Rabbit haemorrhagic disease: applying Occam's razor to competing hypotheses

Rabbit haemorrhagic disease virus (RHDV) is a highly virulent lagovirus endemic in Europe and Australasian populations of the European rabbit, Oryctolagus cuniculus. It has also caused several unexplained disease outbreaks in domestic European rabbits in North America. Non-pathogenic spread of RHDV leading to persistent infection which later reactivated has recently been proposed as the cause of overt disease and death of a pet rabbit in Canada, the first confirmed case of Rabbit haemorrhagic disease in that country. We suggest that there is little evidence to support non-pathogenic spread of virulent RHDV, some evidence that is contradictory, and evidence to support a simpler alternative hypothesis. RHDV can be spread over long distances between sparse rabbit populations by fomites or flying insects. Although highly pathogenic, RHDV can be limited in its spread within rabbit populations, or its presence masked by closely related but non-pathogenic lagoviruses which can provide protection against acute disease. In the absence of any evidence from clinical studies to support reactivation of persistent RHDV infection, the simpler explanation seems more likely to be correct.     

The distribution and dispersal of the introduced millipede, Ommatoiulus moreletii (Diplopoda: Iulidae), in Australia

The millipede, Ommatoiulus moreletii, is indigenous to South-Western Europe. The species has been introduced to Australia where it is in large numbers and constitutes a severe nuisance.
The distribution of O. moreletii in Australia is increasing. Man appears to be a major factor in the establishment of new outbreaks of the species. Individual outbreaks spread slowly but steadily (an expansion of up to 200 metres in radius per year).
The abundance of O. moreletii in Australia relative to its apparent rarity in Southwestern Europe cannot be explained by differences in general climate.
The variety of habitats colonized by O. moreletii in Australia is already wide and there is no reason to doubt that the species will continue to spread much further.     

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.     

Reeves' Muntjac Muntiacus reevesi in Britain: their history, spread, habitat selection, and the role of human intervention in accelerating their dispersal

The origins, early history, captive populations, spread and habitat preferences of Reeves' Muntjac in Britain are reviewed. It is suggested that much of the published information on the history of Muntjac in Britain is based on misconceptions, and that each subsequent report has continued to promulgate a false impression on the origins, time-scale and pattern of spread of the species in Britain. Indian and Reeves' Muntjac were introduced to Woburn Park in Bedfordshire within a year of each other, and it appears that the Indian Muntjac did not thrive, at least in the decade following their introduction. How long they survived as a free-living species in Britain is unclear, but it was probably only a few years. However, there is some evidence to suggest that they might have persisted within the Park at Woburn until 1930. Following the first releases from Woburn in 1901, the numbers of free-living Reeves' Muntjac in Britain remained low until the 1920s, when populations were largely confined to the woods around Woburn, and possibly also around Tring in Hertfordshire. However, in the 1930s and 1940s there were further deliberate introductions in selected areas some distance from Woburn. As a consequence, the subsequent spread of Reeves' Muntjac was from several main foci, i.e. from the vicinity of Woburn, the Norfolk/Suffolk border, three sites in Northamptonshire, two sites in Oxfordshire and two in Warwickshire. The spread in the second half of this century has been aided by further deliberate and accidental releases, and by these means new populations continue to be established outside the main range. Thus the natural spread has been much less impressive than previously assumed; even in areas with established populations it takes a long time for Muntjac to colonize all the available habitat. Data from a number of areas in Britain suggest that the natural rate of spread is about 1 km a year, which is comparable to other species of deer in Britain. The many introductions have complicated an analysis of the habitat preferences of Reeves' Muntjac, and no clear trends could be found. It would appear that Reeves' Muntjac are less dependent on specific types of habitat than previously believed. Examination of the land-class preferences using resource selection indices showed that arable land classes were predominantly selected for, and that marginal upland land classes tended to be avoided. Subsequent logistic regression models based on the land classes selected (and to a lesser extent avoided) by Muntjac were able to predict accurately the current distribution of Reeves' Muntjac in Britain, and one of these, together with our knowledge of their history and spread, was used to infer those areas most likely to be colonized by Muntjac in the near future. The greatest potential for range expansion is in Kent and Sussex, and to a lesser extent north into Lincolnshire, Nottinghamshire and south Yorkshire, and west into Cheshire and north Shropshire. However, long-established populations in areas such as Betws-y-Coed in Wales show that Muntjac may persist in low numbers in atypical habitats. Future habitat changes, such as the planting of new woodlands, and continued deliberate and accidental releases, are likely to lead to population changes in addition to those predicted by the logistic regression model.     

