Dynamic risk from Mexican wolves and mountain lions influences elk foraging behavior

Published Mar 03, 2026 on Dryad. https://doi.org/10.5061/dryad.0000000hx

Data files

Mar 03, 2026 version files 1.96 MB

Focal_repeats.csv

977.31 KB
FocalCovars.csv

602.07 KB
ISSCovars.csv

342.61 KB
README.md

36.31 KB

Abstract

Foraging time is a major component of ungulate activity budgets but can be limited by anti‐predator behaviors (e.g., vigilance). Multitasking can reduce the nutritional costs of vigilance under heightened predation risk, but this may depend on the response of prey to risk from multiple predators across a complex spatiotemporal landscape. Mexican gray wolves (Canis lupus baileyi) and mountain lions (Puma concolor) are primary predators for elk (Cervus canadensis) in the Mexican wolf experimental population area in east‐central Arizona and west‐central New Mexico.

Authors: Olson, Julia E.; Thompson, Cara J.; Farley, Zachary J.; Martinez, Samuel I.; Boyle, Scott T.; Tatman, Nicole M.; DeVos, James G.; Liley, Stewart D.; and Cain, James W. III

Date Created: 04/08/2025

Corresponding author: Dr. James W. Cain; james.cain@ag.tamu.edu

This repository includes data files used in modeling and analysis of five data types:
Individual behavior, herd behavior, individual Forage Intake Rate (FIR), individual Accepted Food Abundance (AFA), and individual multitasking probability. Descriptions of these data types can be found in the introduction and methods sections of manuscript.

The data files include:

FocalCovars.csv: behavioral data and covariates for individual/focal elk.

ISSCovars.csv: behavioral data and covariates for elk herds collected as Instantaneous Scan Samples (ISS).

Focal_repeats.csv: raw focal behavior repeat data used to create binary multitasking/vigilant variable prior to modeling.

Variable descriptions and structure for FocalCovars.csv:
All continuous covariates were centered and scaled prior to modeling, and all categorical covariates were converted to factors.

