This thesis quantifies aspects of shade tolerance in tree seedlings of species native to the Caledonian pinewood ecosystems of Glen Affric (Highland Region, Inverness-shire). Growth, allocation and morphological responses of 15 species to irradiance under simulated forest canopy light were investigated in a nursery-based shade house experiment. The same responses of four of the 15 species (Ilex aquifolium, Alnus glutinosa, Sorbus aucuparia and Betula pubescens) to different developmental stages of Pinus sylvestris woodland were investigated in the field. The spatial and temporal growth responses of naturally regenerating S. aucuparia seedlings to shade and gap microhabitats were also studied. Data from the shade house experiment enabled further detailed exploration of the relationship between relative growth rates (RGR) and irradiance and potential cross-overs of ranks of growth in high and low light conditions.

Abrupt and persistent reconfiguration of ecosystem’s structure and function has been observed on a wide variety of ecosystems worldwide. While scientist believe that such phenomena could become more common and severe in the near future, little is known about the patterns of regime shifts’ causes and consequences for human well-being. This thesis aims to assess global patterns of regime shifts in social-ecological systems. A framework for comparing regime shifts has been developed as well as a public forum for discussing knowledge about regime shifts, namely the regime shift database. The most common drivers and expected impacts on ecosystem services have been identified by studying the qualitative topology of causal networks as well as the statistical properties that explain their emergent patters. Given that long time series data for ecosystems monitoring is rather sparse, and experimenting with ecosystems at the scales required to understand their feedback dynamics is rarely an option; we also proposed an indirect computationally based method for monitoring changes in ecosystem services. I hope the results here presented offer useful guidance for managers and policy makers on how to prioritize drivers or impacts of regime shifts: one take home message is that well-understood variables are not necessary the ones where most managerial efforts need to be taken. I also hope the scientific community rigorously criticize our results, but also acknowledge that when doing theoretical or empirical work, our methods tend to ignore the multi-causal nature of regime shifts. By bringing back multi-causality to the scientific debate, I hope our results offer new avenues for hypothesis exploration and theory development on the human endeavour of understanding Nature. / Transiciones críticas o cambios de régimen en ecosistemas se definen como reconfiguraciones abruptas de su estructura y función. Estos cambios, en ocasiones inesperados, se han documentado en una gran variedad de ecosistemas en todo el planeta. Algunos científicos proponen que en el futuro cercano dichos fenómenos pueden volverse más frecuentes y severos. Sin embargo, sabemos muy poco sobre las causas y consecuencias potenciales para el bienestar humano. El objetivo de esta tesis es evaluar patrones globales de cambios de régimen en sistemas socio-ecológicos. Un marco conceptual para comparar cambios de régimen y un foro público de discusión sobre el estado del arte en su conocimiento fue desarrollado en la base de datos virtual www.regimeshifts.org. Las causas más comunes y los impactos en servicios ecosistémicos más esperados han sido identificados estudiando las propiedades topológicas de redes causales, así como las propiedades estadísticas que explican sus propiedades emergentes. Dado que experimentar con ecosistemas a la escala adecuada para capturar sus mecanismos causales generalmente no es una opción, y dado que la disponibilidad de datos de largo plazo necesarios para monitorear cambios de régimen son la excepción y no la regla, proponemos un método indirecto computacional para monitorear cambios en servicios ecosistémicos. Espero que los resultados sean de utilidad para actores encargados del diseño de políticas o del manejo de ecosistemas, especialmente espero que ofrezcan una guía sobre cómo priorizar causas y consecuencias de estos cambios de régimen: una lección clave es que las variables que mejor entendemos o las que más monitoreamos no son necesariamente aquellas en las que debemos enfocar las estrategias de manejo. También espero que la comunidad científica critique con rigor nuestros resultados, pero a su vez reconozca que tanto el trabajo empírico y teórico como los métodos que comúnmente se utilizan para estudiar cambios de régimen tienden a ignorar su naturaleza multi-causal. Al enfatizar la diversidad de sus causas, espero que los resultados ofrezcan nuevas posibilidades para la exploración de hipótesis y el desarrollo de teorías para entender mejor la Naturaleza. / Abrupt och ihållande omkonfigurering av ekosystems struktur och funktion har observerats i en mängd olika ekosystem världen över. Forskning visar på att dessa fenomen antas bli vanligare och allvarligare inom vår närmsta framtid. Kunskapen kring dessa s.k. regimskiften är dock bristfällig, framförallt kring dess konsekvenser för mänskligt välbefinnande. Denna avhandling syftar till att bedöma globala mönster av regimskiften. Ett ramverk för att jämföra regimskiften, samt ett offentligt forum, “the regime shifts database”, för att främja diskussion och sprida kunskap om regimskiften, har utvecklats. De mest förekommande drivkrafter och effekter på ekosystemtjänster har identifierats genom att studera kvalitativa topologiska och kausala nätverk, samt de statistiska egenskaperna som förklarar deras framväxande mönster. Då långvariga tidsserier av ekosystemövervakning är få, och då de experiment som krävs för att förstå regimskiftens återkopplingsdynamik sällan är möjliga, föreslås också en indirekt beräkningsmetod för övervakning av förändringar i ekosystemtjänster. Resultaten från denna avhandling ämnar ger värdefull vägledning för beslutsfattare om prioriteringsordningen mellan olika typer av drivkrafter och effekter av regimskiften. En viktig slutsats är att gedigen kunskap om en viss variabel inte nödvändigtvis ger området där insatser bör tillsättas. Vidare, genom att föra tillbaka multi-kausalitet till den vetenskapliga debatten, erbjuder avhandlingen nya vägar för hypotesprövning och teoriutveckling inom vår gemensamma strävan att förstå Naturen. / <p>At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 1: Manuscript. Paper 2: Manuscript. Paper 4: Manuscript.</p>

