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Broadcasted social calls attracted hoary bats during both the spring and fall migration. Broadcasting conspecific social calls increased hoary bat capture rates at netting sites intentionally removed from normal capture locations. We had very low capture rates during control periods, because we intentionally placed nets in locations removed from flyways to reduce incidental captures. Moreover, capture rates of hoary bats tend to be low even in many locations where they are known to occur24,25, and capture rates of approximately one bat per hour in a single mist net suggest a very strong attraction response to broadcasted calls.Hoary bat activity, as measured by acoustic monitoring was not associated with increased capture rates in response to call broadcasting. However, subsequent research has shown that hoary bats periodically use higher frequency, inconspicuous calls, or do not constantly echolocate during the fall, which may mean acoustic monitoring did not effectively measure hoary bat activity in the vicinity of our trials26,27. We recorded substantially higher acoustic activity during the spring migration, which could represent either more hoary bats and/or bat activity, or a seasonal difference in echolocation or flight behavior such as differences in flight altitude27. It remains unknown if hoary bats use inconspicuous calls or fly in silence during spring migration or other times of year other than the fall when these inconspicuous echolocation behaviors were observed, and seasonally variable behavior could affect detectability or exposure to our playback trials in ways not captured by our acoustic activity covariate. In addition, while we did audibly hear social calls of hoary bats during the fall, we did not record any during fieldwork for this study, which may be an artifact or due to differences in social behavior, context, or number of hoary bats present in the area during our trials.We only captured one female during trials in New Mexico, and were unable to locate any females during the fall migration in coastal regions of California, despite high concentrations of males in the area during what is presumably the mating season. In New Mexico, during spring migration, females migrate through the study area before males28, with very little temporal overlap. As a result, we were unable to determine sex specific responses to call playback, however we have subsequently captured several female hoary bats and Ope’ape’a (Hawaiian hoary bat, L. semotus) using call playback during capture and radio-tracking studies (GAR, pers. obs.).It is difficult to elucidate the meaning of social calls based on the behaviors observed in the field. In bats, social call complexity often reflects social behavior complexity, with a range of uses including but not limited to attracting mates, locating pups within colonies, defending roosting or foraging territory, and attracting bats to roosts10. Attraction to conspecific call broadcasting could indicate positive social interactions (e.g., maintaining group cohesion or investigation) or agonistic behavior (e.g., hoary bats approaching to chase conspecific bats), as has been observed in other bat species29 and in hoary bats during the maternity season30. We did not observe any obvious instances of aggressive hoary bat interactions, and the social calls differ from hisses and clicks that hoary bats use defensively (Fig. 2). We would also audibly hear pairs of hoary bats calling in close proximity to each other, with no indication of aggressive or territorial responses, and these calls being low frequency and audible to humans means that they attenuate at greater distances than hoary bat echolocation calls.Aggressive or territorial interactions in many taxa are often driven by seasonally variable contexts, such as mating, defending food resources, or rearing of young. It may be unlikely that migrating hoary bats would expend energy defending territory during migration when they are utilizing roosts or foraging habitat for such limited periods of time (i.e., a few hours to a day). During active migration birds are often not territorial even when foraging at stopover sites31, and there may be benefits to maintaining group cohesion during migration including navigation and identification of favorable habitat. It is unknown if hoary bats utilize stopover sites for refueling during migration. However the silver-haired bat Lasionycteris noctivagans was found to utilize a migration stopover site in Long Point, Canada, where they opportunistically foraged for short periods of time (1 to 2 days32). Tracking studies would be required to determine temporal patterns of site usage by individual bats to examine stopover behavior.As we had recorded most of our initial social calls during late summer and early fall when hoary bats mate21, we had originally hypothesized that these social calls were associated with mating behavior, which would have been consistent with observations in this study had we found both increased attraction during the fall, and less attraction to calls during the spring. However, social calls attracted hoary bats effectively during both the spring and fall migration. In addition, from acoustic recordings and capture observations in the field, hoary bats produced many social calls during the spring migration when only males were present. There is a possibility, due to our lack of understanding of the mating systems of hoary bats that some mating may continue into the spring. However the majority of taxonomic, physiological, and observational data suggests mating behavior ends by the spring migration19,33, and the majority of females are already pregnant when travelling through New Mexico28. While hoary bats may or may not use social calls as a component of mating behavior, social calls recorded during the spring likely serve purposes not associated with mating.Previous studies describe the hoary bat as solitary throughout most of the year, which would imply only brief social interactions limited to mating or association with offspring, and the many historical accounts of aggregations of hoary bats are thought to be related to mating behavior20,33,34. However the use of, and attraction to, social calls during both spring and fall migration supports that these calls are used for social interactions beyond mating behavior. Further research may determine if hoary bats use these social calls to maintain group cohesion during migration, and what, if any, relationships exist between individual hoary bats that appear to be migrating together. Baerwald and Barclay35 found that