Magnitude of Canopy Dieback and Implications for Conservation of Southern Rata-Kamahi (

The amount of conspicuous canopy dieback in all central Westland southern rata-kamahi forests east of the Alpine Fault, between 500 m altitude and treeline, was assessed and mapped from aerial photographs taken in 1984-85 and verified by aerial reconnaissance of selected areas in 1988. At least 20% of all canopy trees, predominantly southern rata (Metrosideros umbellata) and Hall's totara (Podocarpus hallii), were dead in 1984-85. Major catchments with their headwaters east of the Alpine Fault comprised 78% of the study area and were worst affected (5%-44% canopy mortality). Because affected trees decay and eventually disappear from the canopy, the extent of visible dieback underestimated total canopy depletion, particularly where mortality occurred more than c. 15 years ago. Geographical variation in canopy dieback reflected the intensity and duration of browsing by the introduced brush- tailed possum (inferred from patterns of invasion from their liberation sites) and the influences of forest composition and timing of dieback. Although 29% of the forests showed light dieback (<10% mortality), only 11% had not experienced heavier past dieback and could be classified as having canopies largely unmodified by possums. Such widespread and continuing forest depletion, and declining possum control effort over the last decade, indicate the urgent need for a coordinated rata-kamahi forest conservation strategy, involving long-term possum control and monitoring in representative tracts.     

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.     

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.     

Brushtail possum (

We quantified brushtail possum (Trichosurus vulpecula) diet in a mixed Nothofagus fusca-N. menziesii forest in north Westland. Diet comprised 49 food items of which four (Aristotelia serrata, Muehlenbeckia australis and Weinmannia racemosa foliage, and W. racemosa flowers) contributed 68%. The canopy dominant Nothofagus species were a minor diet component (<1%), while wood, fungi and bark were a small but consistent part of diet (10.1%). Our results are similar to previous possum diet studies in Nothofagus forests and suggest that possums are very selective in their feeding, both spatially and temporally, focusing on key preferred species in particular parts of the forest and taking advantage of different food types that become available at different times of the year.     

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.     

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 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.     

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.     

Forest decline and dieback - A global ecological problem

Destruction of tropical forests has gained world-wide attention, and the associated depletion of biodiversity has become a growing concern. Almost simultaneously, forest decline and dieback in the developed countries on both sides of the North Atlantic has become an international worry. Industrial pollution has been implicated as the major cause, but the etiology of forest decline and dieback remains largely unresolved. Decline and dieback can also be related to biotic impoverishment in the sense of reduction of canopy species diversity, simplification of structural diversity and biological mining of soil nutrients. Thus, concern over the depletion of biodiversity is highly justified. It links tropical forest destruction to forest dieback in Europe in a way not yet widely realized. At the same time, it does not diminish tge equally justified concern about industrial pollution. A world-wide internationally coordinated research program is suggested which focuses on comparative, interdisciplinary studies in those countries where forest decline and dieback has surfaced as an unresolved problem.     

Canopy structures and its effect on shoot growth and flowering in subalpine forests

Changes in foliage density distribution with altitude and its effect on shoot growth and flowering were studied in forest section planes (profiles) of subalpine forests and scrubs (Krummholz) in Nepal and Japan.Patterns of foliage in forest canopy were evaluated by an analysis of variance. Foliage densities were very high at high altitude canopies, but the change in spatial patterns along altitude differs in both areas.The spatial pattern of new shoot production was similar to that of current foliage density and was affected by the amount of foliage above the sample probably through light condition. Flowering of tall trees occurred in the surface of the upper canopy, but a shrub species flowered even under tree canopies.     

Microclimatic buffering in forests of the future: the role of local water balance

Forest canopies buffer climate extremes and promote microclimates that may function as refugia for understory species under changing climate. However, the biophysical conditions that promote and maintain microclimatic buffering and its stability through time are largely unresolved. We posited that forest microclimatic buffering is sensitive to local water balance and canopy cover, and we measured this effect during the growing season across a climate gradient in forests of the northwestern United States (US). We found that forest canopies buffer extremes of maximum temperature and vapor pressure deficit (VPD), with biologically meaningful effect sizes. For example, during the growing season, maximum temperature and VPD under at least 50% forest canopy were 5.3°C and 1.1 kPa lower on average, respectively, compared to areas without canopy cover. Canopy buffering of temperature and vapor pressure deficit was greater at higher levels of canopy cover, and varied with water balance, implying that buffering effects are subject to changes in local hydrology. We project changes in the water balance for the mid‐21st century and predict how such changes may impact the ability of western US forests to buffer climate extremes. Our results suggest that some forests will lose their capacity to buffer climate extremes as sites become increasingly water limited. Changes in water balance combined with accelerating canopy losses due to increases in the frequency and severity of disturbance will create potentially non‐linear changes in the microclimate conditions of western US forests.     

