Saltwater Forest is a Dacrydium cupressinum-dominated lowland forest covering 9000 ha in south Westland, South Island, New Zealand. Four thousand hectares is managed for sustainable production of indigenous timber. The aim of this study was to provide an integrated analysis of soils, soil-landform relationships, and soil-vegetation relationships at broad and detailed scales. The broad scale understandings provide a framework in which existing or future studies can be placed and the detailed studies elucidate sources of soil and forest variability.
Glacial landforms dominate. They include late Pleistocene lateral, terminal and ablation moraines, and outwash aggradation and degradation terraces. Deposits and landforms from six glacial advances have been recognised ranging from latest Last (Otira) Glaciation to Penultimate (Waimea) Glaciation. The absolute ages of landforms were established by analysis of the thickness and soil stratigraphy of loess coverbeds, augmented with radiocarbon dating and phytolith and pollen analysis.
In the prevailing high rainfall of Westland soil formation is rapid. The rate of loess accretion in Saltwater
Forest (ca. 30 mm ka⁻¹) has been low enough that soil formation and loess accretion took place contemporaneously. Soils formed in this manner are known as upbuilding soils. The significant difference between upbuilding pedogenesis and pedogenesis in a topdown sense into an existing sediment body is that each subsoil increment of an upbuilding soil has experienced processes of all horizons above. In Saltwater Forest subsoils of upbuilding soils are strongly altered because they have experienced the extremely acid environment of the soil surface at some earlier time. Some soil chronosequence studies in Westland have included upbuilding soils formed in loess as the older members of the sequence. Rates and types of processes inferred from these soils should be reviewed because upbuilding is a different pedogenic pathway to topdown pedogenesis.
Landform age and morphology were used as a primary stratification for a study of the soil pattern and nature of soil variability in the 4000 ha production area of Saltwater Forest. The age of landforms (> 14 ka) and rapid soil formation mean that soils are uniformly strongly weathered and leached. Soils include Humic Organic Soils, Perch-gley Podzols, Acid Gley Soils, Allophanic Brown Soils, and Orthic or Pan Podzols. The major influence on the nature of soils is site hydrology which is determined by macroscale features of landforms (slope, relief, drainage density), mesoscale effects related to position on landforms, and microscale influences determined by microtopography and individual tree effects. Much of the soil variability arises at microscales so that it is not possible to map areas of uniform soils at practical map scales. The distribution of soil variability across spatial scales, in relation to the intensity of forest management, dictates that it is most appropriate to map soil complexes with boundaries coinciding with landforms.
Disturbance of canopy trees is an important agent in forest dynamics. The frequency of forest disturbance in the production area of Saltwater Forest varies in a systematic way among landforms in accord with changes in abundance of different soils. The frequency of forest turnover is highest on landforms with the greatest abundance of extremely poorly-drained Organic Soils. As the abundance of better-drained soils increases the frequency of forest turnover declines. Changes in turnover frequency are reflected in the mean size and density of canopy trees (Dacrydium cupressinum) among landforms. Terrace and ablation moraine landforms with the greatest abundance of extremely poorly-drained soils have on average the smallest trees growing most densely. The steep lateral moraines, characterised by well drained soils, have fewer, larger trees. The changes manifested at the landform scale are an integration of processes operating over much shorter range as a result of short-range soil variability. The systematic changes in forest structure and turnover frequency among landforms and soils have important implications for sustainable forest management.
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.
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.
Mixed conifer, beech and hardwood forests are relatively common in Aotearoa/New
Zealand, but are not well studied. This thesis investigates the coexistence, regeneration
dynamics and disturbance history of a mixed species forest across an environmental
gradient of drainage and soil development in north Westland.
The aim was to investigate whether conifers, beech and non-beech hardwood species were
able to coexist on surfaces that differed in their underlying edaphic conditions, and if so to understand the mechanisms that influenced their regeneration on both poorly drained and
well drained soils. The site selected was an area of high tree species diversity on a lowland
0.8 km² post-glacial terrace at the base of Mount Harata in the Grey River Valley.
My approach was to use forest stand history reconstruction at two spatial scales: an
intensive within-plot study of stand dynamics (chapter 1) and a whole-landform approach
(chapter 2) that examined whether the dynamics identified at the smaller within-plot scale
reflected larger patterns across the terrace.
In chapter 1, three large permanent plots (0.3-0.7 ha) were placed at different points along
the drainage gradient, one plot situated in each of the mainly well-drained, poorly drained
and very poorly drained areas along the terrace. Information was gathered on species age
and size structures, spatial distributions of tree ages, species interactions, microsite
establishment preferences, patterns of stand mortality, and disturbance history in each plot.
