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Summary Zoonotic disease control presents significant costs and challenges in human and wildlife populations. Although spatial variability and temporal variability in host populations play a significant role influencing the spread and persistence of pathogens, their impact on the effectiveness of disease control are not well understood. Field studies are impractical for many zoonotic diseases; thus, simulation modelling is an alternative. Some research has experimented with metapopulation models of host–pathogen systems, with discrete host populations distributed on a network of connections or on a one-dimensional transect of contiguous cells. Little attention has been paid to treating geographic space as a fine-grained two-dimensional continuum, a more appropriate spatial model for many generalist host and vector species. Using raccoon rabies as an example, we apply an individual-based spatially explicit stochastic simulation model to evaluate effectiveness of vaccination barrier strategies to control rabies. Barrier width and immunization levels are varied over landscapes with habitats of varying quality and spatial heterogeneity, resulting in varying degrees of host connectivity. Our results demonstrate that spatial heterogeneity in the landscape does affect vaccination efficacy. The probability that rabies will breach a vaccination barrier is greater and rabies incidence is higher in landscapes with (i) overall good-quality homogeneous habitat and (ii) overall poor-quality habitat with high spatial heterogeneity, than in landscapes with overall good-quality habitat and high spatial heterogeneity. The influence of landscape conditions on disease dynamics decreases with increasing population immunity. Synthesis and applications. Using a spatially explicit stochastic simulation model, we demonstrated that landscape spatial heterogeneity and vaccination control will interact to influence the success of controlling infectious disease outbreaks. Further, under some landscape conditions, insufficient vaccination is counter-productive because immunized individuals (i) reduce the number of disease transmitting contacts, preventing the disease from growing rapidly thus depleting the susceptible population; and (ii) survive to replenish the stock of susceptible animals through reproduction, facilitating disease persistence.

A landscape genetic simulation modelling approach is used to understand factors affecting raccoon rabies disease spread in southern Ontario, Canada. Using the Ontario Rabies Model, we test the hypothesis that landscape configuration (shape of available habitat) affects dispersal, as indicated by genetic structuring. We simulated range expansions of raccoons from New York into vacant landscapes in Ontario, in two areas that differed by the presence or absence of a landscape constriction. Our results provide theoretical evidence that landscape constriction acts as a vicariant bottleneck. We discuss implications for raccoon rabies spread.

We used a time–space clustering approach to estimate the incubation period of raccoon rabies in the wild using data from the 1999–2001 invasion of raccoon rabies into eastern Ontario from northern New York State. The time differences and geographical distances between all possible pairs of rabies cases were computed, classified and assembled into a time–space matrix. The rows of that matrix represented differences in cases in weeks and the columns represent distances between cases in kilometers and the values in the cells of the matrix represent the counts of cases at specific time and distance intervals. There was a significant cluster of pairs 5 weeks apart with apparent harmonics at additional 5-week intervals. These results are explained by assuming the incubation period of raccoon rabies had a mode of 5 weeks. The time clusters appeared consistently at distance intervals of 5 km. We discuss the possibility that the spatial intervals were influenced by the 5 km radius of the point infection control depopulation process used in 1999 and the 10–15 km radial areas used in 2000. With the practical limits of those radii, there was an intensive effort to eliminate raccoons. Our procedure is easy to implement and provides an estimate of the shape of the distribution of incubation periods for raccoon rabies.

Intra- and interspecific contact rates of 12 adult (five females, seven males) raccoons (Procyon lotor) were recorded while these animals fed at a rural garbage dump 40 km north of Kingston, Ontario, Canada from 15 June to 5 September 1995. While raccoons were being observed, they bit, and were bitten, by their conspecifics an average of 0.99 (+/- 0.21) and 1.28 (+/- 0.21) times per hour, respectively, while feeding. Based on mean nightly contact rates (which included time when raccoons were not observed), raccoons bit one of their conspecifics once every 3 nights while feeding. The mean rate of bites made and received per hour for males was not significantly different from lactating females. There was no detectable difference between the mean rate of bites made and received per hour for raccoons which regularly versus occasionally fed at the dump. No interspecific contacts were observed, though raccoons and striped skunks (Mephitis mephitis) often fed at the dump concurrently. The contact rates in this study are the first to be calculated for raccoons directly from field data and will be useful as a point of reference for modeling rabies spread in raccoons in areas with similar site characteristics.