Tropical deciduous forest in Yunnan,southwestern China: Implications for geological and climatic histories from a little-known forest formation

In the southern mountain ranges of Yunnan province, China, deep valleys of several large rivers create rain shadows with hot dry summers, and are locally designated tropical; towards the north, notably in the Lancang (Upper Mekong) valley, these regions may experience frost during winter. The woody forest canopy of these valleys is predominantly deciduous, with evergreen elements in the north, where the canopy is open and the forest savanna-like. However, we here present tall forest with a closed deciduous canopy and semi-evergreen subcanopy observed in hot dry valleys of these rivers and their tributaries in the tropical south. The structure and physiognomy of these forests resemble the tall (moist) deciduous forest formation widespread in South Asia and Indo-Burma. Furthermore, these forests are largely composed of tropical elements at both the generic (80%) and the species level (>70%), indicating that these forests are indeed tropical. We originally hypothesized that these isolated forests represent refugia of a pre-Holocene extension of tall (moist) deciduous forest formation of South Asia and Indo-Burma. The sample plot we established to test this hypothesis confirmed that these forests share the structure and physiognomy of the tall (moist) deciduous forest formation; however, the plots also showed that these forests lack the characteristic and dominant species of the formation''s Indo-Burmese range. The tree flora, in particular, indicates that both deciduous and evergreen elements are instead mostly derived from the adjacent tropical semi-evergreen forests of tropical southern China; yet they also include an important endemic element, which implies that these forests have survived as refuges possibly since the Pliocene. The exceptional representation of evergreen elements in these forests indicates that they have rarely been subject to hot fires or domestic cattle browsing, adding to the unique nature of the forests and further justifying their strict conservation.     

Invasive gorse (<Emphasis Type="Italic">Ulex europaeus</Emphasis>, Fabaceae) changes plant community structure in subtropical forest–grassland mosaics of southern Brazil

Gorse (Ulex europaeus L., Fabaceae) is an invasive species of worldwide concern. However, knowledge of its effects on ecosystem and community structure is still missing from most areas of its invasive range, including species-rich ecosystems such as the forest–grassland mosaics of southern Brazil. We studied the influence of landscape mosaic structure and land use on gorse distribution. At the community scale, we examined the effects of proximate gorse canopies on plant community abundance, diversity, and composition. Vulnerability to gorse invasions was highest in heavily grazed grasslands, where gorse canopy cover often reached 50 %. Gorse did not invade natural forests. However, gorse invaded tree plantations. Gorse cover in relatively undisturbed areas, such as ungrazed natural grasslands, was found to be variable, although the physiognomy and structure of mosaic ecosystems was still affected. Gorse canopies exerted significant effects on community structure and composition. Assemblages in the understory of gorse canopies were comparatively species-poor and more homogeneous than beside gorse canopies. Woody species were more abundant in the understory of gorse canopies than in areas beside gorse canopies, whereas the cover of grasses decreased under gorse canopies. We argue that invasions can differentially affect the components of natural forest–grassland mosaics, which could have long-term consequences. Selecting competitive species for restoring native assemblages can provide successful outcomes in managing risks associated to invasive species like gorse.     

