Introduced red deer reduce tree regeneration in Pureora Forest, central North Island, New Zealand

This study uses data from repeatedly measured forest monitoring plots (20 × 20 m) (n = 32) and nine ungulate exclosures (paired fenced and unfenced plots; 20 × 20 m) to show the effects of introduced ungulates on tree regeneration in Pureora Forest Park, central North Island, between 974 and 2002. Results show that introduced ungulates, particularly red deer (Cervus elaphus scoticus), have suppressed the regeneration of hardwood species such as Coprosma grandifolia, Elaeocarpus dentatus, Griselinia littoralis, Melicytus ramiflorus, Schefflera digitata and Weinmannia racemosa. These species were only common as saplings and small trees in the complete absence of ungulate browsing. The results of this study suggest that red deer will need to be culled to low densities to assure regeneration of palatable tree species in Pureora Forest.     

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.     

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.     

Diet of the opossum Trichosuvus vulpecula (Kerr) in the Orongorongo Valley,Wellington, New Zealand,in relation to food‐plant availability

The food preferences of a population of opossums in 4.4 ha of lowland broadleaf/podocarp forest were studied for 4 years by faecal analysis. A survey and phenological records were made of the vegetation to compare foods eaten with foods available. Metrosideros robusta and Weinmannia racemosa contributed 60% to the leaf diet, and were being defoliated and killed by opossums. Metrosideros fulgens and Ripogonum scandens, both lianes, together contributed 15 % to the leaf diet but were eaten seasonally. About 10 other species contributed small amounts. Flowers, flower buds, fruit, seeds, bark, and petiole made up about 35 % of the total diet. Compared with their status in the same area 30 years ago, Fuchsia excorticata, Alectryon excelsus, Pseudopanax arboreum, Myrsine salicina, and Coriaria arborea are now rare or localised and of much less consequence in the diet.     

Impact of cattle on conservation land licensed for grazing in South Westland, New Zealand

Making use of existing fences as ready-made exclosures, this study aimed to assess the long-term effects of cattle grazing on forest margins. Results indicated: 1) that cattle browsing and trampling has an impact on vegetation species composition, structure and regeneration; 2) that the effects of a particular grazing regime may take many decades to dissipate; and 3) that the impacts of cattle change with stock intensity. Some plant species appeared to be highly palatable to cattle and only occurred on sites without cattle. Such species included pate (Schefflera digitata), broadleaf (Griselinia littoralis), pigeonwood (Hedycarya arborea), supplejack (Ripogonum scandens), mahoe (Melicytus ramiflorus), milk tree (Streblus heterophyllus), lancewood (Pseudopanax crassifolius) and hen and chickens fern (Asplenium bulbiferum). A small group of plants appeared to regenerate better under cattle than in their absence, particularly mountain horopito (Pseudowintera colorata) and prickly shield fern (Polystichum vestitum). A few species were encouraged by cattle at one site but suppressed by them at another: kahikatea (Dacrycarpus dacrydioides), wheki (Dicksonia squarrosa), Coprosma rhamnoidesand Blechnum fluviatile. The impact of cattle on most other plant species was not discernible. The results of this study, while somewhat equivocal, indicate that future grazing licences in South Westland should restrict stock to low numbers and be confined to already modified sites where damage to conservation values would be minimal.     

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.     

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.     

Diet selection in the green ringtail possum (Pseudochirops archeri): A specialist folivore in a diverse forest

Abstract This study investigated the feeding ecology of the green ringtail possum, Pseudochirops archeri (Pseudocheiridae) in a tropical rainforest with 94 plant species in the canopy. Over 50% of tree use was from only four tree species, Aleurites rockinghamensis, Ficus fraseri, Arytera divaricata and Ficus copiosa. These species were used significantly more frequently than would be expected if tree species were selected randomly in proportion to their relative abundance in the forest. Conversely, 88 other tree species present were used less frequently than expected. Possums also favoured particular individual trees within some of the preferred tree species. In 91% of feeding observations, possums consumed mature leaves only. The availability of young leaves, flowers and fruit varied throughout the year, with a peak in availability of these resources during the early wet season. By primarily selecting mature leaves, green ringtail possums reduce their dependence on seasonally variable resources. We suggest that green ringtail possums should be considered as specialist folivores, focusing on only a few of the tree species available, possibly due to advantages associated with limiting the number of plant secondary metabolites in their diet. Furthermore, they favour certain individual trees within species, perhaps due to intraspecific variation in plant secondary metabolites or nutrient content, behaviour that has been well established in eucalypt folivores. We conclude that green ringtail possums are highly specialized in their feeding ecology, limiting their diet to a small number of continuously available food items.     

