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.     

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.     

Ecological and evolutionary conditions for fruit abortion to regulate pollinating seed-eaters and increase plant reproduction

Coevolved mutualisms, such as those between senita cacti, yuccas, and their respective obligate pollinators, benefit both species involved in the interaction. However, in these pollination mutualisms the pollinator's larvae impose a cost on plants through consumption of developing seeds and fruit. The effects of pollinators on benefits and costs are expected to vary with the abundance of pollinators, because large population sizes result in more eggs and larval seed-eaters. Here, we develop the hypothesis that fruit abortion, which is common in yucca, senita, and plants in general, could in some cases have the function of limiting pollinator abundance and, thereby, increasing fruit production. Using a general steady-state model of fruit production and pollinator dynamics, we demonstrate that plants involved in pollinating seed-eater mutualisms can increase their fecundity by randomly aborting fruit. We show that the ecological conditions under which fruit abortion can improve plants fecundity are not unusual. They are best met when the plant is long-lived, the population dynamics of the pollinator are much faster than those of the plant, the loss of one fruit via abortion kills a larva that would have the expectation of destroying more than one fruit through its future egg laying as an adult moth, and the effects of fruit abortion on pollinator abundance are spatially localized. We then use the approach of adaptive dynamics to find conditions under which a fruit abortion strategy based on regulating the pollinator population could feasibly evolve in this type of plant-pollinator interaction.     

Diet of stoats at Okarito Kiwi Sanctuary, South Westland, New Zealand

The diet of 871 stoats (Mustela erminea) caught within the Okarito Kiwi Sanctuary, South Westland, New Zealand, between 2001 and 2004 was studied by assessment of gut contents. Stoat and ship rat (Rattus rattus) captures were used as a measure of relative abundance over time, and rat and mouse (Mus musculus) abundance was indexed using tracking tunnels between spring 2002 and winter 2004. There were major increases in rat captures in spring of 2002 and again in spring of 2003. Stoat captures peaked in the following summers, as rat captures declined. Rats and invertebrates were major components of stoat diet, occurring in 41% and 52% of guts respectively; birds were found in 19%. Changes in these percentages with time indicated that stoats shifted their diet from rats to birds and invertebrates as rat abundance (as indexed by tracking tunnels) decreased. The greatest impact on native species is therefore likely to be when rat abundance is declining after major irruptions, as the increased consumption of birds and invertebrates overlaps with the periods of highest stoat abundance.     

Seasonal diet composition in the Crested bobwhite in savannas of central-eastern Venezuela

The diet of Crested Bobwhite (Colinus cristatus, Phasianidae) in a non-flooded savanna located in central-eastern Venezuela was studied. Food items obtained from crops were sorted, counted, and identified mostly to species level. Birds are omnivorous and their diets comprised mostly seeds, some invertebrates, and green vegetation; besides, they consumed snail shells during the breeding season. Seeds dominated in the diet during most of the year but the relative use of seeds, invertebrates, and green vegetation changed seasonally. Some management guidelines in order to increase the abundance of this important game bird in neotropical savannas are proposed.     

The effectiveness of poison bait stations at reducing ship rat abundance during an irruption in a Nothofagus forest

Abstract

Ship rats exhibit large increases in abundance (irruptions) following heavy beech seedfall in New Zealand's Nothofagus forests. Predation by rats at high density severely damages native fauna populations. In 2006 the Department of Conservation undertook a management experiment in the Eg‐linton Valley to see if they could protect endangered species during a rat irruption. Poison (0.15% 1080, followed by 0.0375% coumatetralyl, or Racumin®) was laid in bait stations, and the consequences for rat abundance and survival were estimated. All 10 radio‐tagged rats died, suggesting that 1080 had a high impact on the rat population. The two rats that made the smallest daily movements survived longer than the others. Live trapping documented a reduced abundance of rats within the poison area (450 ha) after 1 month of poisoning. However, after 4 months of poisoning, the abundance of rats had begun to recover. Further investigation is needed on acceptance of Racumin® to rats, optimum spacing of bait stations for rats, and bait competition between rats and mice when densities of both species are high.     

