Limits to predator regulation of rabbits in Australia: evidence from predator-removal experiments

Summary Predator-prey studies in semi-arid eastern Australia demonstrated that populations of rabbits (Oryctolagus cuniculus) could be regulated by predators. The functional, numerical and total responses of foxes (Vulpes vulpes) to rabbits and the numerical response of feral cats (Felis catus) to rabbits, are described. Measurement of the rabbit component of foxes' stomach contents indicates a Type III functional response. The size of the fox population in summer was dependent on the availability of rabbits over the immediately preceding rabbit breeding season but there appeared to be no density-dependent aggregation of young foxes in areas of surplus food. The total response of foxes, estimated using the short-term numerical response of dispersing foxes, was directly density-dependent for low rabbit densities and inversely density-dependent for high rabbit densities. Two states are possible with this form of total response: a state with low rabbit densities regulated by predators and a state with high rabbit densities which occurs when rabbits escape predator regulation. The boundary between regulation and non-regulation by predators was demonstrated by a predator-removal experiment. In the treated areas, predators were initially culled and rabbits increased to higher densities than in an untreated area where predators were always present. When predators were allowed back into the treated areas, rabbit populations continued to increase and did not decline to the density in the untreated area. This is the critical evidence for a two-state system. When predators were present, rabbits could be maintained at low densities which were in the density-dependent part of the total response curve for foxes. Exceptionally high rabbit recruitment, or artificially reduced predation, could result in rabbits escaping predator-regulation. Under these circumstances, rabbits could move into the inversely density-dependent region of the total response curve for foxes.     

Pests controlling pests: does predator control lead to greater European rabbit abundance in Australasia?

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Optimising the Application of Multiple-Capture Traps for Invasive Species Management Using Spatial Simulation

Internationally, invasive vertebrate species pose a significant threat to biodiversity, agricultural production and human health. To manage these species a wide range of tools, including traps, are used. In New Zealand, brushtail possums (Trichosurus vulpecula), stoats (Mustela ermine), and ship rats (Rattus rattus) are invasive and there is an ongoing demand for cost-effective non-toxic methods for controlling these pests. Recently, traps with multiple-capture capability have been developed which, because they do not require regular operator-checking, are purported to be more cost-effective than traditional single-capture traps. However, when pest populations are being maintained at low densities (as is typical of orchestrated pest management programmes) it remains uncertain if it is more cost-effective to use fewer multiple-capture traps or more single-capture traps. To address this uncertainty, we used an individual-based spatially explicit modelling approach to determine the likely maximum animal-captures per trap, given stated pest densities and defined times traps are left between checks. In the simulation, single- or multiple-capture traps were spaced according to best practice pest-control guidelines. For possums with maintenance densities set at the lowest level (i.e. 0.5/ha), 98% of all simulated possums were captured with only a single capacity trap set at each site. When possum density was increased to moderate levels of 3/ha, having a capacity of three captures per trap caught 97% of all simulated possums. Results were similar for stoats, although only two potential captures per site were sufficient to capture 99% of simulated stoats. For rats, which were simulated at their typically higher densities, even a six-capture capacity per trap site only resulted in 80% kill. Depending on target species, prevailing density and extent of immigration, the most cost-effective strategy for pest control in New Zealand might be to deploy several single-capture traps rather than investing in fewer, but more expense, multiple-capture traps.     

Can increased niche opportunities and release from enemies explain the success of introduced Yellowhammer populations in New Zealand?

