DOI: 10.5061/dryad.sn02v6xfn
title: Does differential habitat selection facilitate coexistence between badgers and hedgehogs?

description: Predicting the spatial and temporal responses of species exhibiting
 intra-guild predation (IGP) relationships is difficult due to variation in
 potential interactions and environmental context. Eurasian badgers (Meles
 meles) are intra-guild predators of European hedgehogs (Erinaceus
 europaeus) and are implicated in their population decline via both direct
 predation and competition for shared food resources. Previous studies have
 shown spatial separation between these species and attributed this to
 hedgehogs experiencing a ‘landscape of fear’, but little is known about
 the potential role of differential habitat use. We estimated the density
 and occupancy of both species at 22 rural study sites in England and
 Wales, to explore whether food availability, habitat, or the presence of
 badgers, explained hedgehog distributions. Hedgehog density varied
 significantly across major rural land uses, whereas badger density did
 not. Although both species co-existed at a regional (1 km²) scale,
 occupancy modelling showed spatial segregation at a finer (individual
 camera trap) scale, associated with differential habitat use. In contrast
 to badgers, hedgehogs were recorded near buildings, and in areas
 supporting lower invertebrate biomass. This is in agreement with IGP
 theory, whereby IG-prey may occupy sub-optimal habitat to avoid predation,
 however hedgehog habitat use did not vary relative to the presence of
 badgers. Badger and hedgehog temporal activity showed no evidence of
 separation. Although these findings are consistent with hedgehogs avoiding
 badgers via a landscape of fear, they are also indicative of differential
 habitat use, highlighting the need for more holistic studies considering
 variation in habitat selection and food availability when investigating
 intra-guild relationships. Future studies exploring alternative hypotheses
 for urban habitat selection by hedgehogs are needed to better understand
 how possible spatial niche partitioning may support their coexistence with
 badgers in some areas.
description: We deployed 645 camera traps across the 22 sites (mean = 29 per
 site), stratified by five key habitat categories: arable, amenity
 grassland (including residential gardens), agricultural grassland, built
 environment (buildings and hardstanding) and woodland (deciduous and
 coniferous). Twenty two rural sites (0.4 – 1.3 km²) were surveyed for
 badgers and hedgehogs between April - September in 2018 or 2019 (Figure
 1). Camera trapping was used to estimate focal species occupancy and
 density, and invertebrate sampling was used to obtain an index of prey
 availability for both species. Rural study sites comprising either
 arable-dominated, pasture-dominated (where a single habitat represented
 ≥66.7% of the total area of a site), or mixed farming habitat were
 selected (Table 1), as these were considered landscapes most likely to be
 occupied by badgers and hedgehogs (Judge et al., 2014; Williams et al.,
 2018).
description: # Does differential habitat selection facilitate coexistence between
 badgers and hedgehogs?
