Globally, the maximum elevations at which treelines are observed to occur coincide with a 6.4 °C soil isotherm. However, when observed at finer scales, treelines display a considerable degree of spatial complexity in their patterns across the landscape and are often found occurring at lower elevations than expected relative to the global-scale pattern. There is still a
lack of understanding of how the abiotic environment imposes constraints on treeline patterns, the scales at which different effects are acting, and how these effects vary over large spatial extents. In this thesis, I examined abrupt Nothofagus treelines across seven degrees of
latitude in New Zealand in order to investigate two broad questions: (1) What is the nature and extent of spatial variability in Nothofagus treelines across the country? (2) How is this variation associated with abiotic variation at different spatial scales? A range of GIS, statistical, and atmospheric modelling methods were applied to address these two questions.
First, I characterised Nothofagus treeline patterns at a 15x15km scale across New Zealand using a set of seven, GIS-derived, quantitative metrics that describe different aspects of treeline position, shape, spatial configuration, and relationships with adjacent vegetation.
Multivariate clustering of these metrics revealed distinct treeline types that showed strong spatial aggregation across the country. This suggests a strong spatial structuring of the abiotic environment which, in turn, drives treeline patterns. About half of the multivariate treeline
metric variation was explained by patterns of climate, substrate, topographic and disturbance variability; on the whole, climatic and disturbance factors were most influential.
Second, I developed a conceptual model that describes how treeline elevation may
vary at different scales according to three categories of effects: thermal modifying effects, physiological stressors, and disturbance effects. I tested the relevance of this model for Nothofagus treelines by investigating treeline elevation variation at five nested scales (regional to local) using a hierarchical design based on nested river catchments. Hierarchical linear modelling revealed that the majority of the variation in treeline elevation resided at the broadest, regional scale, which was best explained by the thermal modifying effects of solar radiation, mountain mass, and differences in the potential for cold air ponding. Nonetheless, at finer scales, physiological and disturbance effects were important and acted to modify the regional trend at these scales. These results suggest that variation in abrupt treeline elevations
are due to both broad-scale temperature-based growth limitation processes and finer-scale stress- and disturbance-related effects on seedling establishment.
Third, I explored the applicability of a meso-scale atmospheric model, The Air
Pollution Model (TAPM), for generating 200 m resolution, hourly topoclimatic data for
temperature, incoming and outgoing radiation, relative humidity, and wind speeds. Initial assessments of TAPM outputs against data from two climate station locations over seven years showed that the model could generate predictions with a consistent level of accuracy for both sites, and which agreed with other evaluations in the literature. TAPM was then used to generate data at 28, 7x7 km Nothofagus treeline zones across New Zealand for January
(summer) and July (winter) 2002. Using mixed-effects linear models, I determined that both
site-level factors (mean growing season temperature, mountain mass, precipitation,
earthquake intensity) and local-level landform (slope and convexity) and topoclimatic factors (solar radiation, photoinhibition index, frost index, desiccation index) were influential in
explaining variation in treeline elevation within and among these sites. Treelines were
generally closer to their site-level maxima in regions with higher mean growing season
temperatures, larger mountains, and lower levels of precipitation. Within sites, higher
treelines were associated with higher solar radiation, and lower photoinhibition and
desiccation index values, in January, and lower desiccation index values in July. Higher treelines were also significantly associated with steeper, more convex landforms.
Overall, this thesis shows that investigating treelines across extensive areas at multiple study scales enables the development of a more comprehensive understanding of treeline variability and underlying environmental constraints. These results can be used to formulate new hypotheses regarding the mechanisms driving treeline formation and to guide the optimal choice of field sites at which to test these hypotheses.
