1. Tree seedling establishment outside forest boundaries is controlled by many interacting factors. Understanding the relative importance of different pressures is essential for improving techniques for forest restoration and for understanding the potential of forest boundaries to shift and adapt to changing climate conditions.2. We investigated constraints on the spread of a key southern hemisphere forest type into grasslands by undertaking a multifactorial experiment sowing Nothofagus cliffortioides (Nothofagaceae) in a historically deforested retired pasture. We manipulated factors to determine the relative effects of sheltered microsites, competition with pasture species, availability of ectomycorrhizal fungi, soil nutrients and rabbit herbivory, on seedling emergence and survival.3. Overall survival was low (11.7% after 6 months and 1.5% after 10 months), but there were strong treatment effects on both seedling establishment and survival.The availability of shelter and competition with pasture species had strongest effects, with the presence of pasture species initially aiding seedling emergence due to a sheltering effect, but negatively affecting survival at later stages. Most remaining seedlings at the conclusion of the experiment had formed ectomycorrhizae regardless of whether inoculum was supplied, and seedling survival and health was positively related to ectomycorrhizal colonization. Fertilizing had less of an effect than other factors, and results regarding rabbit herbivory were inconclusive.
Synthesis and applications.This study provides new insights into how factors interact to limit tree seedling establishment in grasslands and forest expansion into neighbouring ecosystems. This improves understanding of the ability of an ectomycorrhizal keystone forest species to migrate and adapt to climate change. This study also assists restoration practitioners in selecting techniques that will enhance seedling establishment, and highlights that direct seeding approaches in open grasslands are unlikely to result in high rates of Nothofagus establishment. This study was undertaken in the Motatapu Valley, Otago, New Zealand (−44.752°S, 168.910°E), within the 53,000 ha Queen Elizabeth II National Trust Mahu Whenua covenants. Permission to carry out the work was provided by Soho Properties Ltd. Ethics approval was not required. Prior to human arrival (~1300 CE), this area was likely dominated by Nothofagus forest and other woody vegetation (Hall & McGlone, 2006), but waves of burning and then grazing by Māori and European (from ~1800s) settlers have progressively reduced this forest to small remnant patches of mainly Nothofagus cliffortioides. These remnants are now surrounded by native tussock grassland, retired pasture dominated by non-native grasses and Leptospermum scoparium and Discaria toumatou shrubland. The experiment was established within an approximately 330 × 130 m area of retired pasture on the north-west facing side of the Motatapu valley, 560 m a.s.l. Annual rainfall is c. 600-700 mm. At the time o...