Reindeer can have strong effects on the plant species composition and functioning of tundra ecosystems, and often promote a transition towards a graminoid-dominated vegetation type. As a result, they influence many ecological processes, such as nutrient dynamics, soil biotic composition and functioning, and carbon storage. Several studies suggest that the effect of reindeer on vegetation may follow predictable patterns and could induce an alternative stable vegetation state. However, little empirical data on the long-term stability of reindeer effects on vegetation exist, as it is inherently challenging to study these ecological processes experimentally on a sufficiently long timescale. The main objective of this thesis was therefore to gain a better understanding of the long-term ecological processes following reindeer-induced vegetation shifts. In order to gain a more mechanistic insight in what initially drives this transition, I used a field-based grazing simulation experiment in which I separated defoliation, trampling, moss removal and the addition of feces. This allowed me to test the relative contribution of reindeer-related activities to initiating the shift from moss and heath- dominated tundra towards a graminoid-dominated vegetation state. Additionally, I studied the long-term ecological stability following such a vegetation shift. I did this by addressing historical milking grounds (HMGs): sites where high reindeer concentrations associated with historical traditional reindeer herding practices induced a vegetation transition from shrubs towards graminoids several centuries earlier, but which were abandoned a century ago. Studying HMGs allowed me to address: 1. The potential stability of reindeer-induced vegetation shifts; 2. The ecological mechanisms contributing to the long-term stability of these vegetation shifts; and 3. How such long-lasting vegetation changes influence soil carbon- and nutrient cycling. I found that trampling by reindeer is an important mechanism by which reindeer cause vegetation change. Addressing HMGs further revealed that this vegetation change can be hightly persistent, as the studied HMGs showed only a low encroachment at the surrounding borders in the last 50 years. The vegetation in the core areas of all studied HMGs had remained strikingly stable, and were hardly invaded by surrounding shrubs. Interestingly, soil nutrient concentrations and microbial activities were still different from the surrounding area as well, and even comparable to actively grazed areas. Even after many centuries of changed vegetation composition and soil processes, there was no difference in total carbon sequestration. This suggests that the environmental conditions for microbial decomposition were more important than vegetation composition for the soil carbon stocks, in our study site. After studying the contemporary habitat use of HMGs by reindeer and other herbivores, investigating the potential plant-soil feedbacks mechanisms and detailed soil analyses, I concluded that several ecological mechanisms contribute to the long-term stability of HMGs: first, the altered soil biotic and abiotic conditions appear to have a stronger advantage for HMG vegetation than for the surrounding tundra vegetation. Furthermore, I found a clear browsing preference of small rodents on single shrubs proliferating in HMGs, causing a strong limitation on shrub expansion. Moreover, the dense established sward of graminoids likely poses a strong direct competition for space and nutrients, hindering seedling establishment. Finally, I conclude that HMGs are highly stable on relevant ecological timescales, and propose how the concepts of historical contingency and ASS can be applied to understand stability of these reindeer-induced vegetation transitions.