Modelling the initial spread of foot-and-mouth disease through animal movements

Livestock movements in Great Britain (GB) are well recorded and are a unique record of the network of connections among livestock-holding locations. These connections can be critical for disease spread, as in the 2001 epidemic of foot-and-mouth disease (FMD) in the UK. Here, the movement data are used to construct an individual-farm-based model of the initial spread of FMD in GB and determine the susceptibility of the GB livestock industry to future outbreaks under the current legislative requirements. Transmission through movements is modelled, with additional local spread unrelated to the known movements. Simulations show that movements can result in a large nationwide epidemic, but only if cattle are heavily involved, or the epidemic occurs in late summer or early autumn. Inclusion of random local spread can considerably increase epidemic size, but has only a small impact on the spatial extent of the disease. There is a geographical bias in the epidemic size reached, with larger epidemics originating in Scotland and the north of England than elsewhere.     

Resistance to RHD virus in wild Australian rabbits: Comparison of susceptible and resistant individuals using a genomewide approach

Deciphering the genes involved in disease resistance is essential if we are to understand host–pathogen coevolutionary processes. The rabbit haemorrhagic disease virus (RHDV) was imported into Australia in 1995 as a biocontrol agent to manage one of the most successful and devastating invasive species, the European rabbit (Oryctolagus cuniculus). During the first outbreaks of the disease, RHDV caused mortality rates of up to 97%. Recently, however, increased genetic resistance to RHDV has been reported. Here, we have aimed to identify genomic differences between rabbits that survived a natural infection with RHDV and those that died in the field using a genomewide next‐generation sequencing (NGS) approach. We detected 72 SNPs corresponding to 133 genes associated with survival of a RHD infection. Most of the identified genes have known functions in virus infections and replication, immune responses or apoptosis, or have previously been found to be regulated during RHD. Some of the genes identified in experimental studies, however, did not seem to play a role under natural selection regimes, highlighting the importance of field studies to complement the genomic background of wildlife diseases. Our study provides a set of candidate markers as a tool for the future scanning of wild rabbits for their resistance to RHDV. This is important both for wild rabbit populations in southern Europe where RHD is regarded as a serious problem decimating the prey of endangered predator species and for assessing the success of currently planned RHDV variant biocontrol releases in Australia.     

A sharp threshold for disease persistence in host metapopulations

A sharp threshold is established that separates disease persistence from the extinction of small disease outbreaks in an S→E→I→R→S type metapopulation model. The travel rates between patches depend on disease prevalence. The threshold is formulated in terms of a basic replacement ratio (disease reproduction number), ?(0), and, equivalently, in terms of the spectral bound of a transmission and travel matrix. Since frequency-dependent (standard) incidence is assumed, the threshold results do not require knowledge of a disease-free equilibrium. As a trade-off, for ?(0)>1, only uniform weak disease persistence is shown in general, while uniform strong persistence is proved for the special case of constant recruitment of susceptibles into the patch populations. For ?(0)<1, Lyapunov's direct stability method shows that small disease outbreaks do not spread much and eventually die out.     

The impact of sarcoptic mange Sarcoptes scabiei on the British fox Vulpes vulpes population

• 1 Disease epizootics can significantly influence host population dynamics and the structure and functioning of ecological communities. Sarcoptic mange Sarcoptes scabiei has dramatically reduced red fox populations Vulpes vulpes in several countries, including Britain, although impacts on demographic processes are poorly understood. We review the literature on the impact of mange on red fox populations, assess its current distribution in Britain through a questionnaire survey and present new data on resultant demographic changes in foxes in Bristol, UK.
• 2 A mange epizootic in Sweden spread across the entire country in < 10 years resulting in a decline in fox density of up to 95%; density remained lowered for 15–20 years. In Spain, mange has been enzootic for > 75 years and is widely distributed; mange presence was negatively correlated with habitat quality.
• 3 Localized outbreaks have occurred sporadically in Britain during the last 100 years. The most recent large‐scale outbreak arose in the 1990s, although mange has been present in south London and surrounding environs since the 1940s. The questionnaire survey indicated that mange was broadly distributed across Britain, but areas of perceived high prevalence (> 50% affected) were mainly in central and southern England. Habitat type did not significantly affect the presence/absence of mange or perceived prevalence rates. Subjective assessments suggested that populations take 15–20 years to recover.
• 4 Mange appeared in Bristol's foxes in 1994. During the epizootic phase (1994–95), mange spread through the city at a rate of 0.6–0.9 km/month, with a rise in infection in domestic dogs Canis familiaris c. 1–2 months later. Juvenile and adult fox mortality increased and the proportion of females that reproduced declined but litter size was unaffected. Population density declined by > 95%.
• 5 In the enzootic phase (1996–present), mange was the most significant mortality factor. Juvenile mortality was significantly higher than in the pre‐mange period, and the number of juveniles classified as dispersers declined. Mange infection reduced the reproductive potential of males and females: females with advanced mange did not breed; severely infected males failed to undergo spermatogenesis. In 2004, Bristol fox population density was only 15% of that in 1994.