As human activity along coastlines increase, degradation and destruction of coastal marine ecosystems around the globe will increase at an alarming rate. In an effort to mitigate degradation and destruction of coastal marine ecosystems, artificial reefs have been used in restoration and enhancement projects. As artificial reefs are the main method of restoring diversity to a degraded area, it is important to know the mechanisms that drive marine community assembly and diversity on those reefs. Understanding community assembly patterns of foundational species, in particular, may provide insight to community assembly patterns at higher trophic levels. Subtidal macroalgae are commonly seen as foundational species in marine environments and both deterministic and stochastic processes play a role in their assembly. Environmental gradients, which are deterministic processes, play a significant role in structuring subtidal macroalgae communities. Depth, which is negatively correlated with light, is the main driver structuring subtidal macroalgal communities, however, other gradients such as water flow, and distance to a propagule source also impact their assembly. This study sought to determine which environmental gradients play a prominent role in subtidal macroalgal community assembly. To study subtidal macroalgal community assembly, 92 artificial reef units called Reef Balls were deployed east of the Ogden Point Breakwater in Victoria, BC in June 2009. Two years passed to allow for macroalgal growth and early successional processes to occur prior to sampling the communities on thirty Reef Balls via underwater collection in July 2011. Algae were sorted by genus and dry weight was measured. To determine effects of environmental gradients on community assembly light, depth, water flow, distance to the nearest Reef Ball and distance to the breakwater were measured at each Reef Ball. A redundancy analysis revealed that depth was the most significant environmental gradient shaping algae communities and had the greatest effect on upper canopy algae. Spatial plots reveal a depth and coastline zonation of algae genera comprising the canopy. While depth was found to significantly structure algae genera found in the canopy, there was a high degree of unexplained variation in the model. This suggests that unmeasured variables such as colonization and priority effects may be driving algal community structure in the lower canopy. Differences in community structure between upper and lower canopy reveal that multiple mechanisms are responsible for shaping subtidal algal communities. Further study is required to determine the importance of stochastic colonization events and priority effects. / Graduate / 0329 / vmucciar@uvic.ca

Anadromous Pacific salmon (Oncorhynchus spp.) return annually from marine ecosystems to their natal freshwater habitat to spawn and die. Runs of spawning salmon provide an important source of nutrients and energy to watersheds. However, in coastal systems, substantial amounts of salmon-derived nutrients can be exported back to estuaries. Human land use, including agriculture and urban development, also contribute substantial nutrients to coastal ecosystems, and have the potential to confound results from salmon-derived nutrient studies. This thesis examines the influences of spawning salmon and human land use on stream nutrient and particulate dynamics, including export to estuaries. It also investigates the use of the stable isotope composition (δ13C and δ15N) of estuarine clams, the varnish clam (Nuttalia obscurata: Reeve, 1857) and the manila clam (Tapes philippinarum), and their food sources, as indices of the freshwater export of salmon-derived nutrients to estuaries. Samples were collected from three nearby river-estuary systems along Southeast Vancouver Island, British Columbia. Study systems had either a large number of returning salmon and little human land use (Goldstream), few returning salmon and extensive human land use (Shawnigan), or few returning salmon and little human land use (Holland). In Goldstream River, high abundance of salmon carcasses increased concentrations of total nitrogen and total phosphorus stream water below a barrier to upstream salmon migration. Carcasses also contributed substantial amounts of organic matter to the stream, as indicated by high δ13C and δ15N, and corresponding low C:N ratios in suspended particulate organic matter. My calculations indicate that between 51-77% of the phosphorus transported upstream by migrating salmon, was exported back to the estuary. Human land use also increased downstream nutrient concentrations and raised baseline δ15N in stream ecosystems, which is cause for concern and caution for salmon-derived nutrient studies in land use-affected watersheds, or in the reverse situation, for anthropogenic nutrient studies in watersheds that support runs of anadromous salmon. The high δ15N of anthropogenic nitrogen was not evident in the Shawnigan Estuary. In the Goldstream Estuary salmon-derived nutrients appeared to increase the δ15N of clams, and both the δ13C and δ15N of sedimentary organic matter (SOM), with more enrichment in the high intertidal zone near the river mouth, than in the mid-intertidal zone. The stable isotope composition of clams and SOM was relatively constant across the period of salmon spawning and carcass decay, indicating that they may reflect a legacy salmon-derived nutrient input into estuaries. This study demonstrates that substantial amounts of salmon-derived nutrients are exported back downstream to the Goldstream Estuary where they appear to become integrated into the estuarine food web. Data from a series of estuaries receiving a range of nutrients inputs from salmon is needed to confirm indices of salmon-derived nutrients in estuaries. There is also need for more extensive examination regarding the downstream effects of salmon-derived nutrients in areas such as estuarine productivity, community composition, and positive feedback mechanisms that influence salmon populations. This last area of research is of particular importance considering the high number of salmon stocks at risk in B.C.

