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1) Pathogens can spill over and infect new host species by overcoming a series of ecological and biological barriers. Hendra virus (HeV) circulates in Australian flying foxes and provides a data‐rich study system for identifying environmental drivers underlying spillover events. The frequency of spillover events to horses has varied interannually since the virus was first discovered in 1994. These observations suggest that HeV spillover events are driven, in part, by environmental factors, including loss of flying fox habitat and climate variability. 2) We explicitly examine the impact of environmental variation on the risk of HeV spillover at three spatial scales relevant to this system. We use a dataset of 60 spillover events and boosted regression tree methods to identify environmental features (including concurrent and lagged temperature, rainfall, vegetation indices, land cover, and climate indices) at three spatial scales (1‐km, 20‐km, 100‐km radii) associated with horse contacts and reservoir species ecology. 3) We find that temperature, local (1‐km radius) human population density, and landscape (100‐km radius) forest cover and pasture are the most influential environmental features associated with HeV spillover risk. By including multiple spatial scales and temporal lags in environmental features, we can more accurately quantify risk across space and time than with models that use a single scale. For example, high quality vegetation at the local scale and within a foraging radius (20‐km) in the concurrent month and previous years, combined with poorer quality vegetation at the landscape scale in the concurrent month increase risk of HeV spillover. These and other environmental associations likely influence the dynamic foraging behaviour of reservoir flying foxes and drive contacts that facilitate spillover into horse populations. 4) Synthesis and application: Current management of HeV spillover focuses on local‐scale interventions – primarily through vaccination and detection of infected horses. Our study finds that HeV spillover risk is also driven by environmental changes over much larger scales and demonstrates management practices would benefit from incorporating landscape interventions alongside local interventions. 

Abstract During recent decades, pathogens that originated in bats have become an increasing public health concern. A major challenge is to identify how those pathogens spill over into human populations to generate a pandemic threat 1 . Many correlational studies associate spillover with changes in land use or other anthropogenic stressors 2,3 , although the mechanisms underlying the observed correlations have not been identified 4 . One limitation is the lack of spatially and temporally explicit data on multiple spillovers, and on the connections among spillovers, reservoir host ecology and behaviour and viral dynamics. We present 25 years of data on land-use change, bat behaviour and spillover of Hendra virus from Pteropodid bats to horses in subtropical Australia. These data show that bats are responding to environmental change by persistently adopting behaviours that were previously transient responses to nutritional stress. Interactions between land-use change and climate now lead to persistent bat residency in agricultural areas, where periodic food shortages drive clusters of spillovers. Pulses of winter flowering of trees in remnant forests appeared to prevent spillover. We developed integrative Bayesian network models based on these phenomena that accurately predicted the presence or absence of clusters of spillovers in each of the 25 years. Our long-term study identifies the mechanistic connections between habitat loss, climate and increased spillover risk. It provides a framework for examining causes of bat virus spillover and for developing ecological countermeasures to prevent pandemics. 

Old World fruit bats (Chiroptera: Pteropodidae) provide critical pollination and seed dispersal services to forest ecosystems across Africa, Asia, and Australia. In each of these regions, pteropodids have been identified as natural reservoir hosts for henipaviruses. The genus Henipavirus includes Hendra virus and Nipah virus, which regularly spill over from bats to domestic animals and humans in Australia and Asia, and a suite of largely uncharacterized African henipaviruses. Rapid change in fruit bat habitat and associated shifts in their ecology and behavior are well documented, with evidence suggesting that altered diet, roosting habitat, and movement behaviors are increasing spillover risk of bat‐borne viruses. We review the ways that changing resource landscapes affect the processes that culminate in cross‐species transmission of henipaviruses, from reservoir host density and distribution to within‐host immunity and recipient host exposure. We evaluate existing evidence and highlight gaps in knowledge that are limiting our understanding of the ecological drivers of henipavirus spillover. When considering spillover in the context of land‐use change, we emphasize that it is especially important to disentangle the effects of habitat loss and resource provisioning on these processes, and to jointly consider changes in resource abundance, quality, and composition.