Factors influencing brushtail possum (

Damage by introduced brushtail possums (Trichosurus vulpecula) to Pinus radiata trees was assessed in 41 compartments of a commercial forestry plantation on the Coromandel Peninsula, New Zealand. All the trees assessed were less than 3 years old. Possum damage in the compartments was low (median prevalence 3.3%) but highly variable (range 0?30%). Eight of 37 measured habitat factors differed significantly (P < 0.05) between the sites with damaged and undamaged trees. The best predictor of mean damage was stand age, but this explained only 21% of the variation in damage among compartments. Including both stand age and New Zealand bracken (Pteridium esculentum) cover improved the model significantly and explained 36% of variation in damage. Damage was apparently unrelated to compartment size, distance from the compartment boundary, and possum den-site availability. Surprisingly, the relationship between browse damage and a trap-catch index of possum abundance was weakly negative (r_S = −0.53, P = 0.05). The dense understorey associated with young pine stands tends to increase possum damage to associated P. radiata trees, but the possums in such stands may be less mobile at ground level and thus less easily trapped. Assessment of stand age and understorey characteristics, together with visual inspection for early signs of damage, is likely to be more cost-effective than possum surveys for identifying forest compartments at risk from possum browse.     

Thresholds in plant–herbivore interactions: predicting plant mortality due to herbivore browse damage

Patterns of herbivore browse at small scales, such as the rate of leaf consumption or plant preferences, drive the impact of herbivores on whole-plant processes, such as growth or survival, and subsequent changes in plant population structure. However, herbivore impacts are often non-linear, highly variable and context-dependent. Understanding the effect of herbivores on plant populations therefore requires a detailed understanding of the relationships that drive small-scale processes, and how these interact to generate dynamics at larger scales. We derive a mathematical model to predict annual rates of browse-induced tree mortality. We model individual plant mortality as a result of rates of foliage production, turnover and herbivore intake, and extend the model to the population scale by allowing for between-tree variation in levels of herbivore browse. The model is configurable for any broadleaved tree species subject to vertebrate or invertebrate browse, and is designed to be parameterized from field data typically collected as part of browse damage assessments. We parameterized and tested the model using data on foliage cover and browse damage recorded on kamahi trees (Weinmannia racemosa) browsed by possums (Trichosurus vulpecula) in New Zealand forests. The model replicated observed patterns of tree mortality at 12 independent validation sites with a wide range of herbivore densities and browse damage. The model reveals two key thresholds; in plant foliar cover, indicating when individual trees may be at high risk from browse-induced mortality, and in herbivore intake, leading to high rates of mortality across the whole population.     

Browse on mahoe and kamahi leaf-fall as a trigger for possum control

Introduced brushtail possums (Trichosurus vulpecula) are controlled over large parts of New Zealand to protect canopy trees. The condition of canopy trees is one of the cues used to trigger possum control, but selecting an indicator of canopy tree condition is difficult because many factors unrelated to possum browsing can affect canopy condition, and indices based on canopy scoring may not always quickly detect real changes in possum herbivory. We therefore investigated the usefulness of the percentage of fallen mahoe (Melicytus ramiflorus) and kamahi (Weinmannia racemosa) leaves browsed by possums (‘fallen-leaf browse’ or ‘FLB’) as a trigger for control aimed at protecting these tree species. We collected leaves falling from kamahi and mahoe trees every two months for two years at two study areas in the central North Island, one with initially high possum abundance (Oriuwaka) and the other with low possum abundance (Otupaka). We classified each of 92 384 leaves as either not-browsed, possum-browsed or insect-browsed. There was a strong and similar seasonal pattern in the mean number of fallen leaves per tree for both mahoe and kamahi at both study areas; fewest leaves fell in winter, and the most leaves fell in spring and early summer. Mahoe and kamahi FLB exhibited a similar seasonal pattern at both areas, being lowest in winter and highest in spring and early summer. FLB for both mahoe and kamahi declined following control of possums to low densities at Oriuwaka. The proportion of fallen mahoe and kamahi leaves browsed by possums was small compared with those browsed by insects or not browsed. We show that spring/early summer (i.e. September–December) is the best period for sampling FLB and that the mean FLB can be estimated with a CV of 20% if one trap is randomly placed under the canopy of each of 24 randomly located trees. However, CVs were much larger in other seasons and when possum abundance was low. We consider FLB to be a sensible trigger for initiating possum control when the objective of control is to protect canopy trees, but further work is needed to determine the relationships between possum abundance, FLB, canopy condition, and key tree demographic rates.     