There were differences in stand structure, composition and relative abundance of species
found between the well drained plot and the two poorer drained plots. On the well drained
site conifers were scarce, the beeches Nothofagus fusca and N. menziesii dominated the
canopy, and in the subcanopy the hardwood species Weinmannia racemosa and Quintinia
acutifolia were abundant. As drainage became progressively poorer, the conifers
Dacrydium cupressinum and Dacrycarpus dacrydioides became more abundant and
occupied the emergent tier over a beech canopy. The hardwoods W. racemosa and Q.
acutifolia became gradually less abundant in the subcanopy, whereas the hardwood
Elaeocarpus hookerianus became more so.
In the well drained plot, gap partitioning for light between beeches and hardwoods enabled
coexistence in response to a range of different sized openings resulting from disturbances
of different extent. In the two more poorly drained plots, species also coexisted by
partitioning microsite establishment sites according to drainage.
There were several distinct periods where synchronous establishment of different species
occurred in different plots, suggesting there were large disturbances: c. 100yrs, 190-200
yrs, 275-300 yrs and 375-425 yrs ago. Generally after the same disturbance, different
species regenerated in different plots reflecting the underlying drainage gradient. However,
at the same site after different disturbances, different sets of species regenerated,
suggesting the type and extent of disturbances and the conditions left behind influenced
species regeneration at some times but not others. The regeneration of some species (e.g.,
N. fusca in the well-drained plot, and Dacrydium in the poorer drained plots) was periodic
and appeared to be closely linked to these events. In the intervals between these
disturbances, less extensive disturbances resulted in the more frequent N. menziesii and
especially hardwood regeneration. The type of tree death caused by different disturbances
favoured different species, with dead standing tree death favouring the more shade-tolerant
N. menziesii and hardwoods, whereas uprooting created a mosaic of microsite conditions
and larger gap sizes that enabled Dacrycarpus, N. fusca and E. hookerianus to maintain
themselves in the poorly drained areas.
In chapter 2, 10 circular plots (c. 0.12 ha) were placed in well drained areas and 10
circular plots (c. 0.2 ha) in poorly drained plots to collect information on species
population structures and microsite preferences. The aims were to reconstruct species'
regeneration responses to a range of disturbances of different type and extent across the
whole terrace, and to examine whether there were important differences in the effects of
these disturbances.
At this landform scale, the composition and relative abundances of species across the
drainage gradient reflected those found in chapter 1. There were few scattered conifers in well drained areas, despite many potential regeneration opportunities created from a range
of different stand destroying and smaller scale disturbances.
Three of the four periods identified in chapter 1 reflected distinct terrace-wide periods of
regeneration 75-100 yrs, 200-275 yrs and 350-450 yrs ago, providing strong evidence of
periodic large, infrequent disturbances that occurred at intervals of 100-200 yrs. These
large, infrequent disturbances have had a substantial influence in determining forest
history, and have had long term effects on forest structure and successional processes.
Different large, infrequent disturbances had different effects across the terrace, with the
variability in conditions that resulted enabling different species to regenerate at different
times. For example, the regeneration of distinct even-aged Dacrydium cohorts in poorly
drained areas was linked to historical Alpine Fault earthquakes, but not to more recent
storms. The variation in the intensity of different large, infrequent disturbances at different
points along the environmental drainage gradient, was a key factor influencing the scale of
impacts. In effect, the underlying edaphic conditions influenced species composition along
the drainage gradient and disturbance history regulated the relative abundances of species.
The results presented here further emphasise the importance of large scale disturbances as a
mechanism that allows coexistence of different tree species in mixed forest, in particular
for the conifers Dacrydium, Dacrycarpus and the beech N. fusca, by creating much of the
environmental variation to which these species responded. This study adds to our
understanding of the effects of historical earthquakes in the relatively complex forests of
north Westland, and further illustrates their importance in the Westland forest landscape as
the major influential disturbance on forest pattern and history.
These results also further develop the 'two-component' model used to describe
conifer/angiosperm dynamics, by identifying qualitative differences in the impacts of
different large, infrequent disturbances across an environmental gradient that allowed for
coexistence of different species. In poorer drained areas, these forests may even be thought
of as 'three-component' systems with conifers, beeches and hardwoods exhibiting key
differences in their regeneration patterns after disturbances of different type and extent, and
in their microsite preferences.