Environmental stress and disturbance can affect the structure and functioning of marine ecosystems by altering their physical, chemical and biological features. In estuaries, benthic invertebrate communities play important roles in structuring sediments, influencing primary production and biogeochemical flux, and occupying key food web positions. Stress and disturbance can reduce species diversity, richness and abundance, with ecological theory predicting that biodiversity will be at its lowest soon after a disturbance with assemblages dominated by opportunistic species. The Avon-Heathcote Estuary in Christchurch New Zealand has provided a novel opportunity to examine the effects of stress, in the form of eutrophication, and disturbance, in the form of cataclysmic earthquake events, on the structure and functioning of an estuarine ecosystem. For more than 50 years, large quantities (up to 500,000m3/day) of treated wastewater were released into this estuary but in March 2010 this was diverted to an ocean outfall, thereby reducing the nutrient loading by around 90% to the estuary. This study was therefore initially focussed on the reversal of eutrophication and consequent effects on food web structure in the estuary as it responded to lower nutrients. In 2011, however, Christchurch was struck with a series of large earthquakes that greatly changed the estuary. Massive amounts of liquefied sediments, covering up to 65% of the estuary floor, were forced up from deep below the estuary, the estuary was tilted by up to a 50cm rise on one side and a corresponding drop on the other, and large quantities of raw sewage from broken wastewater infrastructure entered the estuary for up to nine months. This study was therefore a test of the potentially synergistic effects of nutrient reduction and earthquake disturbance on invertebrate communities, associated habitats and food web dynamics. Because there was considerable site-to-site heterogeneity in the estuary, the sites in this study were selected to represent a eutrophication gradient from relatively “clean” (where the influence of tidal flows was high) to highly impacted (near the historical discharge site). The study was structured around these sites, with components before the wastewater diversion, after the diversion but before the earthquakes, and after the earthquakes. The eutrophication gradient was reflected in the composition and isotopic chemistry of primary producer and invertebrate communities and the characteristics of sediments across the sample sites. Sites closest to the former wastewater discharge pipe were the most eutrophic and had cohesive organic -rich, fine sediments and relatively depauperate communities dominated by the opportunistic taxa Capitellidae. The less-impacted sites had coarser, sandier sediments with fewer pollutants and far less organic matter than at the eutrophic sites, relatively high diversity and lower abundances of micro- and macro-algae. Sewage-derived nitrogen had became incorporated into the estuarine food web at the eutrophic sites, starting at the base of the food chain with benthic microalgae (BMA), which were found to use mostly sediment-derived nitrogen. Stable isotopic analysis showed that δ13C and δ15N values of most food sources and consumers varied spatially, temporally and in relation to the diversion of wastewater, whereas the earthquakes did not appear to affect the overall estuarine food web structure. This was seen particularly at the most eutrophic site, where isotopic signatures became more similar to the cleaner sites over two-and-a-half years after the diversion. New sediments (liquefaction) produced by the earthquakes were found to be coarser, have lower concentrations of heavy metals and less organic matter than old (existing) sediments. They also had fewer macroinvertebrate inhabitants initially after the earthquakes but most areas recovered to pre-earthquake abundance and diversity within two years. Field experiments showed that there were higher amounts of primary production and lower amounts of nutrient efflux from new sediments at the eutrophic sites after the earthquakes. Primary production was highest in new sediments due to the increased photosynthetic efficiency of BMA resulting from the increased permeability of new sediments allowing increased light penetration, enhanced vertical migration of BMA and the enhanced transport of oxygen and nutrients. The reduced efflux of NH4-N in new sediments indicated that the capping of a large portion of eutrophic old sediments with new sediments had reduced the release of legacy nutrients (originating from the historical discharge) from the sediments to the overlying water. Laboratory experiments using an array of species and old and new sediments showed that invertebrates altered levels of primary production and nutrient flux but effects varied among species. The mud snail Amphibola crenata and mud crab Austrohelice crassa were found to reduce primary production and BMA biomass through the consumption of BMA (both species) and its burial from bioturbation and the construction of burrows (Austrohelice). In contrast, the cockle Austrovenus stutchburyi did not significantly affect primary production and BMA biomass. These results show that changes in the structure of invertebrate communities resulting from disturbances can also have consequences for the functioning of the system. The major conclusions of this study were that the wastewater diversion had a major effect on food web dynamics and that the large quantities of clean and unpolluted new sediments introduced to the estuary during the earthquakes altered the recovery trajectory of the estuary, accelerating it at least throughout the duration of this study. This was largely through the ‘capping’ effect of the new liquefied, coarser-grained sediments as they dissipated across the estuary and covered much of the old organic-rich eutrophic sediments. For all aspects of this study, the largest changes occurred at the most eutrophic sites; however, the surrounding habitats were important as they provided the context for recovery of the estuary, particularly because of the very strong influence of sediments, their biogeochemistry, microalgal and macroalgal dynamics. There have been few studies documenting system level responses to eutrophication amelioration and to the best on my knowledge there are no other published studies examining the impacts of large earthquakes on benthic communities in an estuarine ecosystem. This research gives valuable insight and advancements in the scientific understanding of the effects that eutrophication recovery and large-scale disturbances can have on the ecology of a soft-sediment ecosystem.