Species‐ and site‐specific impacts of an invasive herbivore on tree survival in mixed forests

Invasive herbivores are often managed to limit their negative impact on plant populations, but herbivore density – plant damage relationships are notoriously spatially and temporally variable. Site and species characteristics (both plant and herbivore) must be considered when assessing the potential for herbivore damage, making it difficult to set thresholds for efficient management. Using the invasive brushtail possum Trichosurus vulpecula in New Zealand as a case study, we parameterized a generic model to predict annual probability of browse‐induced mortality of five tree species at 12 sites. We compared predicted and observed tree mortality for each species + site combination to establish herbivore abundance – tree mortality thresholds for each site on a single and combined tree species basis. Model results indicated it is likely that possum browse was the primary cause of all tree mortality at nine of the 12 species‐site combinations, allowing us to estimate site‐specific thresholds below which possum population numbers should be reduced and maintained to keep tree mortality under a predetermined level, for example 0.5% per year. The browse model can be used to set site‐ and species‐specific management action thresholds, and can be adapted easily for other plant or herbivore species. Results for multiple plant or herbivore species at a single site can be combined to create conservative, site‐wide management strategies, and used to: determine which sites will be affected most by changes in herbivore abundance; quantify thresholds for herbivore management; and justify expenditure on herbivore control.     

Inferring changes in foliar mass and area from foliage cover: A mechanistic model

Measuring foliar area or mass directly is destructive, and precludes long‐term, repeated observations of individual trees as they suffer or recover from foliar damage. Instead, foliage cover indices are often used as a proxy for foliar mass. Patterns of fluctuations in foliage cover indices can be used to infer qualitative changes in canopy health. However, foliage cover is not necessarily linearly related to foliar area or mass, and this may confound the detection of significant foliar damage, and comparisons of herbivore browse impacts between individual trees, tree species or sites. I derived a mechanistic model to quantify the relationship between foliar area or mass and foliage cover measured as the proportion of sky occluded by leaves. This one parameter model is close to linear for single‐tiered trees, but increasingly non‐linear for multi‐tiered trees. I compared the non‐linear model to a linear model using foliage cover data from an artificial defoliation experiment on two single‐tiered, sub‐canopy species and from simulated photographic images of single‐ and multi‐tiered canopies. The non‐linear model had lower errors than the linear model, and errors did not increase with foliage density (leaf area per unit area), variation (of leaf sizes within and between canopies) or leaf geometry. The non‐linear model can be easily parameterized from relatively low‐cost observations of foliage cover, independently of empirical measurements of foliar area or mass, and is applicable to a wide range of tree species. It should therefore help managers quantify how changes in foliage cover due to natural fluctuations or foliar damage affect foliar area and mass, and can be used to quantify parameters for models of browse impacts in mixed forest.     

Spatial variation in impacts of brushtail possums on two Loranthaceous mistletoe species

Browsing by introduced brushtail possums is linked to major declines in mistletoe abundance in New Zealand, yet in some areas mistletoes persist, apparently unaffected by the presence of possums. To determine the cause of this spatial variation in impact I investigated the abundance and condition (crown dieback and extent of possum browse) of two mistletoes (Alepis flavida, Peraxilla tetrapetala) and abundance and diet of possums in two mountain beech (Nothofagus solandri var. cliffortioides) forests in the central-eastern South Island of New Zealand. Mistletoe is common and there are long-established uncontrolled possum populations in both forests. Mistletoes were abundant (216?1359 per hectare) and important in possum diet (41?59% of total diet), but possum density was low (c. 2 per hectare) in both areas. Possum impacts were slight with low browse frequencies and intensities over much of the study sites. However, impacts were significantly greater at a forest margin, where possum abundance was highest, and at a high-altitude site where mistletoe density was lowest. Mistletoe crown dieback was inversely proportional to intensity of possum browsing. These results suggest that the persistence of abundant mistletoe populations at these sites is due to mistletoe productivity matching or exceeding consumption by possums in these forests of low possum-carrying capacity, rather than low possum preference for the local mistletoe populations.     

