Diverse hosts, diverse immune systems: evolutionary variation in bat immunology

The ability of multiple bat species to host zoonotic pathogens without showing disease has fostered growing interest in bat immunology, indicating ways immune systems may differ between bats and other vertebrates. However, interspecific variation in immunological diversity among bats has only begun to be recognized. The order Chiroptera accounts for over 20% of all mammal species and shows extreme diversity in a suite of correlated ecological traits, such that bats should not be expected to be immunologically homogenous. Here, we review the ecological and evolutionary diversity of chiropteran hosts and highlight case studies emphasizing the range of immune strategies thus far observed across bat species, including responses to SARS-CoV-2. Next, we synthesize and propose hypotheses to explain this immunological diversity, focused on pathogen exposure, biogeography, host energetics, and environmental stability. Lastly, we analyze immunology citations across bat species to motivate discussion of outstanding research priorities. Broad sampling is needed to remedy current biases, as only a fraction of bat species has been immunologically studied. Such work should integrate methodological advancements, in vitro and in vivo studies, and phylogenetic comparative methods to robustly test evolutionary hypotheses and understand the drivers and consequences of immunological diversity among bats.