Some introduced species succeed spectacularly, becoming far more abundant in their introduced than in their native range. 'Increased niche opportunities' and 'release from enemy regulation' are two hypotheses that have been advanced to explain the enhanced performance of introduced species in their new environments. Using an introduced bird species, the Yellowhammer Emberiza citrinella , which was first released in New Zealand in 1862, as a model, we tested some predictions based on these hypotheses. By quantifying habitat availability and quality, and measuring nest predation rates, we investigated whether increased niche opportunities or release from nest predation could explain the higher density of the Yellowhammer in New Zealand farmland, compared to farmland in their native Britain. Yellowhammer territory densities were over three times higher in New Zealand (0.40 territories per ha) than in comparable British farmland (0.12 territories per ha), and Yellowhammer densities remained significantly higher in New Zealand, after accounting for differences in habitat availability. The density and diversity of invertebrates, a key food resource for nestling Yellowhammers, was significantly lower in New Zealand than in Britain. Hence, these aspects of niche availability and quality cannot explain the higher density of Yellowhammers in New Zealand. Nest predation rates in New Zealand were similar to those in Britain, suggesting that release from nest predation also could not account for the higher density of Yellowhammers in New Zealand. Differences in winter survival, due to differences in winter food supply or the severity of the winter climate, along with release from other types of 'enemy' regulation are possible alternative explanations.     

Modelling the use of NPV for the biological control of Asian gypsy moth Lymantria dispar invading New Zealand

1 The accidental introduction of the Asian strain of gypsy moth (AGM) Lymantria dispar (L.) to New Zealand poses a major threat to New Zealand's forestry industry. To aid eradication and control decisions in the event of its establishment, a model was developed for the effect of nuclear polyhedrosis virus (NPV) as biological control for AGM in New Zealand. 2 The model simulates within‐ and between‐season gypsy moth population dynamics, including temperature‐dependent development, density dependence through predation and resource limitation, and interactions with NPV. 3 Following its introduction to New Zealand, AGM is predicted to increase more quickly and have more severe outbreaks than the European strain. In the absence of predators, the model predicts initial outbreaks then damped oscillations to an equilibrium. 4 In the model, a single application of NPV (2500 GPIB ha^?1) at the time of maximum larval density gave up to 80% suppression of peak larval densities in the following year. The same level of suppression was achieved in the absence of predators. 5 In the long term, the model predicted that spraying when an outbreak was just beginning gave best results (a 50–70% reduction of the following two outbreaks). Simulation of threshold spraying resulted in NPV application, on average, every 7 years and suppression of outbreak densities by 40–70%. Following a single application, NPV was maintained in the population as a classical biological control agent, giving approximately a 20% reduction in outbreak densities. 6 Eradication of AGM using NPV was possible if larval densities were very low.     

Short-term effects of rabbit grazing on a degraded short- tussock grassland in Central Otago

Rabbits are serious economic and environmental pests in New Zealand's semi arid lands, yet there is surprisingly little quantitative information about their grazing impacts. This paper describes the shortterm gains in pasture yield following protection from rabbit grazing in a rabbit-prone, dry tussock grassland community in Central Otago. During the four most productive plant growing months of 1994 (September to December), a six-fold increase in pasture yield was observed after protection from rabbit grazing (139 kg dry weight ha(-1) with rabbits cf. 853 kg DW ha(-1) without rabbits). Rabbit counts were 30 to 42 rabbits per spotlight km. The following four months (January to April) were characterised by reduced pasture growth (3 kg DW/ha with rabbits cf. 337 kg DW ha(-1) without rabbits) and higher rabbit numbers (42-76 rabbits per spotlight km), and was a critical period of herbage depletion. These substantial differences in pasture yield indicate the potential benefits for pastoral production and land conservation following protection from rabbits.     