The province of Ontario (Canada) reported more laboratory confirmed rabid animals than any other state or province in Canada or the USA from 1958-91, with the exception of 1960-62. More than 95% of those cases occurred in the southern 10% of Ontario (approximately 100,000 km2), the region with the highest human population density and greatest agricultural activity. Rabies posed an expensive threat to human health and significant costs to the agricultural economy. The rabies variant originated in arctic foxes: the main vector in southern Ontario was the red fox (Vulpes vulpes), with lesser involvement of the striped skunk (Mephitis mephitis). The Ontario Ministry of Natural Resources began a 5 yr experiment in 1989 to eliminate terrestrial rabies from a approximately 30,000 km2 study area in the eastern end of southern Ontario. Baits containing oral rabies vaccine were dropped annually in the study area at a density of 20 baits/km2 from 1989-95. That continued 2 yr beyond the original 5 yr plan. The experiment was successful in eliminating the arctic fox variant of rabies from the whole area. In the 1980's, an average of 235 rabid foxes per year were reported in the study area. None have been reported since 1993. Cases of fox rabies in other species also disappeared. In 1995, the last bovine and companion animal cases were reported and in 1996 the last rabid skunk occurred. Only bat variants of rabies were present until 1999, when the raccoon variant entered from New York (USA). The success of this experiment led to an expansion of the program to all of southern Ontario in 1994. Persistence of terrestrial rabies, and ease of elimination, appeared to vary geographically, and probably over time. Ecological factors which enhance or reduce the long term survival of rabies in wild foxes are poorly understood.

This chapter reviews the evidence for the impact of natural and anthropogenic barriers on the spread of rabies using evidence mainly drawn from the epidemics of fox and raccoon variant rabies virus over the past 60 years in North America. Those barriers have both directed and inhibited the spread of rabies and, at a regional scale, have been integrated with rabies control efforts in North America. Few studies have been done, however, to examine how the texture (grain) and configuration of the habitat at finer scales affect rabies control, particularly the massive oral vaccination campaigns in operation along the Atlantic coast and southeastern Canada (Ontario, Québec, New Brunswick). To explore these questions, the authors used stochastic simulation. The model of choice was the Ontario Rabies Model (ORM) adapted for use on the high performance computing resources network in Québec (RQCHP-Réseau québécois de calcul de haute performance; http://rqchp.qc.ca). The combination of the ORM and RQCHP allowed us to run many thousands of experiments to explore interactions between nine landscape grain/configuration combinations and vaccination barriers with varying widths and immunity levels. Our results show that breaches of vaccine barriers increase as the grain size of the landscape increases and as the landscape becomes more structured. We caution that mid levels of vaccination can be counterproductive resulting in rabies persistence rather than control. We also note that our model/computing system has the flexibility and capacity to explore a wide range of questions pertinent to improving the efficacy of rabies control.

This chapter reviews the evidence for the impact of natural and anthropogenic barriers on the spread of rabies using evidence mainly drawn from the epidemics of fox and raccoon variant rabies virus over the past 60 years in North America. Those barriers have both directed and inhibited the spread of rabies and, at a regional scale, have been integrated with rabies control efforts in North America. Few studies have been done, however, to examine how the texture (grain) and configuration of the habitat at finer scales affect rabies control, particularly the massive oral vaccination campaigns in operation along the Atlantic coast and southeastern Canada (Ontario, Quebec, New Brunswick). To explore these questions, the authors used stochastic simulation. The model of choice was the Ontario Rabies Model (ORM) adapted for use on the high performance computing resources network in Quebec (RQCHP—Reseau quebecois de calcul de haute performance; http://rqchp.qc.ca ). The combination of the ORM and RQCHP allowed us to run many thousands of experiments to explore interactions between nine landscape grain/configuration combinations and vaccination barriers with varying widths and immunity levels. Our results show that breaches of vaccine barriers increase as the grain size of the landscape increases and as the landscape becomes more structured. We caution that mid levels of vaccination can be counterproductive resulting in rabies persistence rather than control. We also note that our model/computing system has the flexibility and capacity to explore a wide range of questions pertinent to improving the efficacy of rabies control.