Les interactions « lichen-lichénophage » sont médiées par la présence de métabolites au sein des lichens qui peuvent influer sur les préférences alimentaires des lichénophages. Dans le contexte environnemental du Subantarctique, à faible biodiversité animale et « végétale », une étonnante richesse lichénique a contribué au succès de l’unique Gastéropode terrestre endémique, Notodiscus hookeri. Pour mieux comprendre les mécanismes en jeu, nous avons analysé les préférences alimentaires de cet escargot lichénophage et tenté de cibler les traits-clés des lichens qui modulent l’interaction trophique. À partir d’observations et de prélèvements réalisés sur l’île de la Possession (Archipel Crozet, Terres Australes et Antarctiques Françaises), deux étapes décisives de l’interaction ont été étudiées : (i) le comportement alimentaire de l’escargot en situation de choix nutritionnel, (ii) sa réponse gustative face à des métabolites en situation de non choix. Le broutage préférentiel de certaines parties de lichen a été confronté à l’analyse de la combinaison entre la teneur des métabolites secondaires et celle des métabolites primaires. La localisation des métabolites au sein du thalle de trois espèces lichéniques fréquentes sur l’île et consommées par l’escargot (Argopsis friesiana, Pseudocyphellaria crocata et Usnea taylorii) a été effectuée par l’intermédiaire de techniques chromatographiques et spectrométriques (LC-DAD-MS, GC-MS, microdissection couplée à la LC-DAD-MS, DART-MS) ainsi que par des techniques d’imagerie couplées à la spectrométrie de masse ((MA)LDI-MS). N. hookeri se révèle être une espèce généraliste et opportuniste qui se nourrit de la majorité des lichens auxquels il est confronté. Cependant, il opère des choix en se nourrissant des parties ayant une valeur nutritive élevée et ce, en dépit de la présence de métabolites secondaires potentiellement répulsifs. Plusieurs mécanismes permettant à l’escargot de contrer la toxicité de certains métabolites ont également été décrits dans ce travail. Les choix alimentaires de N. hookeri s’avèrent par conséquent répondre à un compromis entre le bénéfice d’absorption de métabolites appétents à forte valeur nutritive et le coût physiologique (détoxication, clivage, excrétion) engendré par l’ingestion de métabolites répulsifs. En conclusion, trois paramètres-clés au moins seraient à prendre en compte lors des futures études d’interactions entre lichens et lichénophages : il s’agit de la nature et de la teneur des métabolites secondaires, de la teneur en azote ainsi que de la quantité de sucres et de polyols. Les techniques d’imagerie ont fourni une voie nouvelle et prometteuse pour cibler les dégâts opérés par un phytophage. En effet, cette approche permet de mieux apprécier la distribution hétérogène au sein du thalle entre les métabolites. Pour cet escargot terrestre, les ratios de composés « appétents/inappétents » semblent jouer un rôle majeur dans la sélection de nourriture. / Lichen-lichenivore interactions are mediated by the presence of metabolites in lichens, which may impact lichenivore food preferences. In the environmental context of the Subantarctic, with a low animal and “plant” biodiversity, the surprising lichen flora contributed to the success of a the only endemic subantarctic land Gastropod, Notodiscus hookeri. To better understand the snail-lichen interactions, we analyzed the food preferences of this lichenivore snail and tried to pinpoint key lichen traits modulating the trophic interaction. From observations and samplings realized on Possession Island (Crozet Archipelago, Terres Australes et Antarctiques Françaises), two decisive stages of the interaction were studied: (i) the snail feeding behavior in situation of nutritional choice, (ii) its gustatory response when facing isolated metabolites in no-choice situations. Preferential grazing of some lichen parts was linked to both secondary and primary metabolites concentrations. Intrathalline localization of metabolites was performed in three lichen species consumed by the snail and widespread on Possession Island (Argopsis friesiana, Pseudocyphellaria crocata and Usnea taylorii), using chromatographic and spectrometric techniques (LC-DAD-MS, GC-MS, microdissection coupled to LC-DAD-MS, DART-MS) and imaging mass spectrometry ((MA)LDI-MS). N. hookeri appears to be a generalist and opportunistic gastropod, feeding on most lichen species encountered. However, it is able to discriminate between lichen parts, preferentially grazing on parts with the highest nutrient value, in spite of the presence of potential deterrent secondary metabolites. We described several mechanisms helping snails to overcome the toxicity of some lichen metabolites. Snail feeding choices result in trade-off between benefits of absorbing appetent metabolites of high nutritive value and the cost of physiological processes (detoxification, cleavage and excretion) caused by the ingestion of deterrent metabolites. To conclude, three main key parameters might be relevant in future to study the interactions between lichens and lichenivores. They correspond to the nature, the histolocalisation and the content of secondary metabolites balanced by the amount of nitrogen along with sugars and polyols. Imaging techniques have provided a new and promising way to visualize the heterogeneous distribution within the thallus between these metabolites. The ratios of appetent/non appetent compounds are likely to play a major role in food selection by this land snail.