geographic and genetic relationships of hoary bats and silver-haired bat carcasses collected at wind turbines were not more closely related than expected by chance, which provides some evidence that groups of migrating hoary bats may not form based on kinship.Many studies hoping to elucidate the causes of fatalities at wind energy facilities have focused only on the fall migration period when bats are most often killed13,20,36. However hoary bats migrate during the spring as well, when they do not suffer high fatality rates. Investigating the spring migration presents a valuable baseline to compare behavioral changes and other factors that may place hoary bats or other impacted species at risk. If social behavior makes a major contribution to the risk of fatalities at wind energy developments, then social behavior should differ between spring and fall migration. We did not find a large difference in response to social calls between seasons. While this represents just an initial study into the social calling behavior of hoary bats during migration, it provides some conclusions to guide subsequent investigations: (1) detecting hoary bat social calls does not necessarily indicate mating behavior, and (2) researchers should be cautious in interpreting evidence of social interactions during the fall at wind energy sites as evidence of mating behavior as in the mating landmarks hypothesis22,37. Because it can separate out mating from other behavioral components, comparing spring and fall migration can benefit the investigation of social and other behaviors in hoary bats and other migratory species. Comparing flight behavior, diet, roost selection, hormonal and physiological changes, and further studies of social interactions including scent and, between the spring and fall migration will allow researchers to elucidate which behaviors change seasonally and which may underlie seasonal patterns of wind turbine fatalities. Additionally, exploring social attraction to audible sounds produced by turbines or other potential signals that could seasonally elicit social attraction could lead to additional insights.Hoary bats have proven challenging to capture and study in many locations across their range24, driven by their solitary tree roosting behavior and as they often fly out of the reach of mist nets or ground-based acoustic monitoring stations36,38. Using call broadcasting to increase capture rates can be a useful research tool, especially in locations where the habitat does not provide any ideal capture locations. Using this technique we have captured hoary bats on coastal sand dunes, in large open fields, and in groves of Eucalyptus trees adjacent to wind energy sites, all of which would normally yield low bat capture success without the use of lures. The ability to capture hoary bats more reliably is a great asset for research and conservation throughout the range of hoary bats.Our study tested the use of social call playback as a methodology to study the social behavior of hoary bats during migration, and the utility of using call playback as a research tool and acoustic lure for hoary bats. Increasing capture rates from conspecific social call playback during mating and non-mating season indicates social interactions during both migratory periods, despite the solitary roosting behavior of this species. Future studies to elucidate the behavioral function of these calls, and response during non-migratory seasons could refine our understanding of social behaviors of this elusive bat species. More

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Here we provide an overview of the key elements of our framework including describing the contact function that links economic activities to contacts, the SIRD (Susceptible-Infectious-Recovered-Dead) model, the dynamic economic model governing choices, and calibration. The core of our approach is a dynamic optimization model of individual behavior coupled with an SIRD model of infectious disease spread. Additional details are found in the SI.Contact functionWe model daily contacts as a function of economic activities (labor supply, measured in hours, and consumption demand, measured in dollars) creating a detailed mapping between contacts and economic activities. For example, all else equal, if a susceptible individual reduces their labor supply from 8 to 4 h, they reduce their daily contacts at work from 7.5 to 3.75. Epidemiological data is central to calibrating this mapping between epidemiology and economic behavior. Intuitively, the calibration involves calculating the mean number of disease-transmitting contacts occurring at the start of the epidemic and linking it to the number of dollars spent on consumption and hours of labor supplied before the recession begins.We use an SIRD transmission framework to simulate SARS-CoV-2 transmission for a population of 331 million interacting agents. This is supported by several studies (e.g.,77,78) that identify infectiousness prior to symptom onset. We consider three health types m ∈ {S, I, R} for individuals, corresponding to epidemiological compartments of susceptible (S), infectious (I), and recovered (R). Individuals of health type m engage in various economic activities ({A}_{i}^{m}), with i denoting the activities modeled. One of the ({A}_{i}^{m}) is assumed to represent unavoidable other non-economic activities, such as sleeping and commuting, which occur during the hours of the day not used for economic activities (see SI 2.3.1). Disease dynamics are driven by contacts between susceptible and infectious types, where the number of susceptible-infectious contacts per person is given by the following linear equation:$${{{{{{{{mathscr{C}}}}}}}}}^{SI}({{{{{{{bf{A}}}}}}}})=mathop{sum}limits_{i}{rho }_{i}{A}_{i}^{S}{A}_{i}^{I}$$
(1)
while similar in several respects to prior epi-econ models15,16,74, a methodological contribution is that ρi converts hours worked and dollars spent into contacts. For example, ρc has units of contacts per squared dollar spent at consumption activities, while ρl has units of contacts per squared hour worked.We also consider robustness to different functional forms in Fig. 6F, G as a reduced-form way to consider multiple consumption and labor activities with heterogeneous contact rates. Formally:$${{{{{{{{mathscr{C}}}}}}}}}^{SI}({{{{{{{bf{A}}}}}}}})=mathop{sum}limits_{i}{rho }_{i}{({A}_{i}^{S}{A}_{i}^{I})}^{alpha },$$
(2)
where α  > 1 (convex) corresponds to a contact function where higher-contact activities are easiest to reduce or individuals with more contacts are easier to isolate. α  More

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