New Zealand's Indigenous Forests and Shrublands
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1.2 INDIGENOUS FORESTS NEW ZEALAND’S INDIGENOUS FORESTS AND SHRUBLANDS Robert B. Allen, Peter J. Bellingham, Robert J. Holdaway, Susan K. Wiser Landcare Research, PO Box 69040, Lincoln 7640, New Zealand ABSTRACT: New Zealand’s remaining indigenous forests and shrublands are of immense cultural, environmental, and economic signifi - cance. Their composition, structure, and function are driven by a diverse array of factors, many of which are complexly interrelated. The imprint of disturbances is pervasive and it is necessary to understand disturbances to interpret anthropogenic impacts. For example, understanding impacts of exotic browsing mammals is only possible if they are placed in the context of forest development and tree demographic processes. Recently, a representative plot-based sample of the country’s indigenous forests and shrublands has allowed an unbiased depiction of their composition and structure that is needed for international reporting, performance assessment and manage- ment prioritisation. There are now extensive areas of shrublands successional to forest, often composed of novel mixtures of indigenous and exotic species. These shrublands provide expanded opportunities for ecosystem services from, for example, carbon sequestration to water quality. An increasing area of indigenous forests and shrublands is managed for distinctive Māori aspirations that include sustain- able use. Key words: composition, disturbance, forests, function, management, structure, trends. INTRODUCTION The current forest area represents a >70% reduction from the New Zealand’s indigenous forests and shrublands currently prehuman state (c. 800 years ago) due to historical fi re, forest cover c. 23% and 10% of New Zealand’s 27-million-hectare clearance, and logging (e.g. Wardle 1991). The expanding infl u- land surface respectively (Thompson et al. 2004). These largely ence of humans continues to have wide-ranging stand-level, evergreen forests have four major physiognomic elements: local or regional effects, for example those brought about by Nothofagus spp. (beech), broadleaved angiosperm trees, Agathis invasive species (e.g. Allen and Lee 2006), while others have an australis (kauri), and other conifers (predominantly podocarps) even broader infl uence, for example those brought about by an (Cockayne 1928; Wardle 1991). Warm temperate forests in the increasing atmospheric CO2 concentration. It is often diffi cult, north give way to cool temperate forests further south (Cockayne however, to determine why the structure, composition, and func- 1926) and tree species richness also decreases from north to south tioning of forests is changing. Such changes can be correlated (McGlone et al. 2010). Hence some of the dominant trees of with many factors (Table 1), some of which are relatively well warm temperate forests, such as kauri and taraire (Beilschmiedia characterised as they are easy to measure (e.g. precipitation), tarairi), are restricted to north of latitude 38°S (Wardle 1991). while others we know little about (e.g. individual species’ effects Subalpine shrublands occupy the zone between montane forests on ecosystems). Often these factors are themselves correlated and alpine grasslands and there are also extensive areas of so it is challenging to defi ne causal relationships. This chapter lowland and montane shrublands that are successional to forest fi rst describes what we know about how disturbance, and related (Wardle 1991). factors (Table 1), drive change in indigenous forests and shrub- lands, then defi nes the structure and composition of current TABLE 1 Factors commonly shown to infl uence the structure, composition forests, what factors these relate to, and how they are changing, and functioning of forests and shrublands. Some of the variables commonly and fi nally considers the consequences of their management for measured to represent these factors are given, as well as some of the mecha- ecosystem services. Our emphasis is on an ecosystem process nisms through which these factors operate (modifi ed from Allen et al. 2003) perspective, informed by a trait-based approach. Factors Variables Examples of mechanism Disturbance Changes in biomass or Individuals killed of one DISTURBANCE, SUCCESSION, AND ECOSYSTEM number of trees or more species DEVELOPMENT Herbivory Level of defoliation, Reduced photosynthetic Disturbances are a fundamental feature of forest ecosys- individual height ability, nutrient removal tems, promoting their regeneration and the maintenance of growth species diversity, population structure and ecosystem function. Species effects Litter quality, decay Modifi es the abilities of New Zealand’s location at the intersection of large tectonic plates, resistance