Can stoat (

There are currently many attempts in New Zealand to restore native ecosystem functioning through the intensive control of introduced mammalian predators. One system that is faltering is bird pollination of endemic mistletoes (Peraxilla tetrapetala) by bellbirds (Anthornis melanura), apparently because of stoat (Mustela erminea) predation. We used a paired-catchment experiment in Nothofagus solandri var. cliffortioides forest at Craigieburn, central South Island, to measure whether stoat control could restore bellbird densities and mistletoe pollination. Stoat trapping for 10?12 weeks during the 2000/01 and 2001/02 nesting seasons significantly reduced stoat abundance in the treatment area compared with the non-treatment area. As a consequence, bellbird nest survival and densities increased immediately and significantly in the treatment area. Nests in 2000/01 were four times more likely to succeed in the treatment area (66.4%) than in the non-treatment area (16.4%), where video monitoring showed stoats were the key predator. Bellbird numbers per 5-minute count increased 79%. Such a large response following a small-scale stoat control operation suggests that predators limited the Craigieburn bellbird population. Adult bellbirds seem to be less susceptible than eggs and chicks to predation, as bellbird densities were still significantly elevated 24 months after trapping ceased. However, the increase in bellbird densities did not significantly improve mistletoe pollination. Therefore, the stoat trapping was only partially successful in restoring ecosystem functioning.

[Note: This paper was presented by DK as recipient of the 2000 NZ Ecological Society Award at the NZES conference in August 2001.]     

Mass trapping of Prays nephelomima (Lepidoptera: Yponomeutidae) in citrus orchards: optimizing trap design and density

The moth Prays nephelomima (Meirick) (Lepidoptera: Yponomeutidae) is a significant pest of citrus (Citrus spp.), and the recent identification of the female sex pheromone has enabled new direct control tactics to be considered. Six trap designs were compared for suitability in mass trapping, and Pherocon III delta traps were chosen to further evaluate mass trapping. A mass trapping field trial was carried out at five lemon, Citrus limon L., orchards to determine the effect of trap density on catch and rind spot damage on fruit. One plot (0.33-1.0 ha) of each of the five trap density treatments (3, 10, 30, 100, and 300 traps/ha) were operated at each orchard over 12 wk. Catch per trap was reduced as trap density increased and a mean of 12,000 and 16,000 males per ha were killed at the trap densities of 100 and 300 traps per ha, respectively. Increased trap density reduced the percentage of flowers infested with P. nephelomima larvae and reduced the number of moths emerging from flowers. The incidence of rindspot damage on fruit decreased from 45 to 16% as the density of traps increased from 3 to 100 traps per ha. Incidence (percentage of fruit with rindspot) and severity (number of rindspots per fruit) was similar at 100 and 300 traps per ha, indicating that the optimal trap density for reducing rindspot damage is likely to be between 30 and 100 traps per ha. Prospects for converting mass trapping to a lure and kill system are discussed.     

Canopy cover and tree regeneration in old-growth cove forests of the Appalachian Mountains

Relationships between canopy cover and tree regeneration were determined for various species in cove forests of the Great Smoky Mountains. Old-growth stands were sampled with six plots covering a total area of 4.8 ha. Each plot was subdivided into contiguous 10×10 m quadrats. Canopy cover overlying each of the 480 quadrats was characterized with three different indices based on visual estimates of cover. Influences of: (1) overlying cover, (2) proximate openings, and (3) total area of proximate openings on quadrat regeneration densities were determined. Most species reproducing by seed and some species reproducing by vegetative means had higher densities in quadrats with openings, but only the intolerants were highly dependent on gaps. Tsuga canadensis, a very shade-tolerant species, was one of the few species with abundant regeneration beneath dense canopy cover. In general, understory areas near gaps had somewhat higher regeneration densities than other areas with overlying cover. Several shade-tolerant species showed a positive regeneration density response to canopy openings and an ability to regenerate in gaps 0.01–0.03 ha in area. These openings were too small for intolerant species. Many species exhibited a positive response to total size of the proximate opening(s). A sharp increase in regeneration density with area of the opening(s) was evident at approximately 0.04 ha for the shade-intolerant species.     

