A multi-scale assessment of animal aggregation patterns to understand increasing pathogen seroprevalence

Understanding how animal density is related to pathogen transmission is important to develop effective disease control strategies, but requires measuring density at a scale relevant to transmission. However, this is not straightforward or well-studied among large mammals with group sizes that range several orders of magnitude or aggregation patterns that vary across space and time. To address this issue, we examined spatial variation in elk (Cervus canadensis) aggregation patterns and brucellosis across 10 regions in the Greater Yellowstone Area where previous studies suggest the disease may be increasing. We hypothesized that rates of increasing brucellosis would be better related to the frequency of large groups than mean group size or population density, but we examined whether other measures of density would also explain rising seroprevalence. To do this, we measured wintering elk density and group size across multiple spatial and temporal scales from monthly aerial surveys. We used Bayesian hierarchical models and 20 years of serologic data to estimate rates of increase in brucellosis within the 10 regions, and to examine the linear relationships between these estimated rates of increase and multiple measures of aggregation. Brucellosis seroprevalence increased over time in eight regions (one region showed an estimated increase from 0.015 in 1991 to 0.26 in 2011), and these rates of increase were positively related to all measures of aggregation. The relationships were weaker when the analysis was restricted to areas where brucellosis was present for at least two years, potentially because aggregation was related to disease-establishment within a population. Our findings suggest that (1) group size did not explain brucellosis increases any better than population density and (2) some elk populations may have high densities with small groups or lower densities with large groups, but brucellosis is likely to increase in either scenario. In this case, any one control method such as reducing population density or group size may not be sufficient to reduce transmission. This study highlights the importance of examining the density-transmission relationship at multiple scales and across populations before broadly applying disease control strategies.