Diet and impacts of brushtail possum populations across an invasion front in South Westland, New Zealand

Impact of the irruptive fluctuation in abundance of brushtail possum populations since their initial colonisation was investigated in the forests of South Westland, New Zealand. Possum abundance, fecundity, and diet, the condition of common possum-palatable tree species, and the abundance of common forest birds were measured at three sites occupied by possums for c. 10, 20, and 30 years. Possum densities were highest at the site where possums had been present for c. 20 years. Possum fecundity was higher at the site they had recently colonised than at the two sites where they had been present for c. 20–30 years. Diet of possums where they had recently colonised was dominated by highly preferred foods (72% of total diet), whereas these same foods contributed just 36% of total diet at the site where possums had been present for more than two decades. Canopy condition of common possum-preferred trees was scored progressively lower in areas with increasing length of possum occupation, especially at the site where the possum population had apparently declined from its maximum density. Native forest bird abundance also declined with increasing length of possum occupation. The implications of these results for management are discussed.     

Long‐term response of temperate canopy trees to removal of browsing from an invasive arboreal herbivore in New Zealand

Defoliation of forest tree canopies by herbivores and other agents, leading to tree mortality and reduced productivity, threatens the ecological stability of forests globally. This study shows that long‐term control of a mammalian arboreal folivore (brushtail possums; Trichosurus vulpecula Phalangeridae) reduces crown dieback and increases foliage cover in browsing‐damaged canopy trees. We monitored indices of possum density, possum browsing, tree foliage cover and crown dieback for 20 years following initiation of possum control in 1994 that repeatedly reduced possum densities to near zero every 5–6 years and kept the population below 35% of pre‐control levels over the entire period. Observable possum browsing was recorded on 20–49% of individuals of three palatable tree species at the time of first control. Those percentages fell to zero after control and never exceeded 2–10% for individual species over the next 19 years. We recorded significant increases in foliage cover attributable to recovery from defoliation by possums for all three species during the first 10 years. Large increases in foliage cover occurred on individuals that were heavily browsed in 1994 (mean increases: 36–89%), but mean population increases were modest (3–19%) because only 10–19% of trees were initially heavily browsed. Twenty‐year mortality rates were similar for plants with, or without, initial possum browsing, indicating no residual impact of pre‐control browsing on tree mortality. Times for full recovery of crown foliage cover varied from 10 years for the youngest trees and faster growing species to more than 20 years for mature individuals of the slowest growing species.     

Individual tree traits shape insect and disease damage on oak in a climate‐matching tree diversity experiment

Diversifying planted forests by increasing genetic and species diversity is often promoted as a method to improve forest resilience to climate change and reduce pest and pathogen damage. In this study, we used a young tree diversity experiment replicated at two sites in the UK to study the impacts of tree diversity and tree provenance (geographic origin) on the oak (Quercus robur) insect herbivore community and a specialist biotrophic pathogen, oak powdery mildew. Local UK, French, and Italian provenances were planted in monocultures, provenance mixtures, and species mixes, allowing us to test whether: (a) local and nonlocal provenances differ in their insect herbivore and pathogen communities, and (b) admixing trees leads to associational effects on insect herbivore and pathogen damage. Tree diversity had variable impacts on foliar organisms across sites and years, suggesting that diversity effects can be highly dependent on environmental context. Provenance identity impacted upon both herbivores and powdery mildew, but we did not find consistent support for the local adaptation hypothesis for any group of organisms studied. Independent of provenance, we found tree vigor traits (shoot length, tree height) and tree apparency (the height of focal trees relative to their surroundings) were consistent positive predictors of powdery mildew and insect herbivory. Synthesis. Our results have implications for understanding the complex interplay between tree identity and diversity in determining pest damage, and show that tree traits, partially influenced by tree genotype, can be important drivers of tree pest and pathogen loads.     