No description available.

No description available.

<p>White-tailed deer (Odocoileus virginianus; hereafter deer) are a keystone herbivore within forest ecosystems. While deer rely on plant species for growth, reproduction, and survival, multiple external factors can dictate browsing behavior. These factors ultimately drive browsing selection, browsing intensity, and diet composition, which in turn can shape the influence deer have on forest ecosystems. To better understand the complex relationship between deer populations, their habitat, and public perception of deer, the Indiana Department of Natural Resources partnered with Purdue University to initiate the Integrated Deer Management Project (IDMP). As part of the IDMP, this dissertation evaluated the ecological condition of deer habitat to assess the influence deer have on woody and herbaceous plant species within Indiana forests. Our study aimed to i) rank woody species according to their selection by deer and evaluate how the ranking of individual species varies across the state (Chapter 2); ii) evaluate variables and spatial extents associated with differences in browsing intensity, and evaluate different indices used to assess differences in browsing intensity (Chapter 3); iii) quantify winter deer diet composition using DNA barcoding to evaluate how diets vary across a gradient of deer densities, browsing intensities, non-native plant densities, and landscape characteristics (Chapter 4); and iv) evaluate the interactive effects of deer, non-native plant species, and landscape characteristics on the herbaceous layer of forests, while further evaluating the spatial extent at which landscape characteristics are most strongly related to herbaceous-layer composition and diversity (Chapter 5). To do this, I sampled 152 woodlots over three years across three regions of Indiana, collecting data on the browsing selection of individual woody species, the browsing intensities on all woody species, and the composition of vegetation communities (Chapters 2, 3, and 5, respectively). To address diet composition (Chapter 4), we collected deer pellet groups to analyzed diet components. We ranked a total of 63 woody species regarding their browsing selection by deer. While most of these remained consistent from region to region, 16 varied greatly in selection, as deer often showed increased selection for a given species when it resided in an area that provided greater browsing opportunities. Browsing intensity was most associated with food availability, however, it was also influenced by deer density in the region with the lowest forest cover. The twig age index of browsing intensity showed promise as the most efficient and effective index for use in Indiana woodlots. Although diet composition did not differ across regions of Indiana, we found 16 that deer consumed several uncommon taxa when the greater landscape exhibited homogenous patch composition. Similarly, deer consumed different native taxa in forested landscapes with greater deer densities in comparison to agricultural landscapes with lower deer densities. Lastly, though browsing varied within and across regions landscape characteristic, and not deer, were the most influential suite of variables. Additionally, the spatial extent at which these variables exhibited their best fit varied depending on the dependent variable being evaluated and the region of analysis. Together, our results highlight that variables ranging from the woodlot to the landscape-scale influence browsing behavior. This showcases that deer respond to variables at varying scales when browsing and in general, browse more in areas that offer the greatest benefit, whether these areas offer greater food availability or quality, or offer lower risks associated with anthropogenic development. This suggests that when managing forests for deer both woodlot and landscape context should play a role in the decision process. Although differences in browsing were observed, deer had less impact on the herbaceous layer compared to other variables we examined. This suggests that, in contemporary forests, landscape characteristics may be the drivers of changes, and species composition may reflect a long-term history of deer herbivory with less variability resulting from differences in contemporary deer abundance within and across regions.  </p>

Knowledge of the impacts of white-tailed deer (Odocoileus virginianus; hereafter deer) as dominant herbivores throughout the Southeastern United States of America is lacking. To address this, three paired experimental units of exclosures and controls were constructed in 2000 on three Wildlife Management Areas across Mississippi within the ecoregions of the Upper Coastal Plain, Lower Coastal Plain, and the Mississippi Alluvial Valley. Vegetation was sampled in the summers of 2000, 2005, and 2021 including vegetation structure, canopy coverage, basal area, and species composition. Additionally, in 2005 and 2021, biomass was sampled to determine potential impacts on nutritional carrying capacity. Among the three study sites, vegetation metrics followed successional trends and were not influenced by herbivory or lack thereof. Additionally, regional differences in nutritional carrying capacity seemed to be of greater importance than herbivory. This research illustrates the difference in effects of deer by region and forest type.