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.     

Sex differences in movement between natural feeding and mating sites and tradeoffs between food consumption,mating success and predator evasion in Mediterranean fruit flies (Diptera: Tephritidae)

Summary Systematic quantitative observations of the location and diel pattern of adult Mediterranean fruit fly, Ceratitis capitata (Wiedemann), activities were carried out in an orange grove and surroundings on the island of Chios in Greece. Natural fly foods were assessed for their contribution to fly longevity, fecundity and fertility. There were diel shifts in male and female location. Females required a substantial and varied diet to realize peak fecundity. This diet was acquired away from the primary host, orange. Foraging for food throughout most of the day on fig and non-host foliage (including feeding on bird droppings) as well as on fig fruit and grapes, females dispersed and fed more than males. A diet of grapes alone did not support any fecundity, contributing only to longevity. A diet of figs alone, on the other hand, sustained both longevity and egg production. Bird feces alone supported neither egg production nor longevity. However, when added to a diet of figs, bird feces significantly increased fly fecundity. Throughout most of the day, males aggregated in leks within the inner canopy of the primary host, orange. The arrival here during the warmest hours of the day of receptive females, followed by pair formation, reinforced the lek mating system on host foliage. In the afternoon, females shifted to orange fruit where they suffered from high predation mortality while ovipositing. Soon after, males also shifted to orange fruit, where they attempted matings with non-receptive ovipositing females. Male feeding on fig fruit occurred late in the day, a time when they were least likely to find a mate. Male survival did not differ between the natural diets. Tradeoffs between food consumption, mating success and predator evasion are discussed for each sex and related to fruit fly mating systems.     

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.     

Fallback Foods of Red Leaf Monkeys (Presbytis rubicunda) in Danum Valley, Borneo

Animals in Southeast Asia must cope with long periods of fruit scarcity of unpredictable duration between irregular mast fruiting events. Long-term data are necessary to examine the effect of mast fruiting on diet, and particularly on the selection of fallback foods during periods of fruit scarcity. No such data is available for colobine monkeys, which may consume substantial amounts of fruits and seeds when available. We studied the diet of red leaf monkeys (Presbytis rubicunda, Colobinae) in Danum Valley, Sabah, northern Borneo, using 25 mo of behavioral observation, phenology and vegetation surveys, and chemical analysis to compare leaves eaten with nonfood leaves. The monkeys spent 46% of their feeding time on young leaves, 38% on seeds, 12% on whole fruits, 2.0% on flowers, 1.0% on bark, and 1.2% on pith. They spent more time feeding on seeds and whole fruit when fruit availability was high and fed on young leaves of Spatholobus macropterus (liana, Leguminosae) as fallback foods. This species was by far the most important food, constituting 27.9% of the total feeding time, and the feeding time on this species negatively correlated with fruit availability. Consumed leaves contained more protein than nonconsumed leaves, and variation in time spent feeding on different leaves was explained by their abundance. These results suggest that red leaf monkeys show essentially the same response to the supra-annual increase in fruit availability as sympatric monogastric primates, increasing their seed and whole-fruit consumption. However, they depended more on young leaves, in particular Spatholobus macropterus, as fallback foods during fruit-scarce periods than did gibbons or orangutans. Their selection of fallback food appeared to be due to both nutrition and abundance.     

Are smaller subspecies of common brushtail possums more omnivorous than larger ones?