Predicting Summer Site Occupancy for an Invasive Species,the Common Brushtail Possum (Trichosurus vulpecula), in an Urban Environment

Invasive species are often favoured in fragmented, highly-modified, human-dominated landscapes such as urban areas. Because successful invasive urban adapters can occupy habitat that is quite different from that in their original range, effective management programmes for invasive species in urban areas require an understanding of distribution, habitat and resource requirements at a local scale that is tailored to the fine-scale heterogeneity typical of urban landscapes. The common brushtail possum (Trichosurus vulpecula) is one of New Zealand’s most destructive invasive pest species. As brushtail possums traditionally occupy forest habitat, control in New Zealand has focussed on rural and forest habitats, and forest fragments in cities. However, as successful urban adapters, possums may be occupying a wider range of habitats. Here we use site occupancy methods to determine the distribution of brushtail possums across five distinguishable urban habitat types during summer, which is when possums have the greatest impacts on breeding birds. We collected data on possum presence/absence and habitat characteristics, including possible sources of supplementary food (fruit trees, vegetable gardens, compost heaps), and the availability of forest fragments from 150 survey locations. Predictive distribution models constructed using the programme PRESENCE revealed that while occupancy rates were highest in forest fragments, possums were still present across a large proportion of residential habitat with occupancy decreasing as housing density increased and green cover decreased. The presence of supplementary food sources was important in predicting possum occupancy, which may reflect the high nutritional value of these food types. Additionally, occupancy decreased as the proportion of forest fragment decreased, indicating the importance of forest fragments in determining possum distribution. Control operations to protect native birds from possum predation in cities should include well-vegetated residential areas; these modified habitats not only support possums but provide a source for reinvasion of fragments.     

The impact of brodifacoum on non-target wildlife: gaps in knowledge

To assess the effect of possum browse on plant growth, an index of the amount of foliage on about 50 trees of Fuchsia excorticata and the number of trees that died or were completely defoliated was measured at five sites in South Westland over 5 years. This index was compared to possum density indices taken at each site each year. At one site, possums were reduced from a high density about 6 months before the final measurement. The degree of defoliation of fuchsia was significantly related to the density of possums at each site. Possums defoliate and kill fuchsia within two years once the catch-rate index of their density exceeds about 25%. However, this impact can be halted and quickly reversed by reducing possum density.     

Opposing deer and caterpillar foraging preferences may prevent reductions in songbird prey biomass in historically overbrowsed forests

Overbrowsing by ungulates decimates plant populations and reduces diversity in a variety of ecosystems, but the mechanisms by which changes to plant community composition influence other trophic levels are poorly understood. In addition to removal of avian nesting habitat, browsing is hypothesized to reduce bird density and diversity through reduction of insect prey on browse‐tolerant hosts left behind by deer. In this study, we excluded birds from branches of six tree species to quantify differences in songbird prey removal across trees that vary in deer browse preference. Early in the breeding season, birds preyed on caterpillars at levels proportional to their abundance on each host. Combining these data with tree species composition data from stands exposed to experimentally controlled deer densities over 30 years ago, we tested whether overbrowsing by white‐tailed deer reduces prey biomass long after deer densities are reduced. Our analysis predicts total prey availability in the canopy of regenerating forests is fairly robust to historic exposure to high deer densities, though distribution of prey available from host species changes dramatically. This predicted compensatory effect was unexpected and is driven by high prey abundance on a single host tree species avoided by browsing deer, Prunus serotina. Thus, while we confirm that prey abundance on host trees can act as a reliable predictor for relative prey availability, this study shows that quantifying prey abundance across host trees is essential to understanding how changes in tree species composition interact with ungulate browse preference to determine prey availability for songbirds.     

Possum (

To assess the effect of possum browse on plant growth, an index of the amount of foliage on about 50 trees of Fuchsia excorticata and the number of trees that died or were completely defoliated was measured at five sites in South Westland over 5 years. This index was compared to possum density indices taken at each site each year. At one site, possums were reduced from a high density about 6 months before the final measurement. The degree of defoliation of fuchsia was significantly related to the density of possums at each site. Possums defoliate and kill fuchsia within two years once the catch-rate index of their density exceeds about 25%. However, this impact can be halted and quickly reversed by reducing possum density.     

