DOI: 10.5061/dryad.5mkkwh7hw
title: Data from: Startling ravens (Corvus corax) at foraging: differences in anti-predator behaviour can be explained by age rather than personality

description: Background: This study investigated boldness-shyness as a personality
 trait in free-flying ravens. Through the utilisation of audio playbacks of
 aerial or ground-dwelling predators, designed to elicit a startle response
 in the birds, we sought to elucidate the influence of between-individual
 behavioural differences, stimulus type, age and sex on ravens’ startle
 responses. Analysis: We analysed nine boldness-shyness related behaviour
 variables for (1) consistent between-individual differences by using the R
 functions ICC and rptr, (2) for influences of stimulus type, age and sex
 through linear mixed models (LMMs) and bootstrap-coupled estimation plots
 and (3) possiblem habituation effects by calculating the delta between
 each trial a bird participated in and afterwards entering these deltas
 into LMMs as well. Results: Our findings do not corroborate
 boldness-shyness as a consistent personality trait in free- flying ravens,
 however, they indicate that the manifestation of boldness in this species
 could be shaped by age and type of stimulus.
description: # Data from: Startling ravens (Corvus corax) at foraging: differences in
 anti-predator behaviour can be explained by age rather than personality
 Dataset DOI: [10.5061/dryad.5mkkwh7hw](10.5061/dryad.5mkkwh7hw) ##
 Description of the data and file structure Analysis: We analysed nine
 boldness-shyness related behaviour variables for (1) consistent
 between-individual differences by using the R functions ICC and rptr, (2)
 for influences of stimulus type, age and sex through linear mixed models
 (LMMs) and bootstrap-coupled estimation plots, and (3) possible
 habituation effects by calculating the delta between each trial a bird
 participated in and afterwards entering these deltas into LMMs as well.
 These variables were: Feeding latency, Return latency, Pre-stimulus
 arrival latency, Frequency of anti-predator behaviours, Time spent
 feeding, Time spent watching, Duration of presence in total, Duration of
 presence after stimulus and Locomotion duration. Dataset contents: The
 data contains all raw data measured during the conduction of our playback
 experiment and behavioural coding of the resulting video footage (Dataset
 S1). It also contains curated datasets extracted from this raw data, which
 was used in the statistical analyses (Dataset S2, Dataset S3, ICC files
 with extracted data of only the first two individual trials per individual
 for each behaviour variable and measured data organized to run the
 bootstrap analysis in R). ### Files and variables Note: All .csv files are
 semicolon-delimited with a comma for a decimal. #### File: Dataset_S3.csv
 **Description:** Data used to calculate habituation effects. Cells
 annotated as "n/a" indicate missing data (i.e. the individual
 did not participate in this trial). ##### Variables * Individual_Trial:
 Reports the order of trials from an individual view, as none of the 12
 individuals joined all 10 trials. I.e., The 3rd trial that was conducted
 within the scope of this study might have been the first trial in which
 individual "Gluco" partook in. Therefore it would be marked with
 "Trial=3", but "Individual_Trial=1". * Focal_ID: Name
 of the focal bird. * Feeding_Latency: Latency until the focal bird resumed
 feeding on site after the playback [sec]. * Delta_Feeding_Latency:
 Difference in "Feeding_Latency" between two consecutive
 individual trials [sec]. * Return_Latency: Latency until the focal bird
 returned to the feeding site after flying away/ on a tree after hearing
 the startle playback [sec]. * Delta_Return_Latency: Difference in
 "Return_Latency" between two consecutive individual trials
 [sec]. * Arrival_Latency: Latency from the start of the trial until the
 focal bird landed in the experimental area for the first time [sec]. *
 Delta_Arrival_Latency: Difference in "Arrival_Latency" between
 two consecutive individual trials [sec]. * Events_anti-predator_corrected:
 Sum of all scored antipredator behaviours [counts], divided by the total
 video length. * Delta_Events_anti-predator_corrected: Difference in
 "Events_anti-predator_corrected" between two consecutive
 individual trials [counts/sec]. * Duration_Feeding[sec]_corrected: Sum of
 all scored manipulate/eat food [sec] and pick/dig/walk and search [sec]
 instances, divided by the total video length. *
 Delta_Duration_Feeding[sec]_corrected: Difference in
 "Duration_Feeding[sec]_corrected" between two consecutive
 individual trials [sec]. * Duration_watching[per_trail_sec]_corrected: Sum
 of all scored "wait and watch" instances [sec], divided by the
 total video length. * Delta_Duration_watching[per_trail_sec]_corrected:
 Difference in "Duration_watching[per_trail_sec]_corrected"
 between two consecutive individual trials [sec]. *
 Duration_present_total_corrected: Trial duration [sec] minus "out of
 view" events and "Arrival_Latency" for the focal bird until
 the end of the trial or until "take off" and "move
 away" without returning occurred, divided by the total video length.