Landscape barriers influence movement patterns of animals, which in turn, affect spatio-temporal spread of infectious wildlife disease. We compare genetic data from computer simulations to those acquired from field samples to measure the effect of a landscape barrier on raccoon (Procyon lotor) movement, enabling risk assessment of raccoon rabies disease spread across the Niagara River from New York State into Ontario, an area currently uninfected by rabies. An individual-based spatially explicit model is used to simulate the expansion of a raccoon population to cross the Niagara River, for different permeabilities of the river to raccoon crossings. Since the model records individual raccoon genetics, the genetic population structure of neutral mitochondrial DNA haplotypes are characterised in the expanding population, every 25 years, using a genetic distance measure, phi ST, Mantel tests and a gene diversity measure. The river barrier effect is assessed by comparing genetic measures computed from model outputs to those calculated from 166 raccoons recently sampled from the same landscape. The "best fit" between modelled scenarios and field data indicate the river prevents 50% of attempts to cross the river. Founder effects dominated the colonizing genetic population structure, and, as the river barrier effect increased, its genetic diversity decreased. Using gene flow to calibrate the effect of the river as a barrier to movement provides an estimate of the effect of a river in reducing the likelihood of cross-river infection. Including individual genetic markers in simulation modelling benefits investigations of disease spread and control.

The control of rabies in wild carnivores has evolved from culling operations targeting vector species, to parenteral and oral vaccination of free-ranging vector populations. In order for a rabies control program to be successful, many variables must be evaluated including the timing of control tactic implementation, the ecology of the rabies vector species, and the density of vaccine-baits required to immunize the target population at a level that will result in the control or eradication of rabies from the target zone. Key to the effective control of rabies in wildlife populations, and prevention of re-initiation of outbreaks following control, will be restrictions on the importation of wildlife species that are at risk of importing disease pathogens such as rabies. It has been proven, both in Europe and in North America, that large-scale oral rabies vaccination programs targeting wild carnivore populations can be cost-effective as well as effective at controlling and eliminating rabies. Key to continued success of wildlife rabies control operations will be the design of effective contingency plans to respond to re-emergence of the disease as well as rabies management plans formulated by multi-jurisdictional collaborations at the international level.

We describe a method based on time series analysis that divided the rabies enzootic area of southern Ontario into 13 regions using data collected at the township level, the smallest available geographical unit for Ontario (Canada). The intent was to discover ecogeographic patterns if such existed. For the period 1957-89, the quarterly time series of fox rabies cases for each of the 423 townships in the study area was correlated with the time series of its adjacent neighbors. Townships were then linked to adjacent townships provided the pair-wise correlations had significant correlation coefficients. This procedure produced 13 clusters that remained stable when additional lead/lag relationships between townships were examined. Furthermore, those clusters, which we then termed "rabies units," had different behaviors in terms of species distribution, persistence, and periodicity. Time series in adjacent units were not synchronous. We discuss how our findings influenced the rabies control program in Ontario, how they relate to recent findings about the distribution of fox rabies virus subtypes, and how they lend support for the role of metapopulation structulre in persistence of disease.

Publisher Summary This chapter discusses the concept of controlling rabies in wildlife. It reviews the diagnosis of suspect wildlife, and the oral rabies vaccine development. The development of the rabies vaccines for oral administration to wildlife is complex. It involves the considerations such as—efficacy testing in the target vector species first by intramuscular, and then by oral routes. The release of any new biologically active agent like—a vaccine, an antibiotic, or a pesticide, has its own associated safety concerns. The release of a pesticide molecule, engineered in a laboratory, is under direct human control when released, and its further release can be limited even though it may persist in the biologic web— like DDT, whereas an organism capable of reproduction, such as a live-virus vaccine, has the potential for amplification and divergence in nature. The safety testing in animals, including humans, that may contact the vaccine bait in the field is therefore a prime consideration.

Thirteen adult Raccoons (Procyon lotor) (six females, seven males) that fed at a garbage dump north of Kingston, Ontario were radio-tracked from 21 June to 16 October 1995 to assess their seasonal home ranges and movements. Average Minimum Convex Polygon (MCP) summer and fall home ranges for the collared Raccoons were 78.4 ha (SD=46.2 ha) and 45.6 ha (SD=29.7 ha), respectively. Average grid cell summer and fall home ranges for the collared Raccoons were 143.3 ha (SD=40.0 ha) and 116.9 ha (SD=24.9 ha), respectively. Summer ranges of the Raccoons were significantly larger than fall ranges using both the MCP method (P=0.05) and the grid cell method (P=0.073). Yearling Raccoons travelled an average summer maximum distance from the dump of 2608 m (SD=1964, n=3), more than double the distance of adults (≥2 yr) at 1239 m (SD=547, n=10). The population density for the study area in late August 1995 was estimated at 1 Raccoon/12 ha based on an effective area surrounding the dump of 234 ha. Home range and movement data may be useful to design a strategy to control Raccoon rabies in Ontario.