<p dir="ltr">North American beaver are ecosystem and habitat altering mammals with a wide distribution in North America. Beaver are associated with bodies of fresh water while utilizing riparian habitat by foraging on woody and herbaceous, terrestrial and aquatic vegetation, as well as creating lodges, dams, dens, and scent mounds. Chain O’Lakes State Park was chosen as a study area for surveying beaver activity and woody species communities. The aim of the study was to better understand the relationship between beaver, and specifically, the woody vegetation in their foraging and home range. There is a large swath of habitat that beaver can utilize and alter within Chain O’Lakes State Park including the lakes, streams, and forests that attract visitors. The factors that have been found to influence beaver utilization in Chain O’Lakes State Park (and North America at large) are woody species community composition, abundance of heavily preferred woody species, as well as a combination of both an absence of predators and an abundance of aquatic vegetation. Based on the distribution of heavily, occasionally, and rarely preferred woody species across the understory (new individuals) and overstory (mature individuals) stratum an inference can be made that most of the riparian habitat around the lakes will continue to and/or shift into a higher proportion of species that beaver heavily select. A minority of the habitat shall shift away from those species and towards rarely and/or occasionally selected species. I will provide USDA APHIS Wildlife Services that play a wildlife management role in Chain O’Lakes State Park with the survey information detailing how beaver and their structures interact with the ecosystem, hypothesized methods for preserving the woody species that are targeted by beaver foraging within the park, and supplementary information that may aid in maintaining the riparian habitats for the benefit of park’s biodiversity and wildlife persistence while continuing to provide an enriched experience within Northwest Indiana with the potential to inform the greater collection of parks and maintained land at large.</p>

Forest canopies are complex and highly diverse environments. Their diversity is affected by pronounced gradients in abiotic and biotic conditions, including variation in leaf chemistry. We hypothesised that branch-localised defence induction and vertical stratification in mature oaks constitute sources of chemical variation that extend across trophic levels. To test this, we combined manipulation of plant defences, predation monitoring, food-choice trials with herbivores and sampling of herbivore assemblages. Both induction and vertical stratification affected branch chemistry, but the effect of induction was stronger. Induction increased predation in the canopy and reduced herbivory in bioassays. The effects of increased predation affected herbivore assemblages by decreasing their abundance, and indirectly, their richness. In turn, we show that there are multiple factors contributing to variation across canopies. Branch-localised induction, variation between tree individuals and predation may be the ones with particularly strong effects on diverse assemblages of insects in temperate forests.

Grape root borer, Vitacea polistiformis (Harris), is an oligophagous pest of grapevines in the eastern USA. Neonates must burrow into the soil to find grape roots. In Virginia, larvae feed on roots for ~2 years, then pupate just beneath the soil surface. Emerging adults leave an empty pupal exuviae at the soil surface around the vine base. There was no relationship between weekly captures in pheromone traps and pupal exuviae counts, indicating that exuviae sampling is most appropriate to assess infestations. Exuviae sampling in Virginia vineyards revealed infestations that ranged from light to very heavy. Eighteen biotic and abiotic variables were measured and used in analyses that assessed their relative contributions to differences in exuviae density. Water holding capacity and clay/sand ratio were most strongly associated with pupal exuviae density; these variables were used to develop a model for predicting the extent of infestation of individual vineyards. The spatial distribution of pupal exuviae was characterized using non-spatial and geospatial techniques. Although the non-spatial method (Taylor's Power Law) indicated that exuviae showed an aggregated distribution in all blocks, spatial methods (variograms, SADIE) revealed aggregated distributions only in blocks with ≥ 0.5 pupal exuviae per vine. Independent pupal exuviae samples for population assessment in vineyards can be achieved using sampling points separated by >8.8 m. Combined results from geospatial analyses and the temporal distribution of pupal exuviae within years enabled the development of a practical and quantitative sampling protocol. Bioassays used to measure the behavioral response of larvae to host stimuli revealed that neonates were attracted to grape root volatiles. In soil column bioassays, larvae moved vertically and horizontally over distances of up to 120 cm and apparently perceived the presence of grape roots from a distance of 5 cm in soil. Results are discussed in relation to their potential implications for monitoring and managing grape root borer. / Ph. D.