Differential susceptibility to Eucalyptus secondary compounds explains feeding by the common ringtail (Pseudocheirus peregrinus) and common brushtail possum (Trichosurus vulpecula)

The effect of two plant secondary metabolites, tannins and formylated phloroglucinol compounds (FPCs), on the intake of Eucalyptus foliage by common ringtail (Pseudocheirus peregrinus) and common brushtail possums (Trichosurus vulpecula) was studied. We manipulated the amount of tannin that was free to bind with protein by coating foliage with polyethylene glycol 4000 (PEG) and relied on natural intraspecific variation in FPC concentrations. In contrast to ringtail possums, brushtail possums showed a greater tolerance to FPCs and ate more foliage when it was coated with PEG, suggesting that tannins limited their food intake. Brushtails detected the effects of tannins through immediate oral sensations rather than through systemic effects. Ringtail possums appeared highly tolerant of foliar tannins yet susceptible to low concentrations of FPCs. We could not detect any interaction between tannins and FPCs that affected the intake of Eucalyptus foliage by either species of possum. Although ringtail and brushtail possums are widely regarded as specialist and generalist folivores, respectively, their differential susceptibility to co-occurring secondary metabolites suggests greater complexity. Each possum species appears to be a specialist in its own right, which leads to a partitioning of available foliage. Brushtails avoid tannins and ringtails avoid FPCs.     

Limited ecological connectivity of an arboreal marsupial across a forest/plantation landscape despite apparent resilience to fragmentation

Demographic and genetic replenishment of populations through the exchange of individuals is essential for their persistence. Habitat loss and fragmentation can reduce the permeability of landscapes, hinder dispersal and compromise the genetic integrity of populations over time. We examined ecological connectivity in an arboreal marsupial, the common ringtail possum (Pseudocheirus peregrinus) in fragmented forests of southeastern Australia. This species is potentially robust to fragmentation based on its presence in degraded landscapes and known use of plantations for foraging and nesting. Using 312 individuals screened at 15 microsatellites, we measured dispersal and gene flow across seven native Eucalyptus forest remnants surrounded by exotic Pinus radiata plantations and three sites within a large continuous forest. The permeability of the pine matrix to dispersing possums was significantly lower than that of the native forest. Small, isolated patches exhibited signatures of genetic drift, having lower heterozygosity and allelic richness than possums in large patches. Most (87%) possums were born in their sampled patch or dispersed only short distances into neighbouring remnants. The continuous forest was identified as an important source of immigrants only for proximate patches (within 2.5 km), thus emphasizing for the common ringtail possum and more specialized arboreal mammals the need to conserve large, proximate forest remnants. Our findings highlight the importance of using genetic tools to understand the long-term biological consequences of fragmentation for effective management.     

The contribution of cacao agroforests to the conservation of lower canopy ant and beetle diversity in Indonesia

The ongoing destruction of tropical rainforests has increased the interest in the potential value of tropical agroforests for the conservation of biodiversity. Traditional, shaded agroforests may support high levels of biodiversity, for some groups even approaching that of undisturbed tropical forests. However, it is unclear to what extent forest fauna is represented in this diversity and how management affects forest fauna in agroforests. We studied lower canopy ant and beetle fauna in cacao agroforests and forests in Central Sulawesi, Indonesia, a region dominated by cacao agroforestry. We compared ant and beetle species richness and composition in forests and cacao agroforests and studied the impact of two aspects of management intensification (the decrease in shade tree diversity and in shade canopy cover) on ant and beetle diversity. The agroforests had three types of shade that represented a decrease in tree diversity (high, intermediate and low diversity). Species richness of ants and beetles in the canopies of the cacao trees was similar to that found in lower canopy forest trees. However, the composition of ant and beetle communities differed greatly between the agroforest and forest sites. Forest beetles suffered profoundly from the conversion to agroforests: only 12.5% of the beetle species recorded in the forest sites were also found in the agroforests and those species made up only 5% of all beetles collected from cacao. In contrast, forest ants were well represented in agroforests, with 75% of all species encountered in the forest sites also occurring on cacao. The reduction of shade tree diversity had no negative effect on ants and beetles on cacao trees. Beetle abundances and non-forest ant species richness even increased with decreasing shade tree diversity. Thinning of the shade canopy was related to a decrease in richness of forest ant species on cacao trees but not of beetles. The contrasting responses of ants and beetles to shade tree management emphasize that conservation plans that focus on one taxonomic group may not work for others. Overall ant and beetle diversity can remain high in shaded agroforests but the conservation of forest ants and beetles in particular depends primarily on the protection of natural forests, which for forest ants can be complemented by the conservation of adjacent shaded cacao agroforests.     

