Six stands located on different land forms in mixed old-growth Nothofagus forests in the Matiri Valley (northwest of South Island, New Zealand) were sampled to examine the effects of two recent large earthquakes on tree establishment and tree-ring growth, and how these varied across land forms. 50 trees were cored in each stand to determine age structure and the cores were cross-dated to precisely date unusual periods of radial growth. The 1968 earthquake (M = 7.1, epicentre 35 km from the study area) had no discernible impact on the sampled stands. The impact of the 1929 earthquake (M = 7.7, epicentre 20 km from the study area) varied between stands, depending on whether or not they had been damaged by soil or rock movement. In all stands, the age structures showed a pulse of N. fusca establishment following the 1929 earthquake, with this species dominating establishment in large gaps created by landslides. Smaller gaps, created by branch or tree death, were closed by both N. fusca and N. menziesii. The long period of releases (1929-1945) indicates that direct earthquake damage was not the only cause of tree death, and that many trees died subsequently most likely of pathogen attack or a drought in the early 1930s. The impacts of the 1929 earthquake are compared to a storm in 1905 and a drought in 1974-1978 which also affected forests in the region. Our results confirm that earthquakes are an important factor driving forest dynamics in this tectonically active region, and that the diversity of earthquake impacts is a major source of heterogeneity in forest structure and regeneration.
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.
Tree mortality is a fundamental process governing forest dynamics, but understanding tree mortality patterns is challenging because large, long-term datasets are required. Describing size-specific mortality patterns can be especially difficult, due to few trees in larger size classes. We used permanent plot data from Nothofagus solandri var. cliffortioides (mountain beech) forest on the eastern slopes of the Southern Alps, New Zealand, where the fates of trees on 250 plots of 0.04 ha were followed, to examine: (1) patterns of size-specific mortality over three consecutive periods spanning 30 years, each characterised by different disturbance, and (2) the strength and direction of neighbourhood crowding effects on sizespecific mortality rates. We found that the size-specific mortality function was U-shaped over the 30-year period as well as within two shorter periods characterised by small-scale pinhole beetle and windthrow disturbance. During a third period, characterised by earthquake disturbance, tree mortality was less size dependent. Small trees (,20 cm in diameter) were more likely to die, in all three periods, if surrounded by a high basal area of larger neighbours, suggesting that sizeasymmetric competition for light was a major cause of mortality. In contrast, large trees ($20 cm in diameter) were more likely to die in the first period if they had few neighbours, indicating that positive crowding effects were sometimes important for survival of large trees. Overall our results suggest that temporal variability in size-specific mortality patterns, and positive interactions between large trees, may sometimes need to be incorporated into models of forest dynamics.
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.
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.
