Question: Does canopy tree regeneration response to different large disturbances vary with soil drainage? Location: Old-growth conifer (Dacrydium and Dacrycarpus), angiosperm (Nothofagus and Weinmannia) rain forest, Mount Harata, South Island, New Zealand. Methods: Trees were aged (1056 cores) to reconstruct stand history in 20 (0.12 - 0.2 ha) plots with different underlying drainage. Spatial analyses of an additional 805 tree ages collected from two (0.3 - 0.7 ha) plots were conducted to detect patchiness for five canopy tree species. Microsite preferences for trees and saplings were determined. Results: There were clear differences in species regeneration patterns on soils with different drainage. Conifer recruitment occurred infrequently in even-aged patches (> 1000 m²) and only on poorly drained soils. Periodic Nothofagus fusca and N. menziesii recruitment occurred more frequently in different sized canopy openings on all soils. Weinmannia recruitment was more continuous on all soils reflecting their greater relative shade-tolerance. Distinct periods of recruitment that occurred in the last 400 years matched known large disturbances in the region. These events affected species differently as soil drainage varied. Following earthquakes, both conifers and N. menziesii regenerated on poorly drained soils, while Nothofagus species and Weinmannia regenerated on well-drained soils. However, Dacrydium failed to regenerate after patchy storm damage in the wetter forest interior; instead faster-growing N. fusca captured elevated microsites caused by uprooting. Conclusions: Underlying drainage influenced species composition, while variation in the impacts of large disturbance regulated relative species abundances on different soils.
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.
We examined the stratigraphy of alluvial fans formed at the steep range front of the Southern Alps at Te Taho, on the north bank of the Whataroa River in central West Coast, South Island, New Zealand. The range front coincides with the Alpine Fault, an Australian-Pacific plate boundary fault, which produces regular earthquakes. Our study of range front fans revealed aggradation at 100- to 300-year intervals. Radiocarbon ages and soil residence times (SRTs) estimated by a quantitative profile development index allowed us to elucidate the characteristics of four episodes of aggradation since 1000 CE. We postulate a repeating mode of fan behaviour (fan response cycle [FRC]) linked to earthquake cycles via earthquake-triggered landslides. FRCs are characterised by short response time (aggradation followed by incision) and a long phase when channels are entrenched and fan surfaces are stable (persistence time). Currently, the Te Taho and Whataroa River fans are in the latter phase. The four episodes of fan building we determined from an OxCal sequence model correlate to Alpine Fault earthquakes (or other subsidiary events) and support prior landscape evolution studies indicating ≥M7.5 earthquakes as the main driver of episodic sedimentation. Our findings are consistent with other historic non-earthquake events on the West Coast but indicate faster responses than other earthquake sites in New Zealand and elsewhere where rainfall and stream gradients (the basis for stream power) are lower. Judging from the thickness of fan deposits and the short response times, we conclude that pastoral farming (current land-use) on the fans and probably across much of the Whataroa River fan would be impossible for several decades after a major earthquake. The sustainability of regional tourism and agriculture is at risk, more so because of the vulnerability of the single through road in the region (State Highway 6).
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.
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.