Density-dependent grazing impacts of introduced European rabbits and sympatric kangaroos on Australian native pastures

Little information is available on relationships between pest animal density and damage in natural ecosystems. Introduced European rabbits, Oryctolagus cuniculus, cause severe damage to Australian native vegetation but density–damage relationships are largely unexplored. There are no recognized simple methods to estimate their impacts on native pastures, due in part to confusion with grazing impact of other herbivores. We tested simple quantitative sampling methods using multiple small quadrats to detect site differences in pasture cover, pasture species richness and dung pellet density of herbivores, from which rabbit density and relative abundance of larger herbivores were estimated. Native pasture cover and species richness declined exponentially with increasing rabbit density, within the range of 0–5 rabbits ha^?1, while cover of unpalatable exotic pasture species increased. By contrast, kangaroo abundance was positively related to palatable native pasture cover and negatively related to cover of unpalatable weeds, and had no negative effect on native pasture cover or species richness that was discernable against a background of low to moderate rabbit densities. Perennial native forbs and perennial grasses replaced invasive Wards weed as the dominant ground cover at low rabbit densities. We conclude that, regardless of previous grazing history, contemporary kangaroo grazing pressure and weed invasion, the severely degraded state of native pastures was perpetuated by rabbits. The effect of rabbits on native pasture can be recorded in a simple manner that is suitable for identifying density–damage relationships in the presence of other herbivores and changes over time. This method is seen as particularly useful in setting target densities below which rabbits must be managed to maintain native plant communities and ecosystem function in southern Australia. It may also be useful to demonstrate rabbits’ impacts in other regions, including optimum densities for plant biodiversity benefits in their native European range.     

Does a HDL injection reduce the development of serum hyperlipidemia and progression of fatty streaks in cholesterol fed rabbits?

The influences of homologous (rabbit) or heterologous (human) high density lipoprotein (HDL) on the development of serum hyperlipidemia and progression of fatty streaks were studied in cholesterol fed rabbits. Three groups of New Zealand rabbits were fed a 0.5% cholesterol rich diet for 8 weeks. Additionally into these animals the following solutions were injected intravenously two times per week: group 1 (control): saline; group 2: human HDL dissolved in saline; group 3: rabbit HDL dissolved in saline. The animals of group 2 had lower serum cholesterol levels during the dietary period than rabbits of group 1 (p < 0.05) but the surface of intima covered with fatty streaks was the same as in group 1. On the other hand, the serum cholesterol level in rabbits of group 3 was the same as in group 1 during the whole experimental period, but the surface of aorta covered with fatty streaks was significantly lower (p < 0.05) in group 3 than in group 1. The results of this study support the hypothesis of an antiatherogenic action of HDL, which seems to be independent of the influence of HDL on the serum lipids but depends on the source of HDL.     

Water and sodium intake of wild and New Zealand Rabbits following angiotensin

Two rabbit strains, New Zealand (laboratory) rabbits and Australian wild rabbits, both members of the Oryctolagus cuniculus genus were studied. New Zealand rabbits under control conditions consumed 2-5 times more water and 8-30 times more 0.5 M NaCl/kg body weight than wild rabbits. Single injections of angiotensin II or III administered ICV did not induce water drinking in either strain. Acute ICV infusion of angiotensin II also did not influence water intake, but after several days of administration, induced increased sodium intake. Intravenous infusion of graded doses of angiotensin II induced diuresis only at the higher doses in both strains. In New Zealand rabbits, this was accompanied by a commensurate and concurrent increase in water intake. Intravenous infusion of angiotensin II also induced urinary sodium loss that was either accompanied or followed by increased sodium intake. The development of salt appetite in both strains was preceded by sodium loss.     

Rabbit (Oryctolagus cuniculus) abundance and protected areas in central-southern Spain: why they do not match?