Column header	Data type	Description
HerdKey	Categorical	Identifier for unique elk herds
DateTime_Start	Date	Date and time observation began, formatted %M/%D/%Y %H:%M
Elk_Latitude	Coordinates	Latitude of approximated location of elk herd in WGS84
Elk_Longitude	Coordinates	Longitude of approximated location of elk herd in WGS84
FocalKey	Categorical	Identifier for unique focal elk within herd
Focal_DateTime_Start	Date	Time focal observation began, formatted %H:%M.%S
Focal_Steps	Numerical	Total number of steps taken by focal elk during observation
Focal_Duration	Numerical	Total duration of focal observation in seconds
Propforage	Numerical	Proportion of focal duration that the focal elk spent actively foraging
Propmulti	Numerical	Proportion of focal duration that the focal elk spent multitasking
Propvig	Numerical	Proportion of focal duration that the focal elk spent intensely vigilant
Propother	Numerical	Proportion of focal duration that the focal elk spent in miscellaneous other behaviors
Proptravel	Numerical	Proportion of focal duration that the focal elk spent traveling between forage patches
Focal_Steps_AFA	Numerical	Total number of steps focal elk took while actively foraging, vigilant, or multitasking
Focal_Stepspermin_AFA	Numerical	Focal_Steps_AFA divided by Focal_Duration_AFA, divided by 60
Focal_Duration_AFA	Numerical	Time focal elk spent foraging, intensely vigilant, or multitasking during focal observation in seconds
Season	Categorical	Season that the observation took place during
SeasonWolf	Categorical	Seasonal designations as defined for Mexican wolf Habitat Selection Function and Utilization Distributions, used to extract these values from the appropriate time period for observations
Month	Categorical	Numerical month that the observation took place during
Focal_Diel	Categorical	Diel period during which focal observation began
Diel_Period	Categorical	Diel period during which overall observation began
Observer	Categorical	Initials of field personnel actively observing elk
Wind_Speed	Categorical	Wind speed estimated as none, low, medium, or high, or NA if not recorded
Focal_Sex	Categorical	Sex of focal elk (aCow = adult female > 2 years, aBull = adult male > 2 years, yearling = male or female 1-2 years old)
Focal_Peripheral	Binary	Variable indicating whether focal elk was on the periphery of the herd (1) or not (0)
Focal_Exposure	Numerical	Average number of focal elk's four sides that were exposed to the edge of the herd
Veg	Categorical	Vegetation type cover type (grassland = 0, aspen = 1, oak/shrub = 3, pinon-juniper = 4, ponderosa = 5, mixed conifer = 6, and wet meadow or pasture = 8)
VH	Categorical	Vegetation height reclassified as unobstructed = 0, obstructed low = 1, and obstructed high = 2
VC	Binary	Vegetation cover reclassified as open (<30% cover) = 0 and closed (>30% cover) = 1
HC	Categorical	Combined vegetative cover and height into horizontal cover (no cover = 0, partial cover = 1, high cover = 2)
Distance_Obs	Numerical	Distance between estimated location of elk herd and the observer's location in meters
N_Total	Numerical	Total number of elk (calves, yearlings, and adults) in herd
Ratio_Calf_Cow	Numerical	Total number of calves in the herd divided by the total number of adult female elk in the herd
Ratio_Bull_Cow	Numerical	Total number of adult male elk divided by the total number of adult female elk in the herd
Focal_AllSides_Av	Numerical	Average number of neighboring elk on each side of the focal elk
Focal_Pos_Within5_Av	Numerical	Average number of neighboring elk within five bodylengths of the focal elk
N_Humans	Numerical	Number of cars recorded passing on road during observation
Water	Numerical	Distance to perennial water bodies in meters
R3Density	Numerical	Density of type 3 (primitive/low use) roads in a 1.5 km^2 area
Road3	Numerical	Distance to type 3 (primitive/low use) road in meters
AllRDensity	Numerical	Density of all road types in a 1.5 km^2 area
AllRoad	Numerical	Distance to road of all types in meters
HumanDensity	Numerical	Human population density
Road1	Numerical	Distance to type 1 (paved/high use) road in meters
Road2	Numerical	Distance to type 2 (dirt or gravel/moderate use) road in meters
Road1Density	Numerical	Density of type 1 (paved/high use) roads in 1.5 km^2 area
Road2Density	Numerical	Density of type 2 (dirt or gravel/moderate use) roads in 1.5 km^2 area
Elev	Numerical	Vertical elevation in meters
Northness	Numerical	Index of slope northness
Slope	Numerical	Slope where elk herd was located
VRM	Numerical	Vector Ruggedness Measurement
Open500	Numerical	Number of pixels with open canopy (<30% canopy cover) in a rolling 500 m window created with 2016 LANDFIRE data
RecTrl	Numerical	Distance to recreation sites or trails in meters
Edge	Numerical	Distance to forest ecotone in meters with a 50 m buffer
Private	Numerical	Distance to private land boundary in meters
Open250	Numerical	Number of pixels with open canopy (<30% canopy cover) in a rolling 250 m window created with 2016 LANDFIRE data
NDVI	Numerical	Normalized Difference Vegetation Index obtained from pre-processed MODIS satellite data
irg	Numerical	Instantaneous Rate of Greenup of vegetation modeled from NDVI data
Max_BS	Numerical	Severity of burn in past 17 years (0 if unburned)
Burn	Binary	Burned in past 17 years = 1, unburned in past 17 years = 0
FireHist	Categorical	Combined fire severity and age ( 0 = unburned in past 17 years, 1 = low-severity in past 5 years, 2 = moderate/high-severity in past five years, 3 = low-severity in past 6-17 years, 4 = moderate/high-severity in past 6-17 years)
TPI	Numerical	Topographic Position Index
TRI	Numerical	Topographic Ruggedness Index