Mitigating the cascade of environmental damage caused by the movement of excess reactive nitrogen (N) from land to sea is currently limited by difficulties in precisely and accurately measuring N fluxes due to variable rates of attenuation (denitrification) during transport. This thesis develops the use of the natural abundance isotopic composition of nitrate (δ15N and δ18O of NO₃-) to integrate the spatialtemporal variability inherent to denitrification, creating an empirical framework for evaluating attenuation during land to water NO₃- transfers. This technique is based on the knowledge that denitrifiers kinetically discriminate against 'heavy' forms of both N and oxygen (O), creating a parallel enrichment in isotopes of both species as the reaction progresses. This discrimination can be quantitatively related to NO₃- attenuation by isotopic enrichment factors (εdenit). However, while these principles are understood, use of NO₃- isotopes to quantify denitrification fluxes in non-marine environments has been limited by, 1) poor understanding of εdenit
variability, and, 2) difficulty in distinguishing the extent of mixing of isotopically distinct sources from the
imprint of denitrification. Through a combination of critical literature analysis, mathematical modelling, mesocosm to field scale experiments, and empirical studies on two river systems over distance and time, these short comings are parametrised and a template for future NO₃- isotope based attenuation measurements
outlined.
Published εdenit values (n = 169) are collated in the literature analysis presented in Chapter 2. By evaluating these values in the context of known controllers on the denitrification process, it is found that the magnitude of εdenit, for both δ15N and δ18O, is controlled by, 1) biology, 2) mode of transport through the denitrifying zone (diffusion v. advection), and, 3) nitrification (spatial-temporal distance between nitrification and denitrification). Based on the outcomes of this synthesis, the impact of the three factors identified as controlling εdenit are quantified in the context of freshwater systems by combining simple mathematical modelling and lab incubation studies (comparison of natural variation in biological versus physical expression). Biologically-defined εdenit, measured in sediments collected from four sites along a temperate stream and from three tropical submerged paddy fields, varied from -3‰ to -28‰ depending on the site’s antecedent carbon content. Following diffusive transport to aerobic surface water, εdenit was found to
become more homogeneous, but also lower, with the strength of the effect controlled primarily by diffusive distance and the rate of denitrification in the sediments. I conclude that, given the variability in fractionation dynamics at all levels, applying a range of εdenit from -2‰ to -10‰ provides more accurate measurements of attenuation than attempting to establish a site-specific value. Applying this understanding of denitrification's fractionation dynamics, four field studies were conducted to measure denitrification/ NO₃- attenuation across diverse terrestrial → freshwater systems. The development of NO₃- isotopic signatures (i.e., the impact of nitrification, biological N fixation, and ammonia volatilisation on the isotopic 'imprint' of denitrification) were evaluated within two key agricultural regions: New Zealand grazed pastures (Chapter 4) and Philippine lowland submerged rice production (Chapter 5). By measuring the isotopic composition of soil ammonium, NO₃- and volatilised ammonia following the bovine urine deposition, it was determined that the isotopic composition of NO₃ - leached from grazed pastures is defined by the balance between nitrification and denitrification, not ammonia volatilisation. Consequently, NO₃- created within pasture systems was predicted to range from +10‰ (δ15N)and -0.9‰ (δ18O) for non-fertilised fields (N limited) to -3‰ (δ15N) and +2‰ (δ18O) for grazed fertilised fields (N saturated). Denitrification was also the dominant determinant of NO₃- signatures in the Philippine rice paddy. Using a site-specific εdenit for the paddy, N inputs versus attenuation were able to be calculated, revealing that >50% of available N in the top 10 cm of soil was denitrified during land preparation, and >80% of available N by two weeks post-transplanting. Intriguingly, this denitrification was driven by rapid
NO₃- production via nitrification of newly mineralised N during land preparation activities.
Building on the relevant range of εdenit established in Chapters 2 and 3, as well as the soil-zone confirmation that denitrification was the primary determinant of NO₃- isotopic composition, two long-term
longitudinal river studies were conducted to assess attenuation during transport. In Chapter 6, impact and recovery dynamics in an urban stream were assessed over six months along a longitudinal impact gradient using measurements of NO₃- dual isotopes, biological populations, and stream chemistry. Within 10 days of the catastrophic Christchurch earthquake, dissolved oxygen in the lowest reaches was <1 mg l⁻¹, in-stream denitrification accelerated (attenuating 40-80% of sewage N), microbial biofilm communities changed, and several benthic invertebrate taxa disappeared. To test the strength of this method for tackling the diffuse, chronic N loading of streams in agricultural regions, two years of longitudinal measurements of NO₃- isotopes were collected. Attenuation was negatively correlated with NO₃- concentration, and was highly
dependent on rainfall: 93% of calculated attenuation (20 kg NO₃--N ha⁻¹ y⁻¹) occurred within 48 h of rainfall.
The results of these studies demonstrate the power of intense measurements of NO₃- stable isotope for distinguishing temporal and spatial trends in NO₃ - loss pathways, and potentially allow for improved catchment-scale management of agricultural intensification. Overall this work now provides a more cohesive understanding for expanding the use of NO₃- isotopes measurements to generate accurate understandings of the controls on N losses. This information is becoming increasingly important to predict ecosystem response to future changes, such the increasing agricultural intensity needed to meet global food demand, which is occurring synergistically with unpredictable global climate change.