Effects of red deer on tree regeneration and growth in Aorangi Forest, Wairarapa

New Zealand forests have been substantially modified by introduced red deer over the past century. New Zealand’s indigenous forest managers need to know if regeneration of palatable tree species can be restored following control or eradication of browsing ungulates. Aorangi Forest, Wairarapa, suffered dramatic changes in forest understorey composition by the 1950s after more than seven decades of colonisation by red deer (Cervus elaphus), feral goats (Capra hircus) and pigs (Sus scrofa). Since then feral goats have been eradicated, but red deer and pigs persist throughout Aorangi Forest despite ongoing recreational hunting. This study uses data from paired fenced and unfenced plots established at seven sites in Aorangi Forest between 1981 and 1987, and re-measured in 2004, to show the effects of ungulates on tree (≥ 2 cm diameter at breast height) regeneration and growth. Our results show that browsing by red deer has prevented regeneration of kanono (Coprosma grandifolia), a highly palatable, fast-growing sub-canopy hardwood tree. Deer reduced the growth of mahoe (Melicytus ramiflorus) trees, probably by directly browsing epicormic shoots. The regeneration of other less palatable sub-canopy trees (e.g. porokaiwhiri, Hedycarya arborea), and slower-growing canopy species (e.g. hinau, Elaeocarpus dentatus and rewarewa, Knightia excelsa) appears to have been unaffected by deer browsing. Accordingly, tree species composition does not appear to have been greatly affected by browsing from deer populations in Aorangi Forest over the past two decades.     

C-methyl-flavonoids from the leaf waxes of some Myrtaceae

The thin waxy coatings on leaves of nine species of Callistemon, two of Melaleuca and one species of Metrosideros, have been studied for the occurrence of leaf surface flavonoids. The Callistemon species and Metrosideros robusta exhibit only C-methylated flavonoids, while O-methyl flavonoids were detected in Melaleuca huegelii. The new natural C-methyl flavonol, 5,7-dihydroxy-3,8,4'-trimethoxy-6-C-methylflavone, was isolated from Metrosideros robusta. The leaf wax of Callistemon coccineus contains the novel C-methylflavonoid, 5,4'-dihydroxy-8-C-methyl-7-methoxy flavanone.     

Long-term impacts of grazing on indigenous forest remnants on North Island hill country, New Zealand

Small isolated patches of native forest surrounded by extensive pastoral grasslands, characteristic of many New Zealand rural landscapes, represent an important reservoir of lowland biodiversity. Improved management of them is a major focus of biodiversity conservation initiatives in New Zealand. We quantified the long-term impacts of grazing on indigenous forest remnants in hill country at Whatawhata, western Waikato, North Island. Structure and composition were compared between forest fragments grazed for >50 years and nearby ungrazed continuous forest. Grazed fragments had shorter and less shady canopies, sparser understoreys, tree populations with larger mean diameters, and ground layers with lower cover of litter and higher cover of vegetation and bare soil than continuous forest. Fragments also had lower indigenous-plant species richness, especially in sapling and seedling populations, and almost no palatable indigenous shrubs, terrestrial orchids, and ferns that require high humidity (e.g. Hymenophyllum spp.), but contained many indigenous and adventive herbaceous species. A transition appears to be occurring in grazed fragments from tall, long-lived trees like Beilschmiedia tawa and Dysoxylum spectabile to short and shorter-lived trees like Kunzea ericoides, Melicytus ramiflorus, and Dicksonia squarrosa. Because grazing inhibits most regeneration processes, unfenced remnants of conifer–broadleaved forest are unlikely to be maintained in grazed pasture in the long term.     

Ecology of Ship Rats (

The impact of browsing by introduced brushtail possums on mixed beech—broadleaved forests in South Westland was estimated from the amount of conspicuous canopy dieback present in 1989- 1990. Aerial and ground-based reconnaissance in all catchments indicated most canopies (84%) were intact. The remaining 16% of canopies were affected by conspicuous dieback, principally of southern rata (Metrosideros umbellata) and/or fuchsia (Fuchsia excorticata). Major dieback nuclei were located in the three areas with the longest history of browsing by possums, which had spread from three known liberation centres. At each dieback nucleus, the amount of dieback reflected the abundance of possum-preferred canopy species. Because South Westland forests contain lower proportions of such species, they are less susceptible to dieback than the conifer-broadleaved forests of central Westland. However, the present low amounts of dieback in South Westland mainly reflect low overall possum densities and a short period of occupation. The occurrence of key possum- preferred species indicates that about one-third of the forests could develop conspicuous canopy dieback if possum numbers increase and 44-94% are susceptible to canopy and/or understorey depletion. By 1992, the few areas selected for sustained possum control effort in Westland under-represented the range of forest composition. However, recently boosted funding for possum control has provided the opportunity to protect representative forest tracts before the onset of widespread ecosystem depletion in South Westland.     