Openness_2021	Numerical	Percent canopy openness from 2021 LANDFIRE data
Open480_2021	Numerical	Number of pixels with open canopy (<30% canopy cover) in a rolling 480 m window created with 2021 LANDFIRE data
Open240_2021	Numerical	Number of pixels with open canopy (<30% canopy cover) in a rolling 240 m window created with 2021 LANDFIRE data
Open90_2021	Numerical	Number of pixels with open canopy (<30% canopy cover) in a rolling 90 m window created with 2021 LANDFIRE data
VHClass1	Binary	Vegetation height reclassified into two categories (0 = unobstructed, 1 = obstructed > 0.5 m)
Lion_RP	Numerical	Mountain lion risky places model, unweighted by population utilization distribution
Lion_RSF	Numerical	Mountain lion habitat selection function, unweighted by population utilization distribution
Lion_PopUD	Numerical	Mountain lion population utilization distribution for all years combined
LionYearlyUD	Numerical	Mountain lion population utilization distribution estimated by year
LionSeasonYearUD	Numerical	Mountain lion population utilization distribution estimated by season and year
LionRP_PopUD	Numerical	Mountain lion risky places model weighted by population utilization distribution for all years
LionRP_YearUD	Numerical	Mountain lion risky places model weighted by population utilization distribution estimated by year
LionRP_SeasonUD	Numerical	Mountain lion risky places model weighted by population utilization distribution estimated by season and year
LionRSF_PopUD	Numerical	Mountain lion habitat selection function weighted by population utilization distribution for all years
LionRSF_YearUD	Numerical	Mountain lion habitat selection function weighted by population utilization distribution estimated by year
LionRSF_SeasonUD	Numerical	Mountain lion habitat selection function weighted by population utilization distribution estimated by season and year
Wolf_RP	Numerical	Mexican wolf risky places model, unweighted by utilization distributions
Wolf_RSF	Numerical	Mexican wolf habitat selection function, unweighted by utilization distributions
WolfPopG	Numerical	Mexican wolf population utilization distribution for all years combined
WolfPackMaxG	Numerical	Mexican wolf pack utilization distributions weighted by maximum annual pack size for all years combined
WolfPackAveG	Numerical	Mexican wolf pack utilization distributions weighted by average annual pack size for all years combined
WolfPopY	Numerical	Mexican wolf population utilization distribution estimated by year
WolfPackMaxY	Numerical	Mexican wolf pack utilization distributions estimated by year and weighted by maximum annual pack size
WolfPackAveY	Numerical	Mexican wolf pack utilization distributions estimated by year and weighted by average annual pack size
WolfPopSY	Numerical	Mexican wolf population utilization distribution estimated by season and year
WolfPackAveSY	Numerical	Mexican wolf pack utilization distributions estimated by season and year and weighted by average seasonal pack size
WolfRP_PopG	Numerical	Mexican wolf risky places model weighted by population utilization distribution for all years combined
WolfRP_PackMaxG	Numerical	Mexican wolf risky places model weighted by pack utilization distributions for all years combined and maximum annual pack size
WolfRP_PackAveG	Numerical	Mexican wolf risky places model weighted by pack utilization distributions for all years combined and average annual pack size
WolfRP_PopY	Numerical	Mexican wolf risky places model weighted by population utilization distribution estimated by year
WolfRP_PackMaxY	Numerical	Mexican wolf risky places model weighted by pack utilization distributions estimated by year and maximum annual pack size
WolfRP_PackAveY	Numerical	Mexican wolf risky places model weighted by pack utilization distributions estimated by year and average annual pack size
WolfRP_PopSY	Numerical	Mexican wolf risky places model weighted by population utilization distribution estimated by season and year
WolfRP_PackAveSY	Numerical	Mexican wolf risky places model weighted by pack utilization distributions estimated by season/year and average annual pack size
WolfRSF_PopG	Numerical	Mexican wolf habitat selection function weighted by population utilization distribution for all years combined
WolfRSF_PackMaxG	Numerical	Mexican wolf habitat selection function weighted by pack utilization distributions for all years combined and maximum annual pack size
WolfRSF_PackAveG	Numerical	Mexican wolf habitat selection function weighted by pack utilization distributions for all years combined and average annual pack size
WolfRSF_PopY	Numerical	Mexican wolf habitat selection function weighted by population utilization distribution estimated by year
WolfRSF_PackMaxY	Numerical	Mexican wolf habitat selection function weighted by pack utilization distributions estimated by year and maximum annual pack size
WolfRSF_PackAveY	Numerical	Mexican wolf habitat selection function weighted by pack utilization distributions estimated by year and average annual pack size
WolfRSF_PopSY	Numerical	Mexican wolf habitat selection function weighted by population utilization distribution estimated by season and year
WolfRSF_PackAveSY	Numerical	Mexican wolf habitat selection function weighted by pack utilization distributions estimated by season/year and average annual pack size
Closest.dist24	Numerical	Minimum distance in meters from the observed elk herd to the nearest collared Mexican wolf in the prior 24 hours
Closewolf24	Binary	Whether a collared Mexican wolf was within three kilometers of the observed elk herd in the prior 24 hours (1) or not (0)
WolfActivity	Numerical	Mexican wolf hourly activity rate at the time of the observation estimated from seasonal wolf activity models

