Coexistence, dynamics and disturbance history of mixed conifer, beech, and hardwood forest, north Westland
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.