To assess the affects of permafrost degradation on key components of the aquatic food web, an in situ manipulative mesocosm experiment was performed in an upland, unslumped Arctic lake located near Inuvik, Northwest Territories. In total, twelve replicate mesocosms were established, 3 control and 3 replicates of 3 treatment levels each dosed with differing amounts of sediments sourced from a nearby thermokarst slumped lake. Findings from the experiment showed that pelagic autotrophic processes had the greatest potential to contribute to higher trophic levels regardless of treatment. Even in the high sediment treatment level, which showed the least pelagic autotrophic production, pelagic autotrophic production was two orders of magnitude greater than pelagic heterotrophic production and 5 times greater than benthic autotrophic or heterotrophic production. Sediment treatment had no significant effect on benthic primary productivity; however, a 500% increase in benthic heterotrophic production was observed. This raises the possibility that increased activity in benthic heterotrophic production is the first step in thermokarst-affected lake ecosystem succession leading to the proliferation of benthic primary production observed in many slumped lakes in the western Canadian Arctic. Water column phosphorus concentrations increased with increasing sediment treatment while pelagic primary production decreased and zooplankton biomass increased. These results suggest that the initial effect of thermokarst disturbance is an enrichment of the system and that top-down predation from zooplankton regulate the abundance of phytoplankton in these systems. If incidences of thermokarst disturbance continue to increase as predicted by current climate models/scenarios, results from this study suggest that the structure and function of Arctic aquatic ecosystems will be significantly impacted. This study highlights the need for further research to obtain a better mechanistic and predictive understanding of the potential effects of thermokarst disturbance on the geochemistry and ecology of Arctic lakes at relevant spatial and temporal scales. / Graduate

As human activity along coastlines increase, degradation and destruction of coastal marine ecosystems around the globe will increase at an alarming rate. In an effort to mitigate degradation and destruction of coastal marine ecosystems, artificial reefs have been used in restoration and enhancement projects. As artificial reefs are the main method of restoring diversity to a degraded area, it is important to know the mechanisms that drive marine community assembly and diversity on those reefs. Understanding community assembly patterns of foundational species, in particular, may provide insight to community assembly patterns at higher trophic levels. Subtidal macroalgae are commonly seen as foundational species in marine environments and both deterministic and stochastic processes play a role in their assembly. Environmental gradients, which are deterministic processes, play a significant role in structuring subtidal macroalgae communities. Depth, which is negatively correlated with light, is the main driver structuring subtidal macroalgal communities, however, other gradients such as water flow, and distance to a propagule source also impact their assembly. This study sought to determine which environmental gradients play a prominent role in subtidal macroalgal community assembly. To study subtidal macroalgal community assembly, 92 artificial reef units called Reef Balls were deployed east of the Ogden Point Breakwater in Victoria, BC in June 2009. Two years passed to allow for macroalgal growth and early successional processes to occur prior to sampling the communities on thirty Reef Balls via underwater collection in July 2011. Algae were sorted by genus and dry weight was measured. To determine effects of environmental gradients on community assembly light, depth, water flow, distance to the nearest Reef Ball and distance to the breakwater were measured at each Reef Ball. A redundancy analysis revealed that depth was the most significant environmental gradient shaping algae communities and had the greatest effect on upper canopy algae. Spatial plots reveal a depth and coastline zonation of algae genera comprising the canopy. While depth was found to significantly structure algae genera found in the canopy, there was a high degree of unexplained variation in the model. This suggests that unmeasured variables such as colonization and priority effects may be driving algal community structure in the lower canopy. Differences in community structure between upper and lower canopy reveal that multiple mechanisms are responsible for shaping subtidal algal communities. Further study is required to determine the importance of stochastic colonization events and priority effects. / Graduate / 0329 / vmucciar@uvic.ca