European rabbits (Oryctolagus cuniculus), a keystone species in the Iberian Mediterranean ecosystem, are the staple prey of the Iberian lynx (Lynx pardinus) and the Spanish imperial eagle (Aquila adalberti). These predators require medium to high rabbit densities and a low degree of human disturbance. We compared rabbit abundances in areas of central-southern Spain under three levels of protection and management: protected areas, intensively managed (nonprotected) hunting estates, and other nonprotected areas. We used pellet abundance indices to estimate rabbit density in 118 surveys conducted during the summers of 2002 and 2003. We observed greater rabbit abundance in intensively managed hunting estates compared to protected areas and other nonprotected areas, perhaps because policy makers did not consider rabbit numbers when selecting priority areas. Alternatively, differences in game management practices (e.g., predator control or habitat management) may explain the higher rabbit densities observed in managed hunting estates. Our results suggest that the best feeding conditions for the Iberian lynx and the Spanish imperial eagle occur in intensively managed hunting areas, where such predators are frequently persecuted. The conservation of these endangered predators may require efforts to increase rabbit densities in protected areas.     

Arterial fatty acid-binding protein activity associated with dietarily-induced and spontaneously occurring atherosclerosis in the rabbit (Oryctolagus cuniculus)

1. Adult WHHL rabbits, or New Zealand rabbits fed either a stock chow diet or a high cholesterol diet were evaluated to assess the relationship between the development of aortic atherosclerosis and arterial FABP activity. 2. Aortic FABP activity was significantly (P less than 0.05) lower in atherosclerotic New Zealand aortas (0.039 +/- 0.008 nmol palmitoyl CoA bound/mg soluble prot) which had developed macroscopic lesions on 80% of the aortic surface as compared to lesion-free New Zealand aortas (0.053 +/- 0.002 nmol palmitoyl CoA bound/mg soluble prot). 3. In spontaneously hyperlipidemic rabbit (WHHL) aortas, FABP activity (0.023 +/- 0.004 nmol palmitoyl CoA bound/mg soluble prot) was significantly lower (P less than 0.05) than in either the normal or atherosclerotic New Zealand aortas. 4. To our knowledge, this study is the first to report a change in arterial FABP with the atherogenic process.     

Emergence behaviour of rabbits, Oryctolagus cuniculus, in Central Otago, New Zealand

The characteristic emergence behaviour of a rabbit population in Central Otago, New Zealand, involves a steady rise in the number of rabbits active throughout the afternoon to a peak near sunset. This differs from populations in Great Britain and Australia, where emergence occurs later and is more rapid. Young rabbits emerged slightly later than adults and were more susceptible to disturbances. Slightly earlier emergence by male rabbits, particularly between May and September, was possibly related to the increased levels of territoriality and social interaction just before and during the early stages of the breeding season. Three emergence indices (25% of the population feeding, 50% of the population emerged, and maximum number of rabbits emerged) were significantly correlated with season. Rabbits emerged earliest before sunset in winter and spring and latest in autumn. Daily variation in emergence times was considerable and was related to weather factors such as temperature and wind direction. A greater level of diurnal activity in New Zealand rabbit populations than in those in Great Britain and Australia is possibly associated with lower levels of diurnal predation in New Zealand.     

Population ecology of Mus musculus on Mana Island, New Zealand

Feral house mice ( Mus musculus ) living on 217 ha Mana Island, New Zealand, with no mammalian predators, were snap-trapped and autopsied. A 7-month breeding season took the population from a spring low to extremely high density in autumn. Litters were largest in the middle of the breeding season, and significantly larger on Mana than on the New Zealand mainland. Litter size in early pregnancy was similar for young and old mice but more embryos were resorbed by old females. The breeding season ended in April when adult females stopped ovulating and young failed to mature. When the population declined over winter no animals bred, they all lost weight, and even previously mature males lost their reproductive ability. Mice continue to grow throughout life and become larger than mice in most populations on the New Zealand mainland. The regular and pronounced seasonal pulse in Mana's mouse population contrasts with longer-term fluctuations generally seen in mainland populations at lower density in indigenous forest. These differences may be explained by absence of predators, habitat features or lack of any chance to disperse on the island.     