Body size is often associated with a dietary divergence within taxonomically related groups so that large animals are often folivorous, while smaller species shift progressively towards omnivory or carnivory. This trend may be influenced by allometric constrains which result in relatively high energetic requirements, but low gut capacities in small animals, compared to their large counterparts. The common brushtail possum (Trichosurus vulpecula, Phalangeridae) has six subspecies ranging widely in weight (1–4 kg). They are not strictly folivorous, but supplement their diet with more nutritious, non‐foliar foods. We predicted that T. vulpecula subspecies diverged in diet in association with body size, with smaller subspecies consuming higher proportions of non‐foliar foods. We assessed this with a review and a meta‐analysis of previous Australian studies. We also investigated the previously unquantified diet of T. v. hypoleucus at three sites in the northern jarrah forest of Western Australia. Results from the meta‐analysis and the review supported our prediction. However, the large variability in the data highlighted their limitations and those of the techniques commonly used to quantify the diet of T. vulpecula. Nonetheless, small subspecies of T. vulpecula appear to consume higher proportions of non‐foliar foods. These results should encourage further research into the body size/diet relationship within T. vulpecula and other possum species. Results from the dietary study of T. v. hypoleucus emphasized their omnivorous diet, which was dominated by foliage and flowers and smaller proportions of invertebrates, seeds and fruits. The common brushtail possum is seldom an exclusive arboreal folivore, but rather ranges from folivory to omnivory.     

Feeding ecology of spider monkeys (Ateles geoffroyi) at Tikal,Guatemala

Field study in tropical dry forest in Tikal National Park, Guatemala, revealed that spider monkeys (Ateles geoffroyi) devoted about 57% of feeding time to fruit (pulp), 20% to the unripe seeds of a single tree species, and the remainder largely to leaves and buds. Fruit, seeds, flowers and leaves ofBrosimum alicastrum (Moraceae) accounted for 56% of the diet, and there was very little predation on invertebrates. A severe drought occurred during 1975, the first year of the study, and a comparison between July of that year with July, 1978 shows that in the drought year there was heavier than normal reliance on fruit of the genusFicus (46% versus 14%) and depressed consumption ofBrosimum (11% vs. 54%). Dietary species diversity (Shannon's measureH) and evenness (Hill'sE) were markedly higher during the drought July than the normal July.     

Small mammals as potential seed dispersers in New Zealand

Weed invasion success is strongly influenced by availability of seed dispersal vectors, which may include animals. We examined the potential of several small introduced mammals (mice, kiore, ship rats and possums) to disperse germinable seeds in New Zealand. Captive animals were fed fleshy fruit of weeds (Berberis glaucocarpa, Cotoneaster spp., Crataegus monogyna, Ilex aquifolium, Leycesteria formosa, Ligustrum sinense, Lonicera japonica, Passiflora mollissima, Pyracantha angustifolia, Sorbus hupehensis) and native species (Coprosma spp., Prumnopitys ferruginea and Solanum aviculare). We recorded the percentage of fruit consumed, seed ingested and gut passage time. Faeces were collected and the seeds extracted and tested for germination potential in an unheated glasshouse (two weed species) or under controlled conditions (11 species). The smallest rodents (mice and kiore) generally destroyed all seeds eaten. Large numbers of viable seeds of the small‐seeded (<1 mg) species, L. formosa and S. aviculare, passed through ship rats. Possums consumed the seeds of all adventive and native fruits except P. ferruginea. The proportion of seeds recovered intact from possum faeces varied with plant species and ranged from 6 to 83%. The time required for 50% of all seeds to be passed by possums ranged from 2.5 to 5.5 days with an average of 3.7 days, and was generally unrelated to simple fruit parameters such as percentage pulp and moisture content. For seeds where germination also occurred in the uneaten controls, the germination of seed from possums ranged from 3 to 78%. Germination was mostly lower in seeds from possums than in the controls, where differences were significant. Possums have major potential to disperse a wide range of fleshy fruit‐producing native and introduced plant species. Ship rats have the potential to disperse those with very small seeds.     

The impact of consumption of fruit by vertebrate and invertebrate frugivores on the germination success of an Australian rainforest seed

Abstract The consumption of fruits by vertebrates and invertebrates can be both advantageous or detrimental to the survival of the seeds they contain. This study investigated the effect of fruit size and consumption of fruit pulp by rodents and beetles on the germination of the seeds of Acmena graveolens, a tropical rainforest canopy tree found in northern Australia. As fruit size increased, germination success and the amount of pulp remaining on the fruits was greater. When beetles were absent, germination success was highest when most of the pulp was removed by rodents, suggesting that they removed an inhibitor of germination. When beetles were present, germination success did not differ significantly across pulp categories, so beetles apparently enhanced germination in seeds with little pulp initially removed, possibly by further removal of fruit pulp. In this study, both rodents and beetles enhanced germination success of A. graveolens seeds by consumption of fruit pulp. Acting as facilitators of germination is a relatively unusual role for both these frugivores that are generally considered to act as seed predators or (in the case of rodents) dispersers.     

