The global distribution of Bacillus anthracis and associated anthrax risk to humans, livestock, and wildlife

Abstract

Bacillus anthracis is a spore-forming, Gram-positive bacterium responsible for anthrax, an acute and commonly lethal infection that most significantly affects grazing livestock, wild ungulates and other herbivorous mammals, but also poses a serious threat to human health1, 2. The geographic extent of B. anthracis endemism is still poorly understood, despite multi-decade research on anthrax epizootic and epidemic dynamics around the world3, 4. Several biogeographic studies have focused on modeling environmental suitability for anthrax at local or national scales5-9, but many countries have limited or inadequate surveillance systems, even within known endemic regions. Here we compile an extensive global occurrence dataset for B. anthracis, drawing on confirmed human, livestock, and wildlife anthrax outbreaks. With these records, we use boosted regression trees10, 11 to produce the first map of the global distribution of B. anthracis as a proxy for anthrax risk. Variable contributions to the model support pre-existing hypotheses that environmental suitability for B. anthracis depends most strongly on soil characteristics such as pH that affect spore persistence, and the extent of seasonal fluctuations in vegetation, which plays a key role in transmission for herbivores12, 13. We apply the global model to estimate that 1.83 billion people (95% credible interval: 0.59--4.16 billion) live within regions of anthrax risk, but most of that population faces little occupational exposure to anthrax. More informatively, a global total of 63.8 million rural poor livestock keepers (95% CI: 17.5--168.6 million) and 1.1 billion livestock (95% CI: 0.4--2.3 billion) live within vulnerable regions. Human risk is concentrated in rural areas, and human and livestock vulnerability are both concentrated in rainfed systems throughout arid and temperate land across Eurasia, Africa, and North America. We conclude by mapping where anthrax risk overlaps with vulnerable wild ungulate populations, and therefore could disrupt sensitive conservation efforts for species like bison, pronghorn, and saiga that coincide with anthrax-prone, mixed-agricultural landscapes. Anthrax is a zoonotic disease caused by the Gram-positive bacterium Bacillus anthracis, a generalist soil-transmitted pathogen found on every inhabited continent14, and several islands including Haiti and parts of the Philippines and Indonesia. Worldwide, an estimated 20,000 to 100,000 cases of anthrax occur annually, mostly in poor rural areas15. In clinical presentations of anthrax, case fatality rates are a function of exposure pathway. Respiratory exposure from spore inhalation is important the context of bioterrorism, but is highly uncommon, and accounts for a negligible fraction of the global burden of anthrax cases. Cutaneous exposure to B. anthracis accounts for the majority of human cases worldwide, and typically presents with low mortality; gastrointestinal exposure accounts for the remainder and presents with intermediate to high fatality rates. Cutaneous and gastrointestinal cases of anthrax are most commonly caused by handling and slaughtering infected livestock, or butchering and eating contaminated meat; untreated gastrointestinal cases likely account for most human mortality from anthrax.14-16