This report reviews the literature on regeneration requirements of main canopy tree species in Westland. Forests managed for production purposes have to be harvested in an ecologically sustainable way; to maintain their natural character, harvesting should facilitate regeneration of target species and ensure that their recruitment is in proportion to the extent of extraction. The reasons for species establishing at any point in time are unclear; however, they are probably related to the availability of suitable microsites for establishment, the size of the canopy openings formed by disturbance, and whether or not seeds are available at or around the time of the disturbance. Age structures from throughout Westland show that extensive, similar-aged, post-earthquake cohorts of trees are a feature of the region. This suggests that infrequent, massive earthquakes are the dominant coarse-scale disturbance agent, triggering episodes of major erosion and sedimentation and leaving a strong imprint in the forest structure. In other forests, flooding and catastrophic windthrow are major forms of disturbance. The findings suggest that, in general, large disturbances are required for conifer regeneration. This has implications for any sustained yield management of these forests if conifers are to remain an important component. Any harvesting should recognise the importance for tree establishment of: forest floor microsites, such as fallen logs and tree tip-up mounds; and the variable way in which canopy gaps are formed. Harvesting should maintain the 'patchy' nature of the natural forest—large patches of dense conifers interspersed with more heterogeneous patches of mixed species.
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.
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.
This thesis investigates landscape disturbance history in Westland since 1350 AD. Specifically, I test the hypothesis that large-magnitude regional episodes of natural disturbance have periodically devastated portions of the landscape and forest, and that these were caused by infrequent earthquakes along the Alpine Fault. Forest stand history reconstruction was used to determine the timing and extent of erosion and sedimentation events that initiated new forest cohorts in a 1412 ha study area in the Karangarua River catchment, south Westland. Over 85 % of the study area was disturbed sufficiently by erosion/sedimentation since 1350 AD to initiate new forest cohorts. During this time four episodes of catchment-wide disturbance impacted the study area, and these took place about 1825 AD ± 5 years (Ruera episode), 1715 AD ± 5 years (Sparkling episode), 1615 AD ± 5 years (McTaggart episode), and 1445 AD ± 15 years (Junction episode). The three most recent episodes disturbed 10 %, 35-40 % and 32-50 % respectively of the study area. The Junction episode disturbed at least 6 % of the study area, but elimination of evidence by more recent disturbances prevented an upper limit being defined. The three earliest episodes correspond to the date-ranges for three Alpine Fault earthquakes from geological data, and are the only episodes of disturbance within each date-range. An earthquake cause is also consistent with features of the disturbance record: large portions of the study area were disturbed, disturbance occurred on all types 'of landforms, and terrace surfaces were abandoned upstream of the Alpine Fault. On this basis erosion/sedimentation induced by Alpine Fault earthquakes has disturbed 14-20 % of the land surface in the study area per century. Storms and other non-seismic erosional processes have disturbed 3-4 % per century. To examine the importance of the Alpine Fault earthquakes to forest disturbance throughout Westland, I collated all available data on conifer stand age structures in the region and identified dates of disturbance events from 55 even-aged cohorts of trees. Three region-wide episodes of forest disturbance since 1350 AD were found in this sample, and these matched the three Alpine Fault earthquake-caused episodes found in the Karangarua. Forest disturbance at these times was widespread across Westland over at least 200 km from Paringa to Hokitika, and originated from both tree fall and erosion processes. This disturbance history can explain the long-observed regional conifer forest pattern in Westland, of a predominance of similar-sized stands of trees and a relative lack of small-sized (young) stands. The many similar-sized stands are a consequence of synchronous forest disturbance and re-establishment accompanying the infrequent Alpine Fault earthquakes, while the dominance of mature stands of trees and relative lack of young small-sized trees in stands is explained by the long lapsed time since the last Alpine Fault earthquake (c. 280 years). I applied the landscape disturbance history information to the existing geological data to reconstruct the paleoseismicity of the Alpine Fault since 1350 AD. Best estimates for the timing of the most recent three rupture events from these data are 1715 AD ± 5 years, 1615 AD ± 5 years and 1445 AD ± 15 years. Earthquake recurrence intervals were variable, ranging from about 100 years to at least 280 years (the lapsed time since the last event). All three events caused forest and geomorphic disturbance over at least a 200 km section of Fault between the Karangarua and Hokitika Rivers, and were probably single rupture events. Suppressions in cross dated tree-ring chronologies in the western South Island suggest that the last rupture occurred in 1717 AD, and extended as a single rupture from Haupiri to Fiordland, a distance along the Fault of 375 km.
