Using social networks to deduce whether residents or dispersers spread parasites in a lizard population

1. Heterogeneity of host behaviour can play an important role in the spread of parasites and pathogens around wildlife populations. Social networks have previously been suggested to represent transmission pathways within a population, but where the dynamics of host-parasite interactions are difficult to observe, networks may also be used to provide insights into transmission processes. 2. Pygmy bluetongue lizards, Tiliqua adelaidensis, occupy individual territories, live exclusively in burrows constructed by spiders in Australian native grasslands and are hosts to a tick, Bothriocroton hydrosauri, and a nematode, Pharyngodon wandillahensis. 3. On five monthly occasions, the locations of all individual lizards in three study plots were used to construct weighted, undirected networks based on proximity of adjacent burrows. 4. The networks were used to explore alternative hypotheses about the spread of each parasite through the population: that stable population members that remained in the same burrow over the study period played a major role in influencing the pattern of infection or that dispersing individuals played a more significant role. 5. For ticks, host individuals that were infected were more connected in the network than uninfected hosts and this relationship remained significant for connections to residents in the population, but not for connections to dispersers. 6. For nematodes, infected and uninfected hosts did not differ in their overall strength of connection in the network, but infected hosts were more connected to dispersers than were uninfected hosts, suggesting that lizards moving across the population are the major agents for the transmission of nematodes. 7. This study shows how network analyses can provide new insights into alternative pathways of parasite spread in wildlife populations, where it is difficult to make direct observations of transmission-related behaviours.     

Brown spider monkeys (Ateles hybridus): a model for differentiating the role of social networks and physical contact on parasite transmission dynamics

Elevated risk of disease transmission is considered a major cost of sociality, although empirical evidence supporting this idea remains scant. Variation in spatial cohesion and the occurrence of social interactions may have profound implications for patterns of interindividual parasite transmission. We used a social network approach to shed light on the importance of different aspects of group-living (i.e. within-group associations versus physical contact) on patterns of parasitism in a neotropical primate, the brown spider monkey (Ateles hybridus), which exhibits a high degree of fission–fusion subgrouping. We used daily subgroup composition records to create a ‘proximity’ network, and built a separate ‘contact’ network using social interactions involving physical contact. In the proximity network, connectivity between individuals was homogeneous, whereas the contact network highlighted high between-individual variation in the extent to which animals had physical contact with others, which correlated with an individual''s age and sex. The gastrointestinal parasite species richness of highly connected individuals was greater than that of less connected individuals in the contact network, but not in the proximity network. Our findings suggest that among brown spider monkeys, physical contact impacts the spread of several common parasites and supports the idea that pathogen transmission is one cost associated with social contact.     

A comparison of methods for estimating raccoon abundance: Implications for disease vaccination programs

Accurate estimates of demographic parameters are critical to the management of wildlife populations, including management programs focused on controlling the spread of zoonotic diseases. Rabies managers in the United States Department of Agriculture (USDA) have applied a simple raccoon (Procyon lotor) abundance index (RAI) based on cumulative catch of unique raccoons per unit area to determine vaccine-bait distribution densities. This approach was designed to allow for both the collection of biological samples and to index raccoon abundance to determine bait densities for oral rabies programs. However, post-baiting surveillance data indicate that, on average, only 30% of raccoons sampled have vaccine induced rabies antibody titers, suggesting that bait densities may not be well calibrated to raccoon densities. We trapped raccoons using both capture-mark-recapture (CMR) and the standard RAI to evaluate the accuracy of the current index-based methodology for estimating raccoon density. We then developed a resource selection function from spatial data collected from radio-collared raccoons to standardize trap placement within the existing RAI protocol, and evaluated the performance of this modified RAI approach relative to CMR for estimating raccoon population size. Both abundance and density estimates derived using the RAI consistently underestimated raccoon population sizes compared with CMR methods. Similarly, although the use of resource selection models to inform trap placement appeared to improve the accuracy of the RAI, the effectiveness of this method was inconsistent because of an inability to account for variance in detection probabilities. Despite the logistical advantages of using indices to estimate population parameters to determine vaccine bait distribution densities, our results suggest that adjustments may be necessary to more accurately quantify raccoon abundance, which should improve the effectiveness of rabies management in the United States. In particular, estimates of detection probabilities are needed to more precisely quantify abundance estimates and ensure appropriate vaccine coverage rates. © 2012 The Wildlife Society.     