of woody seeds of other species to for example, means that its vegetation is prone to disturbances debris germinate and grow that can at times be severe and extensive. Climate Temperature, Changes physiological precipitation processes Disturbance types and ecosystem responses Soil Texture, N availability, Infl uences resources cation availability essential for growth and Most of New Zealand’s current indigenous forests bear the development imprints of natural disturbances long past. The current forests, Dispersal Seed dispersal, Seeds do not arrive at an especially in the South Island, still bear the mark of past glacia- available regeneration otherwise suitable site tions. Successive advances of ice sheets in Westland removed niches beech (Nothofagus spp.) from areas where it was present. Beech Time Tree age, relative Species’ differential takes a long time to recolonise from margins because of slow seed biomass longevity spread and dependence on mycorrhizal fungi for establishment Assembly history Species priority effects Facilitation and growth (e.g. Baylis 1980). As a consequence, the forests of 34 Allen RB, Bellingham PJ, Holdaway RJ, Wiser SK 2013. New Zealand’s indigenous forests and shrublands. In Dymond JR ed. Ecosystem services in New Zealand – conditions and trends. Manaaki Whenua Press, Lincoln, New Zealand. INDIGENOUS FORESTS 1.2 central Westland now lack beech. Glaciation also has a rejuve- 1989). Finally, some earthquakes, even those very distant from nating effect because the glacial till includes ground bedrock, New Zealand, cause tsunamis that have almost certainly caused making key mineral nutrients such as phosphorus more available deforestation in coastal regions (e.g. D’Costa et al. 2011). as glaciers retreat. Hence at Franz Josef a series of forest land- The hilly and mountainous regions of New Zealand are scapes have developed, from very recent advances (the Little Ice prone to landslides caused not only by earthquakes but also by Age of the 1600s) through to terraces that were last under glaciers storms and avalanches. Major storms attend prevailing westerly more than 100 000 years ago (e.g. Wardle et al. 2004). In this wet wind patterns, but many intense storms also affect eastern parts climate leaching of mineral nutrients is rapid, and in the absence of New Zealand. Extra-tropical cyclones affect forests in the of glaciation, phosphorus in particular becomes limiting. Thus the northern North Island disproportionately, but periodically intense forests on recently glaciated soils are highly productive and have cyclones track even to high latitudes (Martin and Ogden 2006). a number of species, especially broadleaved species, that depend All such storms can cause landslides locally, and sometimes on ready access to nutrients. In contrast, with all else equal (e.g. over extensive areas, for example during Cyclone Bola of 1988 climate), the forests on soils last under glaciers 120 000 years ago (Page et al. 1999). Finally, high uplift rates coupled with unstable are dominated by slow-growing conifers that can access scarce bedrock can result in large landslides with no obvious triggers. nutrients (e.g. Richardson et al. 2004; Holdaway et al. 2011). One such event deforested 20 hectares on the slopes of Mt Adams In the North Island, the current forests are strongly infl uenced in Westland in 1999 (Hancox et al. 2005). Intense rainfall events by the effects of past volcanic eruptions, especially in the central in river headwaters, coupled with landslides, and especially the North Island. The Taupō Eruption in AD 232 was one of the largest occlusion of gorges by debris and their subsequent breaching, can in the world over the last 5000 years, depositing pumice over cause signifi cant damage to riparian and fl oodplain forests. most of New Zealand and causing devastating pyroclastic fl ows The new parent material deposited by rivers across fl ood- and outbreaks of fi re that destroyed over 30 000 km2 of forest plains has high available concentrations of some nutrients, such (e.g. Wilmshurst and McGlone 1996). The rhyolitic ignimbrite as phosphorus, but is very low in nitrogen and organic matter. and pumice deposits from past eruptions are relatively infer- Shrubs such as tutu (Coriaria arborea) and native broom tile whereas the more common andesitic ash showers produce (Carmichaelia spp.) have symbionts on their roots that overcome fertile deposits and these have a large infl uence on the kinds of nutrient limitation by fi xing atmospheric nitrogen. These shrubs forests that develop. The ash showers from more recent smaller are often abundant on young fl oodplains; their nitrogen-rich litter eruptions since the Taupō Eruption, such as from Kaharoa in AD enhances soil development (e.g. Bellingham et al. 2005). Canopy 1314 and more recently from Tarawera