The salicinoids are phenolic glycosides (PGs) characteristic of the Salicaceae family and are known defenses against insect herbivory. Common examples are salicin, salicortin, tremuloidin, and tremulacin, which accumulate to high concentrations in the leaves and bark of willows and poplars. Despite their important role in plant defense, their biosynthetic pathway is not known, although recent work has suggested that benzyl benzoate acts as a possible biosynthetic intermediate. We identified three candidate genes encoding BAHD-type acyltransferases that are predicted to produce benzylated secondary metabolites, named PtACT47, PtACT49, and PtACT54. Expression of PtACT47 and PtACT49 generally correlated with PG content in a variety of tissues and organs of wild type hybrid poplar plants. This correlation was also found in transgenic hybrid poplar where PG content varied with the level of expression of the condensed tannin regulator MYB134 transcript. In these plants, a suppression of PtACT47 and PtACT49 expression was correlated with lower PG content. In contrast, PtACT54 exhibited very low expression in all tissues tested, and this level of expression was not affected in MYB134 plants. In order to better understand their possible biochemical functions, cDNA cloning, heterologous expression, and in vitro functional characterization was performed on these three BAHD acyltransferases. Recombinant PtACT47 exhibited a low substrate selectivity and could utilize acetyl-CoA, benzoyl-CoA, and cinnamoyl-CoA as acyl donors with a variety of alcohols as acyl acceptors. This enzyme showed the greatest Km/Kcat ratio (45.8 nM-1 sec-1) and lowest Km values (45.1 µM) with benzoyl-CoA and salicyl alcohol, and was named benzoyl-CoA:salicyl alcohol O-benzoyltransferase (PtSABT). Recombinant PtACT49 utilized a narrower range of substrates, specifically benzoyl-CoA and acetyl-CoA and a limited number of alcohols. Its highest Km/Kcat (31.8 nM-1 sec-1) and lowest Km (55.3 µM) was observed for benzoyl-CoA and benzyl alcohol, and it was named benzoyl-CoA:benzyl alcohol O-benzoyltransferase (PtBEBT). Both enzymes were also capable of synthesizing plant volatile alcohol esters at trace levels, for example hexenyl benzoate. Recombinant PtACT54 shares low sequence identity with PtSABT (52.3%) and PtBEBT (52.5%) and exhibited only moderate BEBT-like properties. PtSABT and PtBEBT appear to be paralogs based on their high sequence identity (90.6%) and closely related yet distinct biochemical functions. They likely arose from gene duplication and subsequent functional diversification possibly by neofunctionalization. Wounding experiments showed that abiotic damage stimulated the synthesis of specific PGs, notably salicin and salicortin within 24-48hrs. This was accompanied by a proportional increase in the expression of PtSABT and PtBEBT. Furthermore, experiments using transgenic RNAi lines with knock-down suppression of PtBEBT, and PtSABT, and both genes simultaneously, provided the first direct evidence that BAHD acyltransferases are important in PG production. PtSABT suppression, both individually and in the double knock-down suppression, significantly lowered salicortin content, particularly in mature leaves. However, a reduced level of PtBEBT expression did not have a significant effect on the PGs measured. This could indicate that BEBT-like activity may be a shared function among closely related BAHDs. The suppression of multiple BEBT-like genes may be necessary to further delineate their functions. / Graduate / rjchedgy@uvic.ca