The number of links to and from the starting node as a predictor of epidemic size in small-size directed networks

Much recent modelling is focusing on epidemics in large-scale complex networks. Whether or not findings of these investigations also apply to networks of small size is still an open question. This is an important gap for many biological applications, including the spread of the oomycete pathogen Phytophthora ramorum in networks of plant nurseries. We use numerical simulations of disease spread and establishment in directed networks of 100 individual nodes at four levels of connectivity. Factors governing epidemic spread are network structure (local, small-world, random, scale-free) and the probabilities of infection persistence in a node and of infection transmission between connected nodes. Epidemic final size at equilibrium varies widely depending on the starting node of infection, although the latter does not affect the threshold condition for spread. The number of links from (out-degree) but not the number of links to (in-degree) the starting node of the epidemic explains a substantial amount of variation in final epidemic size at equilibrium irrespective of the structure of the network. The proportion of variance in epidemic size explained by the out-degree of the starting node increases with the level of connectivity. Targeting highly connected nodes is thus likely to make disease control more effective also in case of small-size populations, a result of relevance not just for the horticultural trade, but for epidemiology in general.     

Age,but not Sex or Genetic Relatedness,Shapes Raccoon Dominance Patterns

Raccoons are generally regarded as solitary, yet several studies have found that raccoons frequently form social affiliations. One benefit to sociality in many mammal species is that relatives and close associates can form coalitions against third parties during agonistic encounters. We tested whether raccoon dominance patterns were influenced by age, sex, genetic relatedness, and association patterns at two anthropogenic feeding stations in an urban forest. We found that genetic relatedness had no significant effect on patterns of agonism at one of the feeding stations. At the second feeding station, raccoons were more likely to act aggressively toward close relatives, which is opposite of the predicted pattern. However, when we controlled for the number of times raccoons arrived at feeding stations in close proximity, the effect of relatedness on dominance patterns was not significant at either feeding station. These results suggest that relatedness plays little or no role in shaping dominance patterns of raccoons. Older raccoons were ranked significantly higher in the dominance hierarchies regardless of sex. This pattern leads us to conclude that age is the primary factor driving the outcome of aggressive interactions in raccoons at our study site. Despite frequent social interactions at the study site, the patterns of raccoon dominance more closely resemble patterns found in solitary animals. To confirm the generality of these results and to better understand the evolution of raccoon social behavior, similar studies need to be undertaken in other raccoon populations.     

Network frailty and the geometry of herd immunity

The spread of infectious disease through communities depends fundamentally on the underlying patterns of contacts between individuals. Generally, the more contacts one individual has, the more vulnerable they are to infection during an epidemic. Thus, outbreaks disproportionately impact the most highly connected demographics. Epidemics can then lead, through immunization or removal of individuals, to sparser networks that are more resistant to future transmission of a given disease. Using several classes of contact networks-Poisson, scale-free and small-world-we characterize the structural evolution of a network due to an epidemic in terms of frailty (the degree to which highly connected individuals are more vulnerable to infection) and interference (the extent to which the epidemic cuts off connectivity among the susceptible population that remains following an epidemic). The evolution of the susceptible network over the course of an epidemic differs among the classes of networks; frailty, relative to interference, accounts for an increasing component of network evolution on networks with greater variance in contacts. The result is that immunization due to prior epidemics can provide greater community protection than random vaccination on networks with heterogeneous contact patterns, while the reverse is true for highly structured populations.     

Contact networks and transmission of an intestinal pathogen in bumble bee (Bombus impatiens) colonies

In socially living animals, individuals interact through complex networks of contact that may influence the spread of disease. Whereas traditional epidemiological models typically assume no social structure, network theory suggests that an individual’s location in the network determines its risk of infection. Empirical, especially experimental, studies of disease spread on networks are lacking, however, largely due to a shortage of amenable study systems. We used automated video-tracking to quantify networks of physical contact among individuals within colonies of the social bumble bee Bombus impatiens. We explored the effects of network structure on pathogen transmission in naturally and artificially infected hives. We show for the first time that contact network structure determines the spread of a contagious pathogen (Crithidia bombi) in social insect colonies. Differences in rates of infection among colonies resulted largely from differences in network density among hives. Within colonies, a bee’s rate of contact with infected nestmates emerged as the only significant predictor of infection risk. The activity of bees, in terms of their movement rates and division of labour (e.g., brood care, nest care, foraging), did not influence risk of infection. Our results suggest that contact networks may have an important influence on the transmission of pathogens in social insects and, possibly, other social animals.     