Dynamics of kanuka (

Exclosure plots established in three separate areas of kanuka (Kunzea ericoides var. ericoides) forest on south Kaipara spit in 1983 to assess the impact of introduced fallow deer (Dama dama) were remeasured in 1993. Kanuka shared canopy dominance with mapou (Myrsine australis), houpara (Pseudopanax lessonii) and mahoe (Melicytus ramiflorus ssp. ramiflorus) in relatively old forest in Lookout Bush, Woodhill, and dominated exclusively in two younger stands at South Head; Coprosma rhamnoides dominated understories throughout. At Lookout Bush cohort senescence continued in kanuka and began in mapou and houpara during the period of the study. Seedling thickets of kanuka self-thinned and were also likely to have been smothered by other species. Massive recruitment of mahoe occurred inside the exclosure, and continued in houpara, mostly outside. Mahoe and another generation of houpara are replacing the existing canopy in the absence of deer, and another generation of kanuka and houpara elsewhere in a partially stalled succession; Canopies are still intact at South Head, and there were no major changes in populations of canopy species. However, similar successional pathways are likely to occur there in future. An influx of highly palatable shrubs, e.g., coastal karamu (Coprosoma macrocarpa) and hangehange (Geniostoma rupestre var. ligus Trifolium), into collapsing forest in the absence of deer, and their scarcity or absence elsewhere, indicates continuing impoverishment of the understorey as well as the canopy by deer. In the long term it is likely that a variety of broadleaved trees will invade these stands and that tall semi- coastal forest, similar to extant relies on the dunes, will develop. In the meantime, the high conservation value of these stands suggests that a major reduction in the deer population—sufficient to allow natural successional changes to proceed unhindered—should be a conservation priority for the region.     

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.     

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.     

Liane distribution within native forest remnants in two regions of the South Island, New Zealand

To determine the susceptibility of different forest types to lianes, and to investigate which ecological factors are limiting for lianes, a field survey covering 28 naturally forested sites in Golden Bay (Northwest Nelson) and on Banks Peninsula (Canterbury) was carried out. Results from Detrended Canonical Correspondence Analysis of liane species abundance data in relation to tree and shrub species abundance data and abiotic site variables, showed that the liane community composition was highly correlated with the composition of the tree and shrub community. Forest remnants with introduced lianes in the canopy were characterised by high soil pH, low altitude and high mean temperatures. Native lianes were more widespread geographically and showed a wider ecological tolerance range than introduced lianes. Native liane species were found in both early successional and mature forest, whereas introduced lianes occurred more often with early- successional vegetation. Forest canopy height was also an important factor affecting liane distribution and abundance. Both climbing mechanism and liane stem longevity seemed to affect the height of canopy accessible to lianes. The three twining lianes with longer-lived woody stems (Muehlenbeckia australis, Parsonsia spp. and Ripogonum scandens) reached higher host canopies than a twiner with herbaceous stems (Calystegia tuguriorum), a tendril climber (Passiflora mollisima), and a hook climber (Rubus cissoides). The susceptibility of individual tree and shrub species to canopy invasion by lianes was affected both by the light environment of the potential host, and the architectural properties (height and support availability) of the host.     

Can Individual and Social Patterns of Resource Use Buffer Animal Populations against Resource Decline?

Species in many ecosystems are facing declines of key resources. If we are to understand and predict the effects of resource loss on natural populations, we need to understand whether and how the way animals use resources changes under resource decline. We investigated how the abundance of arboreal marsupials varies in response to a critical resource, hollow-bearing trees. Principally, we asked what mechanisms mediate the relationship between resources and abundance? Do animals use a greater or smaller proportion of the remaining resource, and is there a change in cooperative resource use (den sharing), as the availability of hollow trees declines? Analyses of data from 160 sites surveyed from 1997 to 2007 showed that hollow tree availability was positively associated with abundance of the mountain brushtail possum, the agile antechinus and the greater glider. The abundance of Leadbeater’s possum was primarily influenced by forest age. Notably, the relationship between abundance and hollow tree availability was significantly less than 1∶1 for all species. This was due primarily to a significant increase by all species in the proportional use of hollow-bearing trees where the abundance of this resource was low. The resource-sharing response was weaker and inconsistent among species. Two species, the mountain brushtail possum and the agile antechinus, showed significant but contrasting relationships between the number of animals per occupied tree and hollow tree abundance. The discrepancies between the species can be explained partly by differences in several aspects of the species’ biology, including body size, types of hollows used and social behaviour as it relates to hollow use. Our results show that individual and social aspects of resource use are not always static in response to resource availability and support the need to account for dynamic resource use patterns in predictive models of animal distribution and abundance.     