Phytophagous insect communities in the canopies of three Eucalyptus forest types in south-eastern Australia

Intensive sampling of tree canopies for phytophagous insects was carried out in three contrasting eucalypt forest types comprised of species widely distributed in sub-alpine forests in Victoria and New South Wales (Eucalyptus delegatensis, E. dives, and E. pauciflora). The number of phytophagous insects present in the canopies of these forest types was low, with a seasonal average of 20 individuals per kg of foliage (dry weight). Numbers were much lower than expected from past literature reporting‘chronically high’levels of defoliation in eucalypt forests. Microlepidoptera, Geometridae, Chrysomelidae, and Curculionidae were the major leaf-chewing groups recorded. Most sap-feeders were either leafhoppers (Cercopidae and Cicadellidae) or in the superfamily Fulgoroidea. Psyllidae and gall-making species were rare. Leafhoppers made up a very large portion of the phytophagous insect communities in each forest type, particularly in the E. dives forest. Microlepidoptera was the most commonly encountered defoliator group in all three forest types making up 33-44% of the total count. Non-phytophagous arthropods accounted for 44-48% of all individuals encountered. The density of insect defoliators was greater in the lower crown than upper crown. The E. dives canopy supported many more phytophagous insects per unit weight of foliage, as well as more per hectare, than the other two forest types. The greatest number of phytophagous species was also encountered in E. dives canopy. The E. delegatensis canopy supported the lowest number of phytophagous insects per unit weight of foliage as Well as numbers per hectare. Abundance of insect defoliators in the eucalypt forest types in this study was similar to published figures of insect defoliators in northern temperate forests.     

The relationship between possum density and browse damage on kamahi in New Zealand forests

To minimize the impacts of introduced pests and to justify and prioritize pest control, managers need to know the relationship between pest density and damage. This relationship can be difficult to quantify because pest impacts can be highly variable. In New Zealand, introduced brushtail possums (Trichosurus vulpecula) browse a wide range of native forest species. However, possum browse is extremely patchy making it difficult to quantify the relationship between density and damage, meaning the benefits of reducing possum densities are poorly understood. We quantified patterns of possum browse on kamahi (Weinmannia racemosa), a common canopy tree species, at 21 forest sites that were repeat‐measured over an 8‐year period in the North Island, New Zealand, during which time possum densities fluctuated widely. We fitted a multilevel statistical model in order to quantify the relationship between possum density and browse damage while simultaneously quantifying how browse varied among trees, sites and years. Higher possum densities were associated with greater browse damage, but browse was also patchily distributed among trees at the same site, and among sites and years for a given possum density. This heterogeneity meant there was no simple density damage relationship, with the relationship differing from tree to tree and among sites and years. Our results show that at most sites reductions in possum density would have little benefit in reducing the probability of heavy browse on kamahi trees, but at a few sites there would be substantial benefits. This approach provides insights into the pattern and potential causes of variability in possum impacts, and a quantitative basis for prioritizing sites for possum control.     

Seed-bank convergence under different tree species during forest development

Seed banks are of vital importance for local plant persistence and recruitment, for maintaining both plant and genetic diversity and for habitat restoration. Yet, seed-bank dynamics, particularly on the long term and in deciduous forests, remain poorly understood. Additionally, information on compositional seed-bank differences under contrasting tree canopies remains scarce.This study aims at quantifying long-term seed-bank dynamics by sampling vegetation and seed banks along a four-stage successional chronosequence (40, 80, 120 and 250 years) using 12 10 m×10 m plots per forest stand age–class under fully developed oak–hornbeam and beech canopies.Seed banks were remarkably abundant and diverse. Species richness and seed density declined steeply with forest stand age, regardless of canopy species. Seed-bank composition differed significantly with stand age, yet also with tree species. Most likely, tree species-dependent ecosystem engineer effects on light availability and possibly also litter quality affect the seed bank through the vegetation. Compositional differences between seed banks from stands with a different canopy diminished with increasing stand age, possibly due to a gradual loss of species with a less persistent seed bank.Long-term seed-bank dynamics in deciduous forests seem to consist predominantly of a unidirectional and predictable depletion of the seed bank as long as large disturbances, which would allow seed-bank replenishment of early-successional species, are lacking. Furthermore, forest seed-banks appear to converge upon a characteristic seed bank in the later stages of forest development, irrespective of canopy composition, driven by seed-bank depletion and limited input from the herb layer.