Characterization of Major Histocompatibility Complex (MHC) DRB Exon 2 and DRA Exon 3 Fragments in a Primary Terrestrial Rabies Vector (Procyon lotor)

The major histocompatibility complex (MHC) presents a unique system to explore links between genetic diversity and pathogens, as diversity within MHC is maintained in part by pathogen driven selection. While the majority of wildlife MHC studies have investigated species that are of conservation concern, here we characterize MHC variation in a common and broadly distributed species, the North American raccoon (Procyon lotor). Raccoons host an array of broadly distributed wildlife diseases (e.g., canine distemper, parvovirus and raccoon rabies virus) and present important human health risks as they persist in high densities and in close proximity to humans and livestock. To further explore how genetic variation influences the spread and maintenance of disease in raccoons we characterized a fragment of MHC class II DRA exon 3 (250bp) and DRB exon 2 (228 bp). MHC DRA was found to be functionally monomorphic in the 32 individuals screened; whereas DRB exon 2 revealed 66 unique alleles among the 246 individuals screened. Between two and four alleles were observed in each individual suggesting we were amplifying a duplicated DRB locus. Nucleotide differences between DRB alleles ranged from 1 to 36 bp (0.4–15.8% divergence) and translated into 1 to 21 (1.3–27.6% divergence) amino acid differences. We detected a significant excess of nonsynonymous substitutions at the peptide binding region (P = 0.005), indicating that DRB exon 2 in raccoons has been influenced by positive selection. These data will form the basis of continued analyses into the spatial and temporal relationship of the raccoon rabies virus and the immunogenetic response in its primary host.     

Raccoon (Procyon lotor) in Iberia: Status update and suitable habitats for an invasive carnivore

Raccoons are American carnivores, considered invasive across several countries worldwide, especially in Europe. In the Iberian Peninsula, previous studies on raccoons documented several breeding populations in Spain a decade ago and only two confirmed records from isolated individuals in Portugal. Given the need for updating its Iberian distribution and identifying suitable areas with higher invasion risk, we compiled presence records from established breeding populations and isolated individuals. By using a Maxent approach based on breeding records, we forecasted the suitable habitats in Iberia with higher invasion risk for raccoons and identified the related environmental drivers. Overall, we collected 1039 records of raccoon presence throughout the Iberian Peninsula, including 980 records from established breeding populations. Their origin is probably linked to escapes from captivity. Climatic conditions, linked to both drier and wetter environments, and proximity to water bodies were the main predictors of suitable areas for raccoon’s expansion from the currently established breeding nuclei in Iberia. The forecasted high probability areas showed a wide, but fragmented distribution concentrated on four main areas: central, central-north, central-east, and north-west Iberia. NW Portugal seems to be the area with higher invasion risk in the country, although field surveys showed no evidence of raccoon presence yet. However, there are several records in Spain near the Portuguese border, comprising isolated individuals and breeding populations. Therefore, it is crucial to ensure regular monitoring of areas with high invasion risk, particularly those near facilities with captive raccoons that often act as a source of feral individuals, to assure early detection and effective control for the expansion of this invasive carnivore.     

Infectious disease transmission and contact networks in wildlife and livestock

The use of social and contact networks to answer basic and applied questions about infectious disease transmission in wildlife and livestock is receiving increased attention. Through social network analysis, we understand that wild animal and livestock populations, including farmed fish and poultry, often have a heterogeneous contact structure owing to social structure or trade networks. Network modelling is a flexible tool used to capture the heterogeneous contacts of a population in order to test hypotheses about the mechanisms of disease transmission, simulate and predict disease spread, and test disease control strategies. This review highlights how to use animal contact data, including social networks, for network modelling, and emphasizes that researchers should have a pathogen of interest in mind before collecting or using contact data. This paper describes the rising popularity of network approaches for understanding transmission dynamics in wild animal and livestock populations; discusses the common mismatch between contact networks as measured in animal behaviour and relevant parasites to match those networks; and highlights knowledge gaps in how to collect and analyse contact data. Opportunities for the future include increased attention to experiments, pathogen genetic markers and novel computational tools.     