Fruit fate,frugivory, and fruit characteristics: a study of the hawthorn,Crataegus monogyna (Rosaceae)

Summary The fate of fruits from a population of European hawthorn (Crataegus monogyna) in western Oregon, USA, was examined over a two-year period. Only one frugivore, the American robin (Turdus migratorius) foraged on the C. monogyna fruits, making this an unusually straightforward fruit-frugivore system. Dispersal efficiency was low, with an average 21% of seeds being dispersed (carried away from parent plants) each year; the most common fate of fruits was to simply fall. Robins dropped 20% of the fruits that they picked, and defecated/regurgitated 40% of the fruits (seeds) that they swallowed, beneath parent plants. One trait, gruit abundance, strongly affected the probability of bush visitation by robins, bushes with larger fruit displays being preferred. Both absolute dispersal success (number of seeds) and dispersal efficiency (proportion of seeds; success per propagule) were also found to be correlated most strongly with initial fruit abundance. Individual plant fecundity and fruit quality were found to vary little between years; as a result, dispersal efficiencies for individual plants were also annully consistent. Larger (older) plants produced more fruits and therefore had higher fitness. These results suggest that the optimal fruiting strategy for C. monogyna is therefore to get as big as possible as quickly as possible by delaying fruiting until later in life.     

The relationship between reproductive success and demographic structure in remnant populations of Primula veris

Plants often suffer reductions in fecundity due to fragmentation, degradation and destruction of populations and their sites. Whether this decrease in seed production has population-level consequences is generally unknown. Here, we aimed to determine the current status of remnant populations in the perennial herb Primula veris in Belgium. Furthermore, we investigated the effects of reduced population size and morph bias on reproductive success and explored if changes in demographic structure could be associated with population fecundity. We studied 69 populations that differed in population size from three to nearly 1500 flowering plants. Three different population types could be distinguished: (a) “dynamic” populations, characterized by high densities of 1 year old juveniles, (b) “normal” populations with adult age-stages prevailing, but still a considerable number of juveniles, and (c) “regressive” populations, in which only flowering adults dominate and rejuvenation hardly occurs. The three population types differed with respect to population size and morph frequency. Dynamic populations were significantly larger and showed a weaker morph bias compared to the intermediate normal and the small regressive populations. Reproductive success, studied the previous year in 26 populations, decreased significantly with decreasing population size and was significantly associated with the demographic structure of the populations. Coefficients of variation for the proportion of flowers setting fruit, the number of seeds per fruit and the total number of seeds per plant decreased significantly with increasing population size. Hence, the observed variability in seed set may be one of the causal factors affecting the observed types of population demographic structure.     

Feeding ecology and dietary differences between blue monkey (Cercopithecus mitis stuhlmanni Matschie) groups in logged and unlogged forest, Budongo Forest Reserve, Uganda

Several studies have investigated the effects of logging on primate population density, and attempted to relate density differences to changes in vegetation composition. As population density in some forest primates may be considerably influenced by dietary quality, it is important to understand how the effects of vegetation changes commonly associated with logging influence primate feeding ecology. Results are presented from a study carried out in the Budongo Forest Reserve in Uganda to investigate differences in blue monkey (Cercopithecus mitis stuhlmanni) feeding ecology in forest blocks with different logging history. Dietary composition differed between logged and unlogged forest, with a higher proportion of unripe fruit consumed in logged forest. By contrast, the proportion of young leaves, invertebrates and seeds in the diet was higher in unlogged forest. Variations in the intake of fruit was also higher in unlogged forest, but feeding patterns on other dietary items were more selective. These differences in diet reflect differences in availability of different categories of food items, as determined by tree species composition, abundance and plant phenological patterns. The observed differences between feeding ecology in logged and unlogged forest are discussed in relation to primate feeding ecology and habitat composition in Budongo.     