Additional variables in ISSCovars.csv:

Column header	Data type	Description
Av_PropForage	Numerical	Proportion of herd actively foraging, averaged across scan samples
Av_PropVigilant	Numerical	Proportion of herd intensely vigilant, averaged across scan samples
Av_PropMulti	Numerical	Proportion of herd multitasking, averaged across scan samples
Av_PropTravel	Numerical	Proportion of herd traveling between forage patches, averaged across scan samples
Av_PropBed	Numerical	Proportion of herd bedded, averaged across scan samples
Av_PropOther	Numerical	Proportion of herd engaged in other behaviors, averaged across scan samples
All_PropForage	Numerical	Proportion of herd actively foraging, from sum of scan samples
All_PropVigilant	Numerical	Proportion of herd intensely vigilnat, from sum of scan samples
All_PropMulti	Numerical	Proportion of herd multitasking, from sum of scan samples
All_PropTravel	Numerical	Proportion of herd traveling between forage patches, from sum of scan samples
All_PropBed	Numerical	Proportion of herd bedded, from sum of scan samples
All_PropOther	Numerical	Proportion of herd engaged in other behaviors, from sum of scan samples
n_Scans	Numerical	Number of instantaneous scan samples collected per scan observation
Scan_Size	Numerical	Number of elk recorded during instantaneous scan sampling (does not include calves < 1 year)
ISS_Diel	Categorical	Diel period during which scan sampling began

Variable descriptions and structure in Focal_repeats.csv:

Column header	Data type	Description
HerdKey	Categorical	Identifier for unique elk herds
FocalKey	Categorical	Identifier for unique focal elk within herd
Behavior_Steps	Numerical	Total steps focal elk had taken at end of behavior
Behavior	Categorical	Behavior of focal elk: GRAZ, BRWS, or FORA = actively foraging; MLTC or NMLT = multitasking; VIGL or NVIG = intensely vigilant; TRVL = traveling between forage patches; OTHR, MLTR, BVIG, BRUM, NURS, BEDD = other behaviors; END = end of focal observation
Behavior_Time_Start	Time	Time at which focal elk initiated a behavior, in %M:%S
Behavior_Time_End	Time	Time at which focal elk began a new behavior, in %M:%S
Behavior_Duration	Numerical	Duration of behaviors in seconds

Dynamic risk from Mexican wolves and mountain lions influences elk foraging behavior

Data files

Abstract

README: Dynamic risk from Mexican wolves and mountain lions influences elk foraging behavior