspecific patterns in wood chemical traits in order to understand 1) the role wood chemical traits play in tropical forest C dynamics, and 2) the adaptive significance of wood chemical traits in tropical trees. I found wood C concentration varies widely among co-occurring tropical tree species, with average C concentration (47.4 ± 0.33% w/w (S.E.)) being significantly lower than values assumed in prominent forest C accounting protocols. Failing to account for this variation leads to overestimates of ~3.3 – 5.3% in tropical forest C accounting, an error that compounds significantly at larger spatial scales. I also show that oven drying samples prior to elemental analysis underestimates wood C concentration by 2.5 ± 0.17%, due to the loss of the “volatile C fraction”. Counter to expectations, I found wood C concentration is not ii phylogenetically conserved nor correlated to species demography or life history traits. Wood chemical traits showed consistent size-dependent patterns: wood C (in 16 of 24 species) and lignin (in 15 of 16 species) was higher in saplings vs. conspecific canopy trees. These patterns, complimented by phylogenetic analyses, suggest saplings require wood chemical traits that confer greater pathogen defense. When analyzed across a continuous size spectrum, I found wood C concentration (and leaf structural traits) increases linearly, while wood starch concentration (and leaf traits associated with C gain) shows “hump-shaped” patterns with peak values closely preceding reproductive onset; the latter result suggests C may limit growth in larger trees. Overall, my dissertation provides one of the first comprehensive examinations of wood chemical trait variation in tropical trees. In doing so it provides novel, timely, and critical insights into how wood chemical traits contribute to tropical forest structure and function.