Heterogeneity in vertebrate and invertebrate herbivory and its consequences for New Zealand mistletoes

Vertebrate herbivores generally have greater effects than invertebrates on plants. However, few studies have investigated the effects of both invertebrate and vertebrate herbivores on a single plant species. In New Zealand, nationwide declines in mistletoe populations have often been attributed to possum herbivory, but never to insect herbivory. The main goal of the present study was to document levels of vertebrate and invertebrate herbivory on endemic New Zealand mistletoe plants to suggest whether herbivory is leading to mistletoe decline. In the present study, the annual amount of leaf loss from herbivory by the brushtail possum (Trichosurus vulpecula), insect herbivory and leaf abscission were measured in two populations each of three mistletoe species (Alepis flavida, Peraxilla colensoi, and Peraxilla tetrapetala, Loranthaceae). In two populations of each species from February 1997 to February 1998, abscission accounted for the most leaf loss (range 10–84% of total mean leaf area, mean 33%), whereas insects and possums usually removed small and similar amounts (less than 3%). Possum browse caused large amounts of abscission in only one population (A. flavida at Eglinton). Observed possum browse was more heterogeneous than insect browse among branches within a plant (possum coefficient of variation = 2.63, insect CV = 1.98, P < 0.001), among plants in a population (possum CV = 2.15, insect CV = 0.69, P < 0.001), and between populations (possum CV = 1.36, insect CV = 1.09). Moreover, insects damaged 100% of the study plants but never removed more than 16% of leaf area on a single plant, whereas possums only browsed 32% of the study plants but severely defoliated some plants. Thus, while the mean amount of biomass removed across a population may have important consequences for mistletoe survival, the effect of possums on mistletoe populations may also depend on the heterogeneity of browse among individuals in the population.     

Secondary poisoning of stoats after an aerial 1080 poison operation in Pureora Forest, New Zealand

Stoats were monitored by three methods through an aerial 1080 poisoning operation at Waimanoa, Pureora Forest in August 1997. Tracking rates and number of live captures were used as indices of abundance, and radio-transmitters were used to follow individual animals. All 13 stoats with radio-transmitters within the poisoned area died between 2-18 days after the operation. No mustelids were tracked or live-trapped after the operation for three months. Of the radio-tracked stoats that died, rat remains occurred in 67%, passerine birds in 17%, cave weta in 17% and possum in 8%. Residues of 1080 were found in 12 of the 13 dead stoats. Our findings have important implications for the management of threatened species. Stoats are known to be a major factor in the continuing decline of some native birds. Previously, the potential of secondary poisoning to control stoats land other predators) in New Zealand had focused on the use of anticoagulants, as these compounds persist and can accumulate in predators over a longer period. However, our results suggest that secondary poisoning with an acute toxin can also be highly efficient. This may also have greater public acceptability.     

Responses of kukupa (

The kukupa or New Zealand pigeon (Hemiphaga novaeseelandiae) is gradually declining on the New Zealand mainland, due mostly to predation by introduced pest mammals including ship rats (Rattus rattus) and brushtail possums (Trichosurus vulpecula). We report on a co-operative project between Maori landowners, the Department of Conservation, and Manaaki Whenua–Landcare Research researchers to restore a Northland kukupa population and to examine kukupa nesting success in relation to pest abundance. Ship rats and possums were targeted by trapping and poisoning throughout Motatau Forest (350 ha) from 1997 to 1999; only possums were targeted in 2000. All 13 kukupa nests located before pest control started in late 1997 failed at the egg stage, but all seven nests located in 1998–99 successfully fledged young when trapping and tracking indices of possums and ship rats were less than 4%. After pest control, counts of kukupa and some other bird species increased at Motatau compared with counts in a nearby non-treatment block, suggesting numbers of adult kukupa can be increased in small forest areas by intensive pest control. This increase is due at least partly to increased nest success. Evidence from time-lapse video cameras, sign remaining at nests, and nest success rates under different pest control regimes suggest both ship rats and possums are important predators at kukupa nests.