     

The return of the frogs: The importance of habitat refugia in maintaining diversity during a disease outbreak

Recent decades have seen the emergence and spread of numerous infectious diseases, often with severe negative consequences for wildlife populations. Nevertheless, many populations survive the initial outbreaks, and even undergo recoveries. Unfortunately, the long‐term effects of these outbreaks on host population genetics are poorly understood; to increase this understanding, we examined the population genetics of two species of rainforest frogs (Litoria nannotis and Litoria serrata) that have largely recovered from a chytridiomycosis outbreak at two national parks in the Wet Tropics of northern Australia. At the wetter, northern park there was little evidence of decreased genetic diversity in either species, and all of the sampled sites had high minor allele frequencies (mean MAF = 0.230–0.235), high heterozygosity (0.318–0.325), and few monomorphic markers (1.4%–4.0%); however, some recovered L. nannotis populations had low N_e values (59.3–683.8) compared to populations that did not decline during the outbreak (1,537.4–1,756.5). At the drier, southern park, both species exhibited lower diversity (mean MAF = 0.084–0.180; heterozygosity = 0.126–0.257; monomorphic markers = 3.7%–43.5%; N_e = 18.4–676.1). The diversity patterns in this park matched habitat patterns, with both species having higher diversity levels and fewer closely related individuals at sites with higher quality habitat. These patterns were more pronounced for L. nannotis, which has lower dispersal rates than L. serrata. These results suggest that refugia with high quality habitat are important for retaining genetic diversity during disease outbreaks, and that gene flow following disease outbreaks is important for re‐establishing diversity in populations where it was reduced.     

Screening mitochondrial DNA sequence variation as an alternative method for tracking established and outbreak populations of Queensland fruit fly at the species southern range limit

Understanding the relationship between incursions of insect pests and established populations is critical to implementing effective control. Studies of genetic variation can provide powerful tools to examine potential invasion pathways and longevity of individual pest outbreaks. The major fruit fly pest in eastern Australia, Queensland fruit fly Bactrocera tryoni (Froggatt), has been subject to significant long‐term quarantine and population reduction control measures in the major horticulture production areas of southeastern Australia, at the species southern range limit. Previous studies have employed microsatellite markers to estimate gene flow between populations across this region. In this study, we used an independent genetic marker, mitochondrial DNA (mtDNA) sequences, to screen genetic variation in established and adjacent outbreak populations in southeastern Australia. During the study period, favorable environmental conditions resulted in multiple outbreaks, which appeared genetically distinctive and relatively geographically localized, implying minimal dispersal between simultaneous outbreaks. Populations in established regions were found to occur over much larger areas. Screening mtDNA (female) lineages proved to be an effective alternative genetic tool to assist in understanding fruit fly population dynamics and provide another possible molecular method that could now be employed for better understanding of the ecology and evolution of this and other pest species.     

Relationship between Sex Distribution and Sporophyte Production in Pleurozium schreberi (Brid.) Mitt.

The distribution of perigonia, perichaetia, and sporophytesof Pleurozium schreberi has been investigated in Britain andin some other areas. Fruiting populations are widespread ina zone extending from northern Scotland across southern Scandinavia,but are rare in southern Britain, and appear to be infrequentin many other parts of western Europe. Less information is availableoutside western Europe: fruiting collections have been examinedfrom scattered localities throughout much of the species' northerncircumpolar range, but there is no evidence of areas where theyare abundant. The rarity of sporophytes is correlated with a rarity of plantsbearing male inflorescences. Barren specimens normally comprisedonly female and sterile plants, while most fruiting specimenswere bisexual but contained more female than male stems. Femaleplants are abundant throughout Great Britain, but plants withmale inflorescences appear to be widespread only in northernScotland and in East Anglia. The latter area is unusual in thatsporophyte production fails in many bisexual colonies. Sporophyte production was stimulated experimentally by transplantingmale plants from East Anglia into female populations in theWest Midlands. The male plants spread vegetatively within thefemale colonies, and perigonia developed during six successiveyears. At an East Anglian station sporophytes developed aftermaterial was transplanted between unisexual male and femalecolonies only 10 m apart.     

