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Summary1. Invasive browsing ungulates can have strong impacts on the structure and composition of forest ecosystems, particularly where ungulates are not native ecosystem components as in New Zealand. Ungulate impacts on plant communities have been considered mostly from an above-ground perspective. However, understanding below-ground effects of these invasive herbivores is critical as they may drive feedbacks to above-ground ecosystem components. 2. We measured growth responses of seedlings of five common tree species in a greenhouse experiment in soils collected from 26 plots fenced to exclude invasive ungulates for at least 17 years and from paired, unfenced control plots. We then further investigated soil-mediated effects of ungulates on one tree species, Melicytus ramiflorus, by partitioning these effects into soil abiotic and biotic components, as well as measuring arbuscular mycorrhizal fungal (AMF) root infection. 3. Biomass of seedlings of all five species was greater in soils from within exclosures, although this was only significant for two species. These soil-mediated effects were partially driven by changes in physical and chemical soil properties; soil bulk densities were lower inside exclosures than in controls. 4. Effects of invasive ungulates on seedling biomass of M. ramiflorus were positively related to effects on per cent AMF root infection. The biomass of M. ramiflorus seedlings was positively related to the AMF infection of its roots, which in turn was related to greater organic matter content and lower bulk density of soils from within exclosures. Results for M. ramiflorus indicated that soilmediated effects of ungulates on seedling biomass were of abiotic origin, but were mediated by the biotic soil component, that is, through effects on AMF. 5. Synthesis. Invasive herbivores may potentially impact on plant performance and community structure not only directly but also indirectly through influencing soil abiotic and biotic properties. Our results show that shifts in plant-soil interactions and feedbacks represent important but understudied pathways by which invasive ungulates can have wide-ranging impacts on forest ecosystems. Future studies should consider the importance of soil-mediated effects of invasive ungulates relative to direct effects of herbivory.

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Browsing of mountain beech seedlings by introduced deer in the central North Island of New Zealand appears to have inhibited canopy regeneration over large areas. In 1998, a trial of high-, medium- and low-intensity deer-culling treatments was initiated in Kaimanawa and Kaweka Forest Parks to test whether mountain beech (Nothofagus solandri var. cliffortioides) forest regeneration could be restored by removing deer. Paired exclosure plots (one plot fenced to exclude deer and the other unfenced) were established within a high-intensity culling area, to monitor the benefits of recreational, commercial and aerial deer culling. Paired plots were also established within low- and medium-intensity culling areas. Medium-intensity culling was allowed through recreational and commercial deer culling. In two low-intensity culling treatment areas, deer management remained substantially unaltered. Annual relative growth rates of tagged seedlings from spring 1998 to spring 2001 from low-, medium- and high-intensity culling areas provide strong evidence that mountain beech seedling growth increases once browsing by deer is removed through fencing. Faecal pellet data indicated that high-intensity deer culling reduced deer abundance by 67% in comparison to medium- and low-intensity culling areas. This apparent reduction in deer abundance appears to have led to a doubling in mountain beech seedling growth in the high-intensity culling area outside fences, in comparison to low- and medium-intensity deer culling areas where there was little or no evidence of benefits for seedling growth.

The effect of herbivory and resource availability on the competitive ability of different plant species has been an area of intense debate amongst plant ecologists for at least two decades, but the interactive effects of herbivory and plant competition between woody and herbaceous plants are rarely studied and theory is poorly developed. This study used experimental manipulations on transplanted and naturally occurring mountain beech (Nothofagus solandri var. cliffortioides) seedlings to show the effects of deer browsing and competition from deer-induced, herbaceous turf communities on mountain beech regeneration in New Zealand. Differences in the species composition of these turfs had little effect on mountain beech seedling establishment, but turf removal increased seedling growth and survivorship, showing that competition with other plants had direct effects on mountain beech regeneration. Deer browsing reduced the establishment and growth of seedlings, but the size of this effect did not vary with light and nutrient availability. There was no immediate compositional response of turf communities following the removal of deer browsing. The addition of nutrients appeared to reduce the intensity of belowground competition (stem growth increased relative to root growth) and increase seedling mortality, but there was no effect of changing levels of light. These results showed simple and direct negative effects of deer browsing on mountain beech regeneration. Indirect negative effects on regeneration were caused by deer-induced turf communities. We found little evidence for interactive effects between herbivory, plant competition and the availability of light or nutrients on seedling regeneration, which suggests that these factors acted independently.

We compared the impacts on forest regeneration of introduced sika (Cervus nippon) and red (Cervus elaphus) deer in New Zealand. Plot data were used to compare mountain beech (Nothofagus solandri var. cliffortioides) regeneration between a region with sika deer, and four regions without sika deer. All regions surveyed had red deer present. In the region where sika deer had been present for more than a decade, there was evidence of poor mountain beech seedling regeneration. In the four regions without sika deer, there was evidence of a strong regenerative response at stands with low occupancy by trees. When compared to larger deer species, sika deer have a digestive morphology allowing greater dietary versatility, which may result in them impeding forest regeneration where red deer do not. In contrast to mountain beech, some small-leaved shrub species may have been competitively advantaged by intensive browsing from sika deer. This is contra to a current view that small-leaved shrub species with interlacing branches were able to tolerate browsing from extinct ratite birds, but not introduced deer. Sika deer have been introduced into countries where other deer species are indigenous, such as Canada, Denmark, Great Britain, Czech Republic, Ireland and the United States of America. Because of their dietary advantage, sika deer may have a greater potential to impede forest regeneration and competitively exclude larger deer species, particularly at low basal area sites where impacts on tree regeneration are likely to be greatest.

Background: Red needle cast caused by Phytophthora pluvialis Reeser, Sutton & E. Hansen, and less frequently P. kernoviae Brasier, Beales & S.A.Kirk, is an important foliar disease of Pinus radiata D.Don (radiata pine) in plantations throughout parts of New Zealand. Significant growth loss occurs following years when severe outbreaks occur. Aerial spraying with a copper-based fungicide has potential for disease control. Research is being carried out to optimise application timing, supported by complementary studies to understand RNC epidemiology. Methods: In order to determine the pathogen infection periods, a field trial was conducted over two years at two forests in the Central North Island of New Zealand. Batches of potted radiata pine seedlings were placed beneath diseased pine stands at fortnightly intervals, before returning them to an open nursery area for assessments of infection every two weeks (based on visual symptoms and qPCR) over a period of three months. A hybrid modelling approach was employed to establish relationships between the proportion of plants showing symptoms and weather conditions during the fortnight of exposure and previous fortnights. Gradient boosting machine learning analyses were used to identify the most important weather variables, followed by analysis of these by generalised mixed effects models, generalised least square models and ordinary least square models. Results: Development of RNC symptoms and detection of Phytophthora pluvialis and P. kernoviae on exchange seedlings was greatest for those exposed between April and September (Southern Hemisphere mid-autumn to early-spring). At this time, temperatures, solar radiation and evapotranspiration were lower, and rainfall and foliage wetness were plentiful. Modelling identified temperature and relative humidity several months before the date of exposure as the most important weather variables explaining infection. Conclusions: Because of autocorrelation, it was not possible to determine those variables that drive sporulation, dispersal, infection and symptom development. This will require more detailed exchange plant studies together with controlled environment inoculation experiments. Nevertheless, results of this and earlier work complement recent research indicating that it may be possible to manage RNC by fungicide applications made in late summer or autumn, early in the annual disease cycle.