The abundance,distribution and structural characteristics of tree‐holes in Nothofagus forest,New Zealand

Tree‐holes provide an important microhabitat that is used for feeding, roosting and breeding by numerous species around the world. Yet despite their ecological importance for many of New Zealand's endangered species, few studies have investigated the abundance or distribution of tree‐holes in native forests. We used complementary ground and climbed tree surveys to determine the abundance, distribution and characteristics of tree‐holes in undisturbed Nothofagus forest in the Lewis Pass, New Zealand. We found that hole‐bearing trees were surprisingly abundant compared with many other studies, including Australian Eucalyptus species and American beech. In fact, we estimated as many as 3906 tree‐holes per hectare, of which 963 holes per hectare were potentially large enough to provide roost sites for hole‐nesting bats in New Zealand, while only eight holes per hectare were potentially suitable for specialist hole‐nesting birds. This was of great interest as primary cavity‐excavating animals are absent from New Zealand forests, compared with North America and Australia. Moreover, tree‐hole formation in New Zealand is likely to be dominated by abiotic processes, such as branch breakage from windstorms and snow damage. As has been found in many other studies, tree‐holes were not uniformly distributed throughout the forest. Tree‐holes were significantly more abundant on the least abundant tree species, Nothofagus fusca, than on either N. menziesii or N. solandri. In addition to tree species, tree size was also an important factor influencing the structural characteristics of tree‐holes and their abundance in this forest. Moreover, these trends were not fully evident without climbed tree surveys. Our results revealed that ground‐based surveys consistently underestimated the number of tree‐holes present on Nothofagus trees, and illustrate the importance of using climbed inspections where possible in tree‐hole surveys. We compare our results with other studies overseas and discuss how these are linked to the biotic and abiotic processes involved in tree‐hole formation. We consider the potential implications of our findings for New Zealand's hole‐dwelling fauna and how stand dynamics and past and future forest management practices will influence the structural characteristics of tree‐holes and their abundance in remnant forest throughout New Zealand.     

Effects of habitat fragmentation on the three-way interaction among ants,aphids and larvae of the giant purple emperor, <Emphasis Type="Italic">Sasakia charonda</Emphasis> (Hewitson), a near-threatened butterfly

Survival rates of both early and middle instar larvae of the nymphalid butterfly, Sasakia charonda, were estimated to be lowest on test trees planted in a meadow (site A), intermediate in a small, narrow secondary deciduous broadleaf forest (small patch, site B) and highest in a large secondary deciduous broadleaf forest (large forest, site C). The larval mortality rates due to predation by tree-climbing predators from the ground (tree climbing predator) such as ants and the larvae of carabids were estimated to be greater at sites A and B than those at site C. The number of predatory ants climbing test trees was significantly greater at sites A and B than at site C, and the ants harvested honeydew from aphids living on tree leaves at those two sites. Aphid densities were significantly higher on trees at sites A and B than at site C, and aphid densities and numbers of predatory ants were significantly and positively correlated at sites A and B. In an experiment controlling aphid density per branch on test trees, the numbers of ants and the mortality rates of S. charonda larvae were greater on branches with high aphid densities than on those with low aphid densities at both sites A and B. These results suggest that the aphid density per host tree was higher in the meadow and the small patch than in the large forest; at both sites these higher aphid densities attracted higher numbers of predatory ants to test trees, and as a result, mortality rates of S. charonda larvae were increased.     

How Herbivore Browsing Strategy Affects Whole-Plant Photosynthetic Capacity

If a browse damage index indicates that a tree has been 50% browsed by herbivores, does this mean half the leaves are entirely eaten or are all the leaves half eaten? Were the affected leaves old or young? Large or small? In sunshine or shade? Understanding what effect browsing will have on the photosynthetic capacity and the plant’s survival ability clearly requires a greater understanding of browsing strategy across the canopy than can be given by a single index value. We developed stochastic models of leaf production, growth and consumption using data from kamahi (Weinmannia racemosa) trees in New Zealand which have been browsed by possums (Trichosurus vulpecula), to ascertain which of six feasible browsing strategies possums are most likely to be employing. We compared the area distribution of real fallen leaves to model output in order to select the best model, and used the model to predict the age distribution of leaves on the tree and thus infer its photosynthetic capability. The most likely browsing strategy that possums employ on kamahi trees is a preference for virgin (i.e. previously unbrowsed) leaves, consistent with the idea that browsing increases the production of chemical plant defences. More generally, our results show that herbivore browsing strategy can significantly change the whole-plant photosynthetic capability of any plant and hence its ability to survive, and therefore, herbivore damage indices should be used in conjunction with more detailed information about herbivore browsing strategy.     