The dynamics of sexual contact networks: effects on disease spread and control

Sexually transmitted pathogens persist in populations despite the availability of biomedical interventions and knowledge of behavioural changes that would reduce individual-level risk. While behavioural risk factors are shared between many sexually transmitted infections, the prevalence of these diseases across different risk groups varies. Understanding this heterogeneity and identifying better control strategies depends on an improved understanding of the complex social contact networks over which pathogens spread. To date, most efforts to study the impact of sexual network structure on disease dynamics have focused on static networks. However, the interaction between the dynamics of partnership formation and dissolution and the dynamics of transmission plays a role, both in restricting the effective network accessible to the pathogen, and in modulating the transmission dynamics. We present a simple method to simulate dynamical networks of sexual partnerships. We inform the model using survey data on sexual attitudes and lifestyles, and investigate how the duration of infectiousness changes the effective contact network over which disease may spread. We then simulate several control strategies: screening, vaccination and behavioural interventions. Previous theory and research has advanced the importance of core groups for spread and control of STD. Our work is consistent with the importance of core groups, but extends this idea to consider how the duration of infectiousness associated with a particular pathogen interacts with host behaviours to define these high risk subpopulations. Characteristics of the parts of the network accessible to the pathogen, which represent the network structure of sexual contacts from the “point of view” of the pathogen, are substantially different from those of the network as a whole. The pathogen itself plays an important role in determining this effective network structure; specifically, we find that if the pathogen’s duration of infectiousness is short, infection is more concentrated in high-activity, high-concurrency individuals even when all other factors are held constant. Widespread screening programmes would be enhanced by follow-up interventions targeting higher-risk individuals, because screening shortens the expected duration of infectiousness and causes a greater relative decrease in prevalence among lower-activity than in higher-activity individuals. Even for pathogens with longer durations of infectiousness, our findings suggest that targeting vaccination and behavioural interventions towards high-activity individuals provides comparable benefits to population-wide interventions.     

Allocation to sexual versus nonsexual disease transmission

Many diseases have both sexual and nonsexual transmission routes, and closely related diseases often differ in their degree of sexual transmission. We investigate the evolution of transmission mode as a function of host social and mating structure using a model in which disease transmission is explicitly dependent on the numbers of sexual and nonsexual contacts (which are themselves a function of population density) and per-contact infection probabilities. Most generally, and in the absence of trade-offs between the degree of sexual transmission and effects on host fecundity and mortality, nonsexual transmission is favored above the social-sexual crossover point (the host density at which the number of nonsexual contacts exceeds the number of sexual contacts), while sexual transmission is favored below this point. When changes in allocation to the two transmission modes are accompanied by changes in mortality or fecundity, both mixed and pure transmission strategies can be favored. If invading genotypes differ substantially from resident genotypes, genetic polymorphism in transmission mode is possible. The evolutionary outcomes are predictable from a knowledge of the equilibrium population sizes in relation to the social-sexual crossover point. Our results also show that predictions about dynamic outcomes, based on rates of invasion for single pathogens into healthy populations, do not adequately describe the resulting disease prevalence nor predict the subsequent evolutionary dynamics; once invasion of a pathogen has occurred, the conditions for spread of a second pathogen are themselves altered. If the host is considered as a single resource, our results show that two pathogens may coexist on a single resource if they use that resource differentially and have differential feedbacks on resource abundance; such resource feedback effects may be present in other biological systems.     

MHC class II DRB diversity in raccoons (<Emphasis Type="Italic">Procyon lotor</Emphasis>) reveals associations with raccoon rabies virus (Lyssavirus)

In North America, the raccoon rabies virus (RRV) is an endemic wildlife disease which causes acute encephalopathies and is a strong selective force on raccoons (Procyon lotor), with estimates of ∼85% of the population succumbing to the disease when epizootic. RRV is regarded as a lethal disease if untreated; therefore, no evolutionary response would be expected of raccoon populations. However, variable immune responses to RRV have been observed in raccoons indicating a potential for evolutionary adaptation. Studies of variation within the immunologically important major histocompatibility complex (MHC) have revealed relationships between MHC alleles and diseases in humans and other wildlife species. This enhances our understanding of how hosts and pathogens adapt and co-evolve. In this study, we used RRV as a model system to study host–pathogen interaction in raccoons from a challenge study and from four wild populations that differ in exposure times and viral lineages. We investigated the potential role of Prlo-DRB polymorphism in relation to susceptibility/resistance to RRV in 113 RRV positive and 143 RRV negative raccoons. Six alleles were found to be associated with RRV negative status and five alleles with RRV positive animals. We found variable patterns of MHC associations given the relative number of selective RRV sweeps in the studied regions and correlations between MHC diversity and RRV lineages. The allelic associations established provide insight into how the genetic variation of raccoons may affect the disease outcome and this can be used to examine similar associations between other rabies variants and their hosts.     