 [https://doi.org/10.5061/dryad.sn02v6xfn](https://doi.org/10.5061/dryad.sn02v6xfn) ## Description of the data and file structure Data relating to camera and invertebrate surveys that were conducted to investigate badger and hedgehog intraguild predation between April 2018 and September 2019. Data and Scripts are stored in the zipped file 'Data_and_scripts_repository' ### Files and variables #### File: Data\_and\_scripts\_repository.zip A description of the folder structure and file contents is given below. ##### Activity: main folder containing data files and R scripts required to reproduce activity analysis for both study species; badger and hedgehog. * badger_small.png – silhouette image of a badger used to produce figure 5. - hog_badger_activity.csv – detection data after conversion to solar time. * hog_small.png – silhouette image of a hedgehog used to produce figure 5. - Rplot.png – copy of ‘Figure 5’ showing diel activity pattern density curves of badger and hedgehog as obtained from camera trap detections. Rplot can be reproduced by running the ‘Temporal_activity_script’. * Temporal_activity.csv – raw detection data captured by camera trapping. - Temporal_activity_script.R – R script required to reproduce activity analysis. ##### Occupancy: main folder containing presence/absence data and R scripts required to reproduce occupancy analysis for both study species; badger and hedgehog. * BADGER.csv – Presence/absence data for badger at each individual camera trap location included in the study. Columns S01 – S10 represent the first 10 nights of camera trapping at each individual camera trap location. A value of ‘1’ depicts the species was detected and a value of ‘0’ depicts the species was not detected. - COVARIATES.csv – Site information corresponding to the location of each individual camera trap deployed during the study. Pitfall trapping was conducted to obtain measures of macroinvertebrate abundance (PITFALL_ABUNDANCE_Z) and biomass (PITFALL_BIOMASS_Z). Relative earthworm abundance (WORM_ABUNDANCE_Z) and biomass (WORM_BIOMASS_Z) was estimated by sampling soil cores (further detail is provided in the manuscript). * HEDGEHOG.csv – Presence/absence data for hedgehog at each individual camera trap location included in the study. Columns S01 – S10 represent the first 10 nights of camera trapping at each individual camera trap location. A value of ‘1’ depicts the species was detected and a value of ‘0’ depicts the species was not detected. - Occupancy_script.R – R script required to reproduce single species, and multi-species occupancy models conducted as part of this study. ##### REM: main folder containing detection data (captured by camera trapping), site information and R scripts required to produce density estimates for badger and hedgehog at each study site. Density estimates were calculated from camera trapping data using the Random Encounter Model (REM). * Camera_deployment.xlsx – Deployment and collection data for individual camera locations at each study site. - Detection.xlsx – Detection data for badger and hedgehog across all study sites. Cells containing ‘n/a’ indicate data not available. * Land_use_%.xlsx – Proportion of major habitat types within the survey area of each study site. - REM_estimates.xlsx – Table of REM density estimates for badger and hedgehog at each study site. ###### Activity\_global *(sub-folder of REM)* – contains files for producing ‘global activity’ estimates for each species. ‘Global activity’ was used to calculate density estimates at sites with low detection rates (&lt;10 detections of a species). * Badger_global_activity.csv - Badger_global_activity.R * Hedgehog_global_activity.csv - Hedgehog_global_activity.R ###### Exemplar\_REM (sub-folder of REM) – provides an example for calculating REM density estimates. Site 1 (Brackenhurst 2018) is included as an exemplar. *Density estimates can be reproduced for other individual sites/species using data contained in 'Detection.xlsx' in the main folder ‘REM’.* * Site_1_*badger (sub-folder of Exemplar*_REM) – contains the detection data for badger at Site 1 (Brackenhurst 2018), R scripts for calculating ‘activity’ at the site level (a parameter required for REM analysis) and ‘density script’ for calculating the density estimate for badger at Site 1 using REM. ·        Activity_script.R ·        Brackenhurst_18.xlsx ·        Brackenhurst_18_activity.csv ·        Brackenhurst_18_percam.csv ·        Density_script.R * Site_1_hedgehog *(sub-folder of Exemplar*_REM) – contains the detection data for hedgehog at Site 1 (Brackenhurst 2018), R scripts for calculating ‘activity’ at the site level (a parameter required for REM analysis) and ‘density script’ for calculating the density estimate for hedgehog at Site 1 using REM. ·        Activity_script.R ·        Brackenhurst_18.xlsx ·        Brackenhurst_18_activity.csv ·        Brackenhurst_18_percam.csv   ·        Density_script.R ###### REM STATS (sub-folder of REM) – contains data files and R scripts required to reproduce all statistical analysis conducted as part of this study. * Chi-square.R - Chitest.csv * IGP_Density_plot.tiff – Rplot showing the relationship between badger and hedgehog densities across all study sites. - Land-use.csv * Land-use_levene.csv - Landuse.R * Linear_regression.csv - Linear_regression.R ## Code/software Scripts were run using R Studio and packages available through the CRAN R repository.