Mistletoe (

The condition of 79 plants of the loranthaceous mistletoe Tupeia antarcticain a podocarp-hardwood forest in the central North Island, New Zealand, was monitored over 4 years during a period of increasing possum density, following previous possum control. Mistletoe comprised 1.2% of total possum diet during the three years following possum control. Incidence of possum browse on mistletoe plants increased from 2.6% of plants when the trap-catch index of possum density was < 3%, to 75.9% of plants when trap-catch rates reached 4.6%. Mistletoe foliage cover declined from 49.8% to 15.6% and mean plant size declined by about 55% over the same period. The mistletoe population was dominated by plants with large haustoria, located in heavily shaded locations in the lower crown of their Carpodetus serratus hosts. Most plants established more than 20 years ago, and the current potential for recruitment of new individuals into the population is severely limited by possum browsing and the senescent nature of the mistletoe population. Intensive management of host crowns and possum populations will be necessary to ensure the long-term viability of mistletoe at the study site.     

Observations on the Ecology of Weaver Ants (Oecophylla smaragdina Fabricius) in a Thai Mangrove Ecosystem and Their Effect on Herbivory of Rhizophora mucronata Lam.

Ants of the genus Oecophylla are predators of other insects and are able to protect a variety of terrestrial plants against pest insects; however, observations on the ecology of these ants in mangrove forests are lacking. General observations on the ecology of Oecophylla smaragdina were carried out in a Thai mangrove forest to determine if these ants can protect their host plants in less favorable mangrove habitats. Leaf herbivory and the density of O. smaragdina ants were measured on Rhizophora mucronata trees at two sites. The results showed a negative correlation between ant density and herbivory. At both sites, the mean percent damaged leaf area was more than four times higher on trees without ants compared to “ant‐trees.” A significant negative correlation was found between tree mean percent leaf damage and the density of ants on the tree. Furthermore, on trees with ants, there was less herbivory on leaves close to ant nests compared to other leaves on the tree. Most damage was caused by chrysomelid beetles (62%) and sesarmid crabs (25%) and both types of herbivory were significantly reduced on ant‐trees.     

Long-term changes and seasonal patterns in possum (

Possum leaf diet from 1976 to 1989 in the Orongorongo Valley was compared with the diet recorded from four previous years, using faecal analysis. There were large differences in the proportions of species eaten in different seasons and between different years. There was little overall change in the proportion of the main diet tree species, Metrosideros robusta and Weinmannia racemosa, and seasonal increases or declines in one were usually balanced by opposite changes in the other. However, the results of a study of forest composition in the same area showed that living stems of both species declined substantially in abundance, the former mostly before 1978. Melicytus ramiflorus maintained both an important contribution to the diet and its stem density in the forest. The lianes Metrosideros fulgens and Ripogonum scandens both increased their proportion of the diet, but no data were available on their abundance in the study area. At present browse levels, Weinmannia racemosa will become extinct locally, and Metrosideros robusta more rare, especially if drought and higher than average possum densities coincide. Given their large contribution to possum diet, the decline of these species will have a significant effect on possum feeding habits, including increased browsing pressure on Melicytus ramiflorus, Ripogonum scaldens and Metrosideros fulgens. The trend for formerly less palatable tree species to replace those previously important in the diet will continue, with consequent changes in forest species composition.     

Community dynamics of evergreen broadleaf forests in southwestern Japan

The process and rate of revegetation in gaps in an evergreen oak forest were studied by comparing the species composition, tree density, frequency distribution of tree height, and relation between diameter at breast height and tree height among different aged stands. For estimating stand ages, the ages of gap indicators, such as,Symplocos prunifolia andAcer rufinerve, were very useful. It took about 70 years for gaps to be filled by large fully-grown trees. Since the mean residence time of the forest canopy was 180 years, the trees that attain the forest canopy were expected to be canopy trees for 110 years on the average. Tree densities of all broadleaved evergreens exceptS. prunifolia, were independent of stand age. On the other hand, densities of gap indicators,S. prunifolia andA. rufinerve, decreased as stand age increased. Other deciduous broadleaf and coniferous species were scarce as a whole. According to the frequency distributions of height of live and dead trees in different aged stands, it was suggested that shorter trees were more susceptible to death than taller trees. The self-thinning in revegetation process in gaps approximately followed the 3/2 power law, though the power was larger (−1.32) than expected from the law.