Contact networks in a wild Tasmanian devil (Sarcophilus harrisii) population: using social network analysis to reveal seasonal variability in social behaviour and its implications for transmission of devil facial tumour disease

The structure of the contact network between individuals has a profound effect on the transmission of infectious disease. Using a novel technology – proximity sensing radio collars – we described the contact network in a population of Tasmanian devils. This largest surviving marsupial carnivore is threatened by a novel infectious cancer. All devils were connected in a single giant component, which would permit disease to spread throughout the network from any single infected individual. Unlike the contact networks for many human diseases, the degree distribution was not highly aggregated. Nevertheless, the empirically derived networks differed from random networks. Contact networks differed between the mating and non-mating seasons, with more extended male–female associations in the mating season and a greater frequency of female–female associations outside the mating season. Our results suggest that there is limited potential to control the disease by targeting highly connected age or sex classes.     

Land-use effects on prevalence of raccoon roundworm (Baylisascaris procyonis)

The raccoon (Procyon lotor) is the definitive host of Baylisascaris procyonis, a large intestinal roundworm that is zoonotic and can result in fatal or severe central nervous system disease in young children. Prevalence of infection among raccoon populations often is high, and in the midwestern United States, B. procyonis has been reported in 68-82% of raccoons. Raccoon populations have increased in response to changes in human land use, and often reach higher densities in urban and suburban landscapes than rural landscapes. However, shifts in foraging behavior among urban raccoons could impact the transmission of B. procyonis if small vertebrate intermediate hosts are not a significant part of the raccoon diet. The objective of this study was to compare prevalence of B. procyonis infection between urban and rural raccoon populations on a regional scale. Necropsy was done on 204 raccoons collected from September through February during 2000-2005 from seven states across the Midwest (regional sample). Baylisascaris procyonis was found in 54% of examined raccoons. Prevalence differed between land-use types (chi2=11.56, df=1, P=0.0007), and was higher among animals collected from rural locations (65%) than those collected in urban locations (41%). Intensity of infection also differed (F=5.52, df=1, P=0.02), with rural raccoons having greater worm burdens (x=29.63+/-36.42) than urban raccoons (x=13.85+/-18.47). Despite high densities of raccoons in urban landscapes, fewer urban raccoons were infected with B. procyonis, suggesting decreased dependence on intermediate hosts as a food source. This possible explanation was supported by a similar trend in prevalence among subsamples of raccoons collected from three Chicago-area populations (local samples) with differing levels of urbanization, population densities, and foraging behavior that had been intensively monitored during 1995-2002. Decreased transmission of B. procyonis in urban landscapes may be due to decreased predation of intermediate hosts, and contact of juvenile raccoons with B. procyonis eggs may be an important factor in maintaining infections within such populations.     

Covariation between the physiological and behavioral components of pathogen transmission: host heterogeneity determines epidemic outcomes

Although heterogeneity in contact rate, physiology, and behavioral response to infection have all been empirically demonstrated in host–pathogen systems, little is known about how interactions between individual variation in behavior and physiology scale‐up to affect pathogen transmission at a population level. The objective of this study is to evaluate how covariation between the behavioral and physiological components of transmission might affect epidemic outcomes in host populations. We tested the consequences of contact rate covarying with susceptibility, infectiousness, and infection status using an individual‐based, dynamic network model where individuals initiate and terminate contacts with conspecifics based on their behavioral predispositions and their infection status. Our results suggest that both heterogeneity in physiology and subsequent covariation of physiology with contact rate could powerfully influence epidemic dynamics. Overall, we found that 1) individual variability in susceptibility and infectiousness can reduce the expected maximum prevalence and increase epidemic variability; 2) when contact rate and susceptibility or infectiousness negatively covary, it takes substantially longer for epidemics to spread throughout the population, and rates of epidemic spread remained suppressed even for highly transmissible pathogens; and 3) reductions in contact rate resulting from infection‐induced behavioral changes can prevent the pathogen from reaching most of the population. These effects were strongest for theoretical pathogens with lower transmissibility and for populations where the observed variation in contact rate was higher, suggesting that such heterogeneity may be most important for less infectious, more chronic diseases in wildlife. Understanding when and how variability in pathogen transmission should be modelled is a crucial next step for disease ecology.     

