DOI: 10.5061/dryad.ksn02v7gj
title: Data from: Diverse hosts, diverse immune systems: evolutionary variation in bat immunology

description: The ability of multiple bat species to host zoonotic pathogens without
 often 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. We then analyze immunology-related citations per species across
 bats to motivate discussion of key 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.
description: # Diverse hosts, diverse immune systems: evolutionary variation in bat
 immunology Dataset DOI: [10.5061/dryad.ksn02v7gj](10.5061/dryad.ksn02v7gj)
 ## Description of the data and file structure The data CSV contains
 species-level data across bats from the COMBINE, PanTHERIA, and VIRION
 databases, standardized to the mammalian taxonomy of Upham et al. 2019.
 The R script contains code for downloading VIRION data, matching to the
 bat taxonomy above, and aggregating bat–virus interactions to viral family
 richness. The R script also aggregates total and immunology-related
 citation counts per bat species. ### Files and variables #### File:
 Becker\_et\_al\_2025\_ANYAS\_R\_script.R **Description:**  #### File:
 Becker\_et\_al\_2025\_ANYAS\_data.csv **Description:** Blank cells are
 coded as NA ##### Variables * tip: bat species name * clade: bat clade
 label from Upham et al. 2019 * fam: bat family from Upham et al. 2019 *
 gen: bat genus  from Upham et al. 2019 * vfams: number of viral families
 (derived from VIRION) * adult_mass_g: adult body mass in grams from
 COMBINE * adult_brain_mass_g: adult brain mass in grams from COMBINE *
 adult_body_length_mm: adult body length in mm from COMBINE *
 adult_forearm_length_mm: adult forearm length in mm from COMBINE *
 max_longevity_d: maximum longevity in days from COMBINE * maturity_d: age
 of sexual maturity in days from COMBINE * female_maturity_d: age of female
 sexual maturity in days from COMBINE * male_maturity_d: age of male sexual
 maturity in days from COMBINE * age_first_reproduction_d: age of first
 reproduction in days from COMBINE * gestation_length_d: duration of
 gestation period in days from COMBINE * teat_number_n: number of teats
 from COMBINE * litter_size_n: number of offspring per female from COMBINE
 * litters_per_year_n: numbers of litters per year from COMBINE *
 interbirth_interval_d: time between reproductive events in days from
 COMBINE * neonate_mass_g: mass of neonates in grams from COMBINE *
 weaning_age_d: age at which weaning ends in days from COMBINE *
 weaning_mass_g: weight at weaning in grams from COMBINE *
 generation_length_d: average age of parents in days from COMBINE *
 dispersal_km: distance from natal origin in km from COMBINE *
 density_n_km2: population density in km2 from COMBINE *
 hibernation_torpor: binary for whether the species hibernates or undergoes
 torpor (1) from COMBINE * fossoriality: binary for ground dwelling or
 fossorial (1) from COMBINE * home_range_km2: home range size in km from
 COMBINE * social_group_n: number of individuals per social group from
 COMBINE * dphy_invertebrate: percent of invertebrates in diet from COMBINE
 * dphy_vertebrate: percent of vertebrates in diet from COMBINE *
 dphy_plant: percent of plants in diet from COMBINE * det_inv: percent of
 invertebrates in diet from COMBINE * det_vend: percent of endotherms in
 diet from COMBINE * det_vect: percent of ectotherms in diet from COMBINE *
 det_vfish: percent of fish in diet from COMBINE * det_vunk: percent of
 unknown in diet from COMBINE * det_scav: percent of scavenge/carcasses in
 diet from COMBINE * det_fruit: percent of fruit in diet from COMBINE *
 det_nect: percent of nectar in diet from COMBINE * det_seed: percent of
 seeds in diet from COMBINE * det_plantother: percent of other plants in
 diet from COMBINE * det_diet_breadth_n: number of dietary categories from
 COMBINE * trophic_level: herbivore (1), omnivore (2), or carnivore (3)
 from COMBINE * foraging_stratum: assignment to one of five foraging strata
 from COMBINE (M: marine, G: ground, S: scansorial, Ar: arboreal, A:
 aerial) * activity_cycle: nocturnal (1), mix (2), or diurnal (3) from
 COMBINE * freshwater: binary for spending significant time in freshwater
 (1) from COMBINE * marine: binary for spending significant time in marine
 habitats (1) from COMBINE * terrestrial_non.volant: binary for spending
 significant time on land (1) from COMBINE * terrestrial_volant: binary for
 capable of powered flight (1) from COMBINE * upper_elevation_m: upper
 elevation of species range in meters from COMBINE * lower_elevation_m:
 lower elevation of species range in meters from COMBINE *