Genetic differentiation of starling (Sturnus vulgaris: Aves) populations in New Zealand and Great Britain

Several hundred starlings (Sturnus vulgaris) were introduced to New Zealand from Great Britain during1860–1880. Allozymic variation at 24 loci was analysed in winter populations sampled at six localities in each country. New Zealand samples had fewer alleles per locus but the same mean heterozygosity (3% per locus) and proportion of polymorphic loci as did British samples. Winter populations in Britain contain European migrants and were genetically homogeneous. Paradoxically, genetic distances among derived New Zealand populations, and between New Zealand and Great Britain were much greater, similar in magnitude to those observed among allopatric populations in other avian species. The geographical pattern of genetic variation in New Zealand suggests that reproductive isolation of populations and random drift have contributed to the development of population differentiation.     

Development of a Novel Rabies Simulation Model for Application in a Non-endemic Environment

Domestic dog rabies is an endemic disease in large parts of the developing world and also epidemic in previously free regions. For example, it continues to spread in eastern Indonesia and currently threatens adjacent rabies-free regions with high densities of free-roaming dogs, including remote northern Australia. Mathematical and simulation disease models are useful tools to provide insights on the most effective control strategies and to inform policy decisions. Existing rabies models typically focus on long-term control programs in endemic countries. However, simulation models describing the dog rabies incursion scenario in regions where rabies is still exotic are lacking. We here describe such a stochastic, spatially explicit rabies simulation model that is based on individual dog information collected in two remote regions in northern Australia. Illustrative simulations produced plausible results with epidemic characteristics expected for rabies outbreaks in disease free regions (mean R₀ 1.7, epidemic peak 97 days post-incursion, vaccination as the most effective response strategy). Systematic sensitivity analysis identified that model outcomes were most sensitive to seven of the 30 model parameters tested. This model is suitable for exploring rabies spread and control before an incursion in populations of largely free-roaming dogs that live close together with their owners. It can be used for ad-hoc contingency or response planning prior to and shortly after incursion of dog rabies in previously free regions. One challenge that remains is model parameterisation, particularly how dogs’ roaming and contacts and biting behaviours change following a rabies incursion in a previously rabies free population.     

Control and spread of Alligator Weed Alternanthera philoxeroides (Mart.) Griseb., in Australia: lessons for other regions

Biological control of alligator weed Alternanthera philoxeroides (Mart.) Griseb. has been successful in limiting growth in water in areas with mild or warm winters, but not on land. Until recently, herbicides have had very limited short term and no long term effectiveness. Several herbicides that now provide better control include: glyphosate over water, and metsulfuron and dichlobenil on land and in shallow water. The latter two are limited by lack of selectivity, contamination of water, and cost. Mechanical or manual control has provided local eradication of the weed at a few locations where infestations were small. Alligator weed is still spreading with new outbreaks on New South Wales, Australia (NSW) coastal beach areas and coastal river systems, and on inland waterbodies. Its use as a cultivated vegetable by some ethnic communities has resulted in many new locations in all eastern Australia states: Queensland to Tasmania. It is predicted that it will spread throughout much of coastal and inland southern Australia. The difficulties with management of this weed indicate that every effort should be made to prevent further invasion of wetlands and, in particular, its introduction to Africa, where it is predicted that all wetlands could support destructive levels of alligator weed growth.     

The Origin and Evolution of Parthenogenesis in Heteronotia Binoei (Gekkonidae): Evidence for Recent and Localized Origins of Widespread Clones

The parthenogenetic form of the gecko lizard species Heteronotia binoei has an unusually broad geographic range and high genetic diversity. Restriction enzyme analysis revealed two basic types of mitochondrial DNA (mtDNA) among the parthenogens. One type is restricted to western populations. The other type, analyzed in detail here, was widespread, being found in populations from central to western Australia. The diversity within this widespread type was low. The variation among parthenogens from central to western Australia was similar to that found within local populations of the sexual species that provided the mtDNA, and was an order of magnitude less than the differentiation shown between sexual populations across the same geographic distance. Phylogenetic analysis revealed that the widespread type of mtDNA in the parthenogens is most closely related to mtDNAs from western populations of the "CA6" sexual parent. These data suggest that these parthenogenetic clones arose recently within a small geographic area, most probably in Western Australia. The parthenogens must have spread rapidly to occupy much of the central and western Australian deserts. This rapid and extensive range expansion provides strong evidence that parthenogenesis can be a successful strategy for lizards in an environment with low and unpredictable rainfall.     

Rabies in urban foxes (Vulpes vulpes) in Britain: the use of a spatial stochastic simulation model to examine the pattern of spread and evaluate the efficacy of different control régimes.