Network metrics reveal differences in social organization between two fission–fusion species, Grevy’s zebra and onager

For species in which group membership frequently changes, it has been a challenge to characterize variation in individual interactions and social structure. Quantifying this variation is necessary to test hypotheses about ecological determinants of social patterns and to make predictions about how group dynamics affect the development of cooperative relationships and transmission processes. Network models have recently become popular for analyzing individual contacts within a population context. We use network metrics to compare populations of Grevy’s zebra (Equus grevyi) and onagers (Equus hemionus khur). These closely related equids, previously described as having the same social system, inhabit environments differing in the distribution of food, water, and predators. Grevy’s zebra and onagers are one example of many sets of coarsely similar fission–fusion species and populations, observed elsewhere in other ungulates, primates, and cetaceans. Our analysis of the population association networks reveals contrasts consistent with their distinctive environments. Grevy’s zebra individuals are more selective in their association choices. Grevy’s zebra form stable cliques, while onager associations are more fluid. We find evidence that females associate assortatively by reproductive state in Grevy’s zebra but not in onagers. The current approach demonstrates the utility of network metrics for identifying fine-grained variation among individuals and populations in association patterns. From our analysis, we can make testable predictions about behavioral mechanisms underlying social structure and its effects on transmission processes.     

Experimental evidence for the effect of habitat loss on the dynamics of migratory networks

Migratory animals present a unique challenge for understanding the consequences of habitat loss on population dynamics because individuals are typically distributed over a series of interconnected breeding and non‐breeding sites (termed migratory network). Using replicated breeding and non‐breeding populations of Drosophila melanogaster and a mathematical model, we investigated three hypotheses to explain how habitat loss influenced the dynamics of populations in networks with different degrees of connectivity between breeding and non‐breeding seasons. We found that habitat loss increased the degree of connectivity in the network and influenced population size at sites that were not directly connected to the site where habitat loss occurred. However, connected networks only buffered global population declines at high levels of habitat loss. Our results demonstrate why knowledge of the patterns of connectivity across a species range is critical for predicting the effects of environmental change and provide empirical evidence for why connected migratory networks are commonly found in nature.     

Limited mitochondrial DNA diversity is indicative of a small number of founders of the German raccoon (Procyon lotor) population

The raccoon (Procyon lotor) has successfully invaded central Europe, despite the population apparently having been founded by a small number of individuals in two distinct populations in Germany. The ecological success of the invasion has been explained by raccoons being an adaptable, truly omnivorous species. However, the German raccoon population might have a larger number of founders and be more genetically diverse than assumed, as accidental or deliberate releases of household pets or individuals from zoos are relatively common. In the present study, we sequenced a 550-base-pair long fragment of the mitochondrial control region in 193 raccoons from Germany and neighbouring countries. We only identified six different haplotypes; of which, five were limited to Germany. Our results support the notions that the population was founded by a small number of females and that the German raccoons originate from two separate release events in central and eastern Germany. Additionally, however, we provide evidence for the presence of a distinct population in Saxony, eastern Germany. Further studies using different genetic markers are necessary to gain additional information on genetic diversity and population genetic structure.     

The effect of predation on the prevalence and aggregation of pathogens in prey

Although pathogens and predators have been widely used as bio-control agents against problematic prey species, little has been done to examine the prevalence and aggregation of pathogens in spatially structured eco-epidemiological systems. Here, we present a spatial model of a predator-prey/host-parasite system based on pair approximation and spatially stochastic simulations, with the predation pressure indicated by predator abundance and predation rates. Susceptible prey can not only be infected by contacting adjacent infected individuals but also by the global transmission of pathogens. The disease prevalence was found to follow a hump-shaped function in response to predation pressure. Moreover, predation pressure was not always negatively correlated with pathogen aggregation as proposed from empirical studies, but depending on the level of predation pressure. Highly connected site network facilitated the parasites infection, especially under high predation pressure. However, the connectivity of site network had no effect on the prevalence and aggregation of pathogens that can infect health prey through global transmission. It is thus possible to better design biological control strategies for target species by manipulating predation pressure and the range of pathogen transmission.