DOI: 10.5061/dryad.stqjq2cfh
title: Expression and mechanisms of behavioral plasticity in large mammals

description: Behavioral plasticity, the alteration of behavior in response to stimuli,
 is becoming increasingly important in the context of human-induced rapid
 environmental change. Theoretical and empirical studies suggest that the
 expression and magnitude of behavioral plasticity are likely facilitated
 or constrained primarily by two factors: environmental variation and
 endogenous traits such as body size. The contextual role of these factors
 on behavioral plasticity, however, is poorly understood; there are
 relatively few studies that have compared the magnitude and potential
 drivers of behavioral plasticity at different levels (i.e., population and
 individual) across species, especially in free-ranging animals with
 diverse behavioral traits such as large mammals. Here, we quantify and
 test potential hypotheses for the mechanisms underpinning behavioral
 plasticity at the individual and population level in response to variation
 in summer temperatures for 1,068 animal years in 17 populations across 9
 species of large mammals. All populations displayed behavioral plasticity
 in response to increased temperatures, modifying their relative selection
 for heat-relieving habitat attributes (e.g., elevation) and
 heat-generating behavior (i.e., movement speed). We found strong support
 for the hypothesis that the variability of the physical environment is an
 important driver of behavioral plasticity—both mean population behavioral
 plasticity and variation among individuals within each population in
 plasticity were lower with increased heterogeneity of habitat attributes
 such as tree cover. Yet, the variability in environmental conditions
 (i.e., the magnitude of the temperature increase) had no effect on
 behavioral plasticity within and among populations. We did not detect an
 effect of endogenous traits on the expression of behavioral plasticity;
 however, we note that data availability limited our tests of this
 hypothesis to a select few endogenous traits (body size, feeding guild,
 and sex of the tracked individuals) that predominantly vary at the species
 level, for which we had 1-3 replicate populations per species. Our results
 provide an integrative and generalizable understanding of the expression
 of behavioral plasticity among populations of large mammals in temperate
 environments and emphasize the important but nuanced role of environmental
 variation in determining the scope of behavioral plasticity in these
 populations.
description: # Expression and mechanisms of behavioral plasticity in large mammals
 Dataset DOI: [10.5061/dryad.stqjq2cfh](10.5061/dryad.stqjq2cfh) ##
 Description of the data and file structure Animal movement data from 1068
 unique animal-years representing 575 individuals of 17 populations of 9
 species of large mammals collected between 2001 and 2019: Rocky Mountain
 bighorn sheep (*Ovis canadensis canadensis*), American bison (*Bison bison
 bison*), North American cougar (*Puma concolor cougar*), Rocky Mountain
 elk (*Cervus canadensis nelsoni*), Shiras moose (*Alces alces shirasi*),
 mountain goat (*Oreamnos americanus*), Rocky Mountain mule deer
 (*Odocoileus hemionus hemionus*), American pronghorn (*Antilocapra
 americana americana*), and gray wolf (*Canis lupus*). ### Files and
 variables #### File: datasets_metadata.xlsx **Description:** metadata for
 each dataset #### File: "..."_result_SC.csv
 **Description:** beta coefficients from Resource Selection Functions for
 each large mammal population. ##### Variables * species: species name *
 population: population name * spec_pop: combined species and population
 name * id_yr: individual ID * n_all: total number of points * n_used:
 total number of used points * n_speed: total number of points used to
 calculate movement speed * relatTmax_HR_Lquant: lower quantile for
 relative Tmax in individual home range (relative daily max. temperature) *
 relatTmax_HR_Uquant: upper quantile for relative Tmax in individual home
 range (relative daily max. temperature) * coefs: names of the RSF
 coefficients  * Estimate: estimate beta coefficient value * SE: standard
 error for the estimated beta coefficient value * z: z-score for the
 estimated beta coefficient value * P: p-value for the estimated beta
 coefficient value #### File: "..."_data.csv
 **Description:** animal location data used to fit the RSF models for each
 study population ##### Variables * id: id * species: species name *
 population: population name * spec_pop: combined species and population
 name * id_yr: individual ID * sex: individual sex (M/F) * pack: pack ID
 (if applicable for species) * timeOfDay: day or night * used: 1 used, 0
 random available locations * tmax: daily max temperature (C) * relatTmax:
 relative daily max temperature  * tmax_HR: daily max temperature across
 the home range (C) * relatTmax_HR:  relative daily max temperature across
 the home range * elev: elevation (meters) * trasp: topographic
 solar-radiation index * cti: compound topographic index * perCover:
 percent tree canopy cover * dist: step length (meters) * dt: time between
 locations (fix rate) * speed: movement speed (ms-1) * speed_log: log of
 movement speed * elev_SC: scaled and centered elevation * trasp_SC: scaled
 and centered topographic solar-radiation index * cti_SC: scaled and
 centered compound topographic index * perCover_SC: scaled and
 centered percent tree canopy cover * speed_log_SC: scaled and centered log
 movement speed #### File:
 10_explain_visualize_variation_in_behavioral_plasticity.R
 **Description:** R code #### File:
 11_more_visualize_variation_in_behavioral_plasticity.R **Description:** R
 code #### File: 04_extract_tmax_GIS_data.R **Description:** R code ####
 File: 07_correlation_matrix_PCA_behavioral_responses.R **Description:** R
 code #### File: 05_calculate_speed_scale_center_data.R **Description:** R
 code #### File:
 09_visualize_individual_behavioral_plasticity_coefficients.R
 **Description:** R code #### File:
 02c_sensitivity_analysis_run_models_visualize.R **Description:** R code
 #### File: 06_run_RSF_speed_models.R **Description:** R code #### File:
 03_generate_BB_HRs_random_points.R **Description:** R code #### File:
 02b_sensitivity_analysis_extract_GIS_tmax.R **Description:** R code ####
 File: 00_prep_GPS_data_bighorn.R **Description:** R code #### File:
 01_clean_filter_GPS_data.R **Description:** R code #### File:
 00_prep_GPS_data_wolf.R **Description:** R code #### File:
 02a_sensitivity_analysis_generate_HR_random_points.R **Description:** R
 code #### File: 00_prep_GPS_data_pronghorn.R **Description:** R code ####
 File: 00_prep_GPS_data_muledeer.R **Description:** R code #### File:
 00_prep_GPS_data_mtngoat.R **Description:** R code #### File:
 00_prep_GPS_data_cougar.R **Description:** R code #### File:
 08_quantify_population_plasticity_metrics.R **Description:** R code ##
 Code/software All data and code can be accessed using the R environment.