Calibration of tunnel tracking rates to estimate relative abundance of ship rats (

Ship rat (Rattus rattus) and mouse (Mus musculus) density and habitat use were estimated by snap trapping and tracking tunnels at Kaharoa in central North Island, New Zealand. Eighty-one ship rats were caught in an effective trapping area of 12.4 ha. Extinction trapping gave an estimated density of 6.7 rats ha(-1) (6.5-7.8 rats ha(-1), 95% confidence intervals). A linear relationship existed between ship rat trapping and tracking rates. Estimating the density of mice was impossible because trapping rates increased rather than decreased during the experiment. Comparisons of density from tracking and trapping rates of mice may be confounded by interference by rats, but this requires further investigation.     

Development and partial metabolic characterization of a dietary cholesterol-resistant colony of rabbits

A colony of New Zealand white rabbits has been developed which, when fed a cholesterol-supplemented diet, exhibit unusual resistance to hypercholesterolemia and atherosclerosis, disorders usually observed in normal cholesterol-fed rabbits. When resistant rabbits (RT) were fed a normal low cholesterol diet (ND), their plasma lipoprotein patterns were significantly different from those of normal rabbits (NR) fed the same diet. The low density lipoprotein cholesterol (LDL-c)/high density lipoprotein cholesterol (HDL-c) ratio and LDL-c/very low density lipoprotein cholesterol (VLDL-c) ratio were lower in the resistant rabbits. The hydrated density of HDL of the normal-responsive rabbits was greater than that of the resistant rabbits. LDL from resistant rabbits contained a lower proportion of esterified cholesterol and protein than LDL from normal rabbits. Peripheral mononuclear cells from resistant rabbits bound about 30% more 125I-labeled rabbit LDL than mononuclear cells from normal rabbits. These results demonstrate that the plasma cholesterol levels of these animals is at least partly under genetic control and that compositional differences exist between the major plasma lipoprotein classes of normal and resistant rabbits even during the ingestion of low-cholesterol diet. The results indicate that at least a part of the difference in the cholesterolemic responses between the two rabbit groups is due to an enhanced LDL uptake by the mononuclear cells, and presumably by other somatic cells of the resistant group.     

Long-term Assessment of the Biological Control of Sitona discoideus by Microctonus aethiopoides and Test of a Model

Autumn densities of the pest weevil Sitona discoideus and its braconid parasitoid Microctonus aethiopoides were monitored from 1996 to 1998 on the Canterbury Plains, New Zealand. M. aethiopoides was introduced as a biological control agent in 1982 and first appeared in the study area in 1986. By 1991 around 50% of autumn weevils were parasitized and weevil density had been reduced by 75%. A model for the system at that time suggested that this level of suppression would be sustained. In agreement with the model, the recent survey confirmed that successful biological control had been maintained, with 75% suppression of weevil density but slightly lower rates of parasitism of around 35%. Weevil densities showed a significant trend longitudinally across the area surveyed, increasing from east to west, probably reflecting soil type. Weevil sex ratio was significantly biased towards females, yet the proportion of males that were parasitized was twice that of females. Percent parasitism in autumn related positively to weevil density over time and space. The scale of homogeneity, and by implication effective annual dispersal, is estimated at 12 +/- 4 km radius for weevils.     

Curative and protective activity in rabbits after reinfection with Schistosoma mansoni: a new model of immunity?

New Zealand rabbits were infected on day 1 and challenged on days 15, 30, and 60 with 1,000 Schistosoma mansoni cercariae/animal/infection. Challenged and control rabbits were perfused 60 days after each infection, corresponding to days 75, 90, and 120 after the first exposure. No decrease in number of adult schistosomes occurred in animals reinfected 15 days after primary infection, but, when the rabbits were challenged 30 and 60 days after the first infection, worm burden reduction of 61.4% and 92.6%, respectively, was observed as compared to infection controls. These data indicate that rabbits submitted to reinfection are able to kill the worms from their primary infection, besides being protected against challenge parasites.