Vegetation recovery in rural kahikatea (

Vegetation was sampled in kahikatea (Dacrycarpus dacrydioides)-dominant forest fragments having different recovery periods since grazing ceased in the Waikato region, North Island, New Zealand. Changes in vegetation were modeled against recovery periods ranging from 0–74 y and in relation to position within fragment (edge or interior). Indigenous plant species richness increased and adventive plant species (mostly pasture herbs) richness declined with increasing recovery period; small tree and sapling density and seedling ground cover increased. Fragment edges had higher adventive species richness, lower basal area, more established seedlings, lower litter cover and higher grass ground cover, than interior plots. Some indigenous species (e.g. Laurelia novaezealandiae and Myrsine australis) increased with recovery period while some adventive species (e.g. Solanum pseudocapsicum) declined. The relative basal area of Alectryon excelsus increased significantly with recovery period at edges, and that of Melicytus ramiflorus everywhere. Twenty years represents a turning point in the recovery period, with the end of the loss phase of adventive pasture species, the start of the re-establishment phase of indigenous ground layer and understorey species, and significant recovery of population structures of major species. In relatively non-weedy rural environments, retirement from grazing may be sufficient to ensure a return to near-natural states in 40–50 y.     

Body size and tree species composition determine variation in prey consumption in a forest‐inhabiting generalist predator

Trophic interactions may strongly depend on body size and environmental variation, but this prediction has been seldom tested in nature. Many spiders are generalist predators that use webs to intercept flying prey. The size and mesh of orb webs increases with spider size, allowing a more efficient predation on larger prey. We studied to this extent the orb‐weaving spider Araneus diadematus inhabiting forest fragments differing in edge distance, tree diversity, and tree species. These environmental variables are known to correlate with insect composition, richness, and abundance. We anticipated these forest characteristics to be a principle driver of prey consumption. We additionally hypothesized them to impact spider size at maturity and expect shifts toward larger prey size distributions in larger individuals independently from the environmental context. We quantified spider diet by means of metabarcoding of nearly 1,000 A. diadematus from a total of 53 forest plots. This approach allowed a massive screening of consumption dynamics in nature, though at the cost of identifying the exact prey identity, as well as their abundance and putative intraspecific variation. Our study confirmed A. diadematus as a generalist predator, with more than 300 prey ZOTUs detected in total. At the individual level, we found large spiders to consume fewer different species, but adding larger species to their diet. Tree species composition affected both prey species richness and size in the spider''s diet, although tree diversity per se had no influence on the consumed prey. Edges had an indirect effect on the spider diet as spiders closer to the forest edge were larger and therefore consumed larger prey. We conclude that both intraspecific size variation and tree species composition shape the consumed prey of this generalist predator.     

Bird species diversity and abundance before and after eradication of possums and wallabies on Rangitoto Island, Hauraki Gulf, New Zealand

Five-minute bird counts were made on Rangitoto Island in 1998 and 1999, 8 and 9 years after the start, and 1 and 2 years after the completion of a 7-year programme that resulted in eradication of the introduced brushtail possum (Trichosurus vulpecula) and brushtailed rock wallaby (Petrogale penicillata). These were compared with counts made in 1990 (immediately before the start of the programme), to assess whether bird species diversity and abundance had increased as a result of the eradications. The number of bird species detected in 1998/99 was similar to 1990. Five-minute counts of all species except silvereye (Zosterops lateralis) and tui (Prosthemadera novaeseelandiae) were also similar in 1998/99 to those in 1990. Silvereye and tui counts increased significantly. This is most likely a response to increased flowering of pohutukawa (Metrosideros excelsa) and rewarewa (Knightia excelsa) as a result of possum (and wallaby) eradication. The most likely reason for no apparent increase in the abundance of other bird species is the continued presence of predators, especially ship rats (Rattus rattus), cats (Felis catus), and stoats (Mustela erminea).