In the context of growing tourism and global warming, the fragile landscape of the Himalayas is under immense pressure because of rapid land cover changes in developing countries like Nepal. Remotely sensed data combined with ethnographic knowledge are useful tools for studying such changes. The quantitative change can be measured analyzing satellite images whereas local people’s perceptions provide supportive information. To measure such changes in Sagarmatha National Park of Nepal, Multispectral Scanner (MSS) and Thematic Mapper (TM) images since 1972 were used. Normalized Difference Vegetation Index (NDVI) was calculated for different elevation classes and land cover types. These measurements, along with land cover change (1992- 2006) analysis, shows a significant conversion of the areas covered by ice, shrub and grass to rock and soil. Factors including political conflict due to a Maoist rebellion group, inactive park management, increasing tourist demand, and consequent natural resources exploitation helped to explain the change in the forested areas. This is supported by the information from short, informal, semi-structured interviews with local people. However, the local people are unaware of global warming, which has caused the ice melting and glacial lake expansion. Although global causes are out of the immediate control of land managers, better management practices and managed tourism might help alleviate deteriorating Himalayan ecosystems.

The secondary productivity of communities is inherently influenced by the availability and quality of food resources. Movement of organic matter (OM) across landscapes can connect adjacent systems by providing subsidies of carbon and nutrients, implying that alterations of environments from their natural state may affect the productivity of neighboring food webs. The intact terrestrial and marine environments of Fiordland provide a setting to study linkages between the land and the sea. The first general objective of this study was to determine if large but nutritionally poor (nitrogen-poor, carbon-rich) inputs of forest litter support marine secondary production, and to identify pathways for incorporation of this material into upper trophic levels. Pools of marine and terrestrial OM had distinct values of [delta]�⁵N, [delta]��C and [delta]�⁴S, providing high power to estimate the relative use of these sources by the food webs of the fjord-head deltas. Deposit feeding invertebrates (e.g. Echinocardium cordatum, Pectinaria australis) directly assimilated plant detritus. Heterotrophic bacteria on the surface of the sediment assimilated forest litter and provided a potential food source for invertebrates. Chemoautotrophic bacteria fix CO₂ that originates from decomposing forest litter, thus providing an indirect pathway for incorporation of forest litter into the food webs. In the deep basins the strength of the flux of uptake by chemoautotrophic bacteria through the benthic food web into the upper trophic levels was demonstrated by hagfish (Eptatretus cirrhatus) obtaining 38-51% of their nutrition from these bacteria. The ability of a community to utilize discrete sources of OM relies on the presence of specific functional feeding guilds. Marine algae provide a highly nutritive (nitrogen-rich) food source for the fjord communities and fluxes of algae into food webs are facilitated by grazing invertebrates and filter feeders. The second general objective of this study was to determine how the effective loss of filter feeders from inner Doubtful Sound would alter the flux of marine-derived OM to the food webs of the delta communities. The low salinity environment imposed by the hydroelectric power station in Doubtful Sound caused a large reduction in the abundance of the infaunal bivalves Austrovenus stutchburyi and Paphies australis from delta habitats. Clams could tolerate periods of freshwater exposure of [less than or equal to]20 days duration, but the constant freshwater conditions in Doubtful Sound decreased survivorship. In 2004/05 the biomass of these species in inner Doubtful Sound (7.28 tonnes) was 29 times smaller than in Bradshaw Sound (214.12 tonnes). The associated loss of biodeposits (~91 tonnes(DW) yr⁻� in Bradshaw Sound vs. 1 tonne(DW) yr⁻� in inner Doubtful Sound) may have also altered the flux of nutritive OM to the infaunal community. The river delta communities in inner Doubtful Sound appear to have a higher reliance on forest litter than those in Bradshaw Sound, which is apparent as low values of [delta]�⁵N and [delta]��C for estuarine fish (Notolabrus celidotus, Hemerocoetes monopterygius), which act as integrators of the benthic community. This study demonstrates important linkages between terrestrial and coastal marine ecosystems and highlights the role of functional diversity in facilitating fluxes of organic material through food webs.

Thesis (Ph.D.) -- McMaster University, 1998. / Includes bibliographical references (p. 223-258). Also available via World Wide Web.

Diss. (sammanfattning) Umeå : Univ., 2003. / Härtill 6 uppsatser.