The threat of rabies being reintroduced into Britain is probably greater now than at any time over the last 60 years. This threat is reviewed with particular regard to the problems that would be posed should rabies be introduced to the high-density fox populations found in many cities in southern England. Computer models can provide a valuable means of understanding the pattern of rabies spread in fox populations and the likely problems of control, so the construction of previous rabies models was reviewed. None were found to be suitable for analysing the particular problems posed by high-density, spatially heterogeneous, urban fox populations. Therefore, a new spatial stochastic simulation model was produced, based on demographic and other data collected during a long-term study on the urban fox population in Bristol, and fox density data collected from a number of cities in southern England. The simulation model was used to analyse the effects of spatial heterogeneity in the fox population on the pattern of rabies spread. Simulations were then used to evaluate the effects of: (i) varying levels of fox control; (ii) changing the size of the control zone; (iii) the onset of the rabies epizooty at different times of the year: and (iv) delay before the commencement of control on the chances of containing the disease. These simulations were run for four cities (Bournemouth and Poole, Bristol, Leicester and the West Midlands conurbation) with different mean fox population densities. It was found that the variance in the monthly velocity of the rabies front was greater for heterogeneous fox populations. In cities with high fox densities, low or moderate levels of control were unsuccessful in containing the disease, but these urban areas had the highest rates of success with the highest levels of control. A three-month delay in the commencement of a rabies control campaign on average reduced the chance of successfully controlling the disease by 10-20%, although this was higher in lower-density fox populations. Rabies outbreaks in the dispersal period were on average 10% less likely to be contained. Increasing the size of the control zone increased the chances of successfully containing the disease, although this effect was density dependent, so the effect was less in low-density fox populations. These results are discussed in relation to the current rabies contingency plans for British urban areas.     

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

相似文献   

Foxes, rabbits, alternative prey and rabbit calicivirus disease: consequences of a new biological control agent for an outbreaking species in Australia

1. Rabbit calicivirus disease (RCD; also known as rabbit haemorrhagic disease) has been introduced recently as a biocontrol agent for rabbits in Australia. The consequences for fox populations that use rabbits as primary prey, for populations of alternative native prey, and for pastures, were examined using a model for rabbit- and fox-prone areas of semi-arid southern Australia.
2. Existing data were used to quantify the interactions of foxes, rabbits and pasture. A generic model for predation on native herbivores was constructed by modifying the density-dependent (Type III) functional response of foxes to rabbits to a depensatory (Type II) response that is appropriate for alternative prey. Similar dependence on pasture biomass was assumed for the dynamics of both rabbits and alternative prey in order to identify clearly the consequences of differing predation. In the absence of quantitative data for Australian conditions, the epidemiology of RCD was simulated empirically to mimic a range of potential patterns of occurrence.
3. For semi-arid Australia the model predicts that as the frequency and intensity of RCD epizootics increases: (i) the mean abundance of rabbits will decline, as will the frequency of eruptions of rabbits; (ii) there may be little increase in mean pasture biomass and a small decrease in periods of very low pasture biomass when competition between herbivores is most intense; (iii) the mean abundance of foxes will decline; (iv) there will be a reduced frequency of occasions when rabbit density is low but fox density is high due to a lag in the response of predator populations; and (v) there is potential for an increase in the mean abundance of alternative prey and in the proportion of time their density exceeds a threshold comparable to that currently required for eruptions of rabbits.     

Wave-like spread of Ebola Zaire

In the past decade the Zaire strain of Ebola virus (ZEBOV) has emerged repeatedly into human populations in central Africa and caused massive die-offs of gorillas and chimpanzees. We tested the view that emergence events are independent and caused by ZEBOV variants that have been long resident at each locality. Phylogenetic analyses place the earliest known outbreak at Yambuku, Democratic Republic of Congo, very near to the root of the ZEBOV tree, suggesting that viruses causing all other known outbreaks evolved from a Yambuku-like virus after 1976. The tendency for earlier outbreaks to be directly ancestral to later outbreaks suggests that outbreaks are epidemiologically linked and may have occurred at the front of an advancing wave. While the ladder-like phylogenetic structure could also bear the signature of positive selection, our statistical power is too weak to reach a conclusion in this regard. Distances among outbreaks indicate a spread rate of about 50 km per year that remains consistent across spatial scales. Viral evolution is clocklike, and sequences show a high level of small-scale spatial structure. Genetic similarity decays with distance at roughly the same rate at all spatial scales. Our analyses suggest that ZEBOV has recently spread across the region rather than being long persistent at each outbreak locality. Controlling the impact of Ebola on wild apes and human populations may be more feasible than previously recognized.