 altitude_breadth_m: difference in elevation of species range in meters
 from COMBINE * island_dwelling: binary for dwelling on islands (1) from
 COMBINE * island_endemicity: score of island endemicity from COMBINE *
 disected_by_mountains: binary for range being dissected by mountains
 (1) from COMBINE * glaciation: binary for historical exposure to
 glaciation (1) from COMBINE * biogeographical_realm: biogeographic realm
 categories * habitat_breadth_n: number of habitat categories from the IUCN
 from COMBINE * X1.1_ActivityCycle: nocturnal only (1), mixed (2), or
 diurnal only (3) from PanTHERIA * X5.1_AdultBodyMass_g: adult body mass in
 grams from PanTHERIA * X8.1_AdultForearmLen_mm: adult forearm length in mm
 from PanTHERIA * X13.1_AdultHeadBodyLen_mm: adult head to base of tail
 length in mm from PanTHERIA * X2.1_AgeatEyeOpening_d: age of eye opening
 in days from PanTHERIA * X3.1_AgeatFirstBirth_d: age at first birth in
 days from PanTHERIA * X18.1_BasalMetRate_mLO2hr: basal metabolic rate
 from PanTHERIA * X5.2_BasalMetRateMass_g: mass of individuals from BMR
 measures from PanTHERIA * X6.1_DietBreadth: number of dietary categories
 from PanTHERIA * X7.1_DispersalAge_d: age of dispersal in days
 from PanTHERIA * X9.1_GestationLen_d: length of gestation period in days
 from PanTHERIA * X12.1_HabitatBreadth: number of habitat layers used
 from PanTHERIA * X22.1_HomeRange_km2: home range size (individuals or
 groups) in km2 from PanTHERIA * X22.2_HomeRange_Indiv_km2: home range size
 of individuals in km2 from PanTHERIA * X14.1_InterbirthInterval_d: days
 between births in days from PanTHERIA * X15.1_LitterSize: individuals per
 litter from PanTHERIA * X16.1_LittersPerYear: numbers of litters per year
 from PanTHERIA * X17.1_MaxLongevity_m: maximum lifespan in months
 from PanTHERIA * X5.3_NeonateBodyMass_g: body mass of neonates in grams
 from PanTHERIA * X13.2_NeonateHeadBodyLen_mm: neonate head to base of tail
 length in mm from PanTHERIA * X21.1_PopulationDensity_n.km2: population
 size in km2 from PanTHERIA * X10.1_PopulationGrpSize: population group
 size from PanTHERIA * X23.1_SexualMaturityAge_d: age of sexual maturity in
 days from PanTHERIA * X10.2_SocialGrpSize: social group size
 from PanTHERIA * X24.1_TeatNumber: number of teats from PanTHERIA *
 X12.2_Terrestriality: binary terrestrial (1) or not (0) from PanTHERIA *
 X6.2_TrophicLevel: herbivore (1), omnivore (2), or carnivore (3)
 from PanTHERIA * X25.1_WeaningAge_d: age when weaning ends in days
 from PanTHERIA * X5.4_WeaningBodyMass_g: body mass when weaning in grams
 from PanTHERIA * X13.3_WeaningHeadBodyLen_mm: weaning head to base of tail
 length in mm from PanTHERIA * X5.5_AdultBodyMass_g_EXT: extrapolated adult
 body mass in grams from PanTHERIA * X16.2_LittersPerYear_EXT: extrapolated
 litters per year from PanTHERIA * X5.6_NeonateBodyMass_g_EXT: extrapolated
 neonate body mass in grams from PanTHERIA * X5.7_WeaningBodyMass_g_EXT:
 extrapolated weaning body mass in grams from PanTHERIA *
 X26.1_GR_Area_km2: geographic range size in km2 from PanTHERIA *
 X26.2_GR_MaxLat_dd: maximum latitude of geographic range in decimal
 degrees from PanTHERIA * X26.3_GR_MinLat_dd: minimum latitude of
 geographic range in decimal degrees from PanTHERIA *
 X26.4_GR_MidRangeLat_dd: median latitudinal extent of range in decimal
 degrees from PanTHERIA * X26.5_GR_MaxLong_dd: maximum longitude of
 geographic range in decimal degrees from PanTHERIA *
 X26.6_GR_MinLong_dd: minimum longitude of geographic range in decimal
 degrees from PanTHERIA * X26.7_GR_MidRangeLong_dd: median longitudinal
 extent of range in decimal degrees from PanTHERIA *
 X27.1_HuPopDen_Min_n.km2: minimum human population density in range in km2
 from PanTHERIA * X27.2_HuPopDen_Mean_n.km2: mean human population density
 in range in km2 from PanTHERIA * X27.3_HuPopDen_5p_n.km2: 5th percentile
 human population density in range in km2 from PanTHERIA *
 X27.4_HuPopDen_Change: mean rate of increase in human population density
 in range (km2) from PanTHERIA * X28.1_Precip_Mean_mm: mean monthly
 rainfall (mm) from PanTHERIA * X28.2_Temp_Mean_01degC: mean monthly
 temperature (0.1°C)  from PanTHERIA * X30.1_AET_Mean_mm: mean actual
 evapotranspiration (AET) rate in mm from PanTHERIA * X30.2_PET_Mean_mm:
 mean potential evapotranspiration (PET) rate in mm from PanTHERIA * cites:
 citation count per species binomial from PubMed * icites:
 immunology-related citation count per species binomial from PubMed ###
 Version changes * 6/9/2025: Updated virus family richness with up-to-date
 VIRION data (see https://github.com/viralemergence/virion)