 * Delta_Duration_present_total_corrected: Difference in
 "Duration_present_total_corrected" between two consecutive
 individual trials [sec]. * Duration_locomotion[per_trail_sec]_corrected:
 Sum of the durations of all instances of Locomotion behaviour for the
 focal bird during this trial [sec], divided by the total video length. *
 Delta_Duration_locomotion[per_trail_sec]_corrected: Difference in
 "Duration_locomotion[per_trail_sec]_corrected" between two
 consecutive individual trials [sec]. *
 Duration_present_afterStimulus_corrected: Sum of the durations of all
 scored behaviours [sec] - excluding "out of view" events - after
 "Stimulus_start" and until the end of the trial or until
 "take off" and "move away" without returning occurred,
 divided by the total video length. *
 Delta_Duration_present_afterStimulus_corrected:Difference in
 "Duration_present_afterStimulus_corrected" between two
 consecutive individual trials [sec]. #### File: Dataset_S2.csv
 **Description:** Raw data ##### Variables * Trial: Reports the order in
 which the trials were conducted; ranges from 1-10. * Individual_Trial:
 Reports the order of trials from an individual view, as none of the 12
 individuals joined all 10 trials. I.e., The 3rd trial that was conducted
 within the scope of this study might have been the first trial in which
 individual "Gluco" partook in. Therefore it would be marked with
 "Trial=3", but "Individual_Trial=1". * Year: Reports
 the year in which the trial was conducted. * Date: Reports the full date
 on which a trial was conducted in the format: DD/MM/YY. * Starting_time:
 Time of day the experiment started. Note: Starting time for trial 1 was
 unfortunately not noted down during conduction of the test and is
 therefore annotated as "n/a". * Stimulus: Information about the
 exact stimulus that was used in a trial. * Stimulus_type: Information on
 whether the stimulus used was natural (=1) or man-made (=2). *
 Speake_position: Reports whether the speaker was attached to a fence pole
 either ~1.5 m (=1) or ~3 m (=2) away from the camera. * Stimulus_start:
 Information about how much time passed between the start of the experiment
 until the startle noise was played [min, sec]. * Weather: If the weather
 was 1=sun, 2=cloudy/foggy, 3=rainy, 4=heavily raining, 5=little snowing,
 6=heavily snowing. * Temp: Temperature [°C] on the day a test was
 conducted. * Food: Type of food that was provided during the trial. *
 Phase: 1=Pre-playback, 2=During Playback (30 sec for raptor playbacks, 20
 sec for gunshots), 3=Post-playback * Birth_year: Year in which the focal
 bird hatched. * Age: Age [years] of the focal bird during experiment
 conduction. * Age_class: If the focal bird was Juvenile, a Subadult or an
 Adult during experiment conduction. * Age_ordered: 1= this year's
 hatchling, 2= previous year's hatchling, 3= subadult, 4= adult * Sex:
 Sex of the focal bird. * Focal_ID: Name of the focal bird. * min:
 Timepoint at which the scored behaviour started in the video. * D/R:
 Whether to focal bird was donor or receiver of the current behaviour (not
 applicable to all scored behaviours). Instances in which the interaction
 partner was not scored due to invisibility of the outcome or because the
 specific behaviour does not involve an interaction partner, are annotated
 as "n/a". * Behav_Category: Which category the scored behaviour
 was assigned to; 0=Self-maintenance, 1=Anti-predator, 2=Foraging, 3=Object
 manipulation, 5=Affiliative, 9=Out of view, 10=Locomotion * Behav_Type:
 Which behaviour from a specific behaviour catergory was scored;
 Self-maintenance 1=resting 2=auto-preening 3=scratching/stretching/shaking
 4=beak wipe 5=panting 10=other Anti-predator 1=alert (upright and thin)
 2=freeze/crowch 3=scold - vocalize 5=jumping jack 6=flee/retreat 10=other
 Foraging 1=manipulate/eat food 2=transport/cache/retrieve 3=pick/dig/walk
 and search 6=steal food: attempt + success 8=wait and watch 10=other
 Object manipulation 1=manipulate/hold/carry object Affiliative 1=cont sit
 (body length) 2=allo-preen 3=touch/hold 4=approach/follow 5=vocalize -
 soft call 10=other Out of view 1=move away 2=re-locate 10=other Locomotion
 1=walk/run/bounce/jump 2=take off 3=landing 4=flying * Description: Which
 behaviour was scored. * Interactor\_ ID: Identity of the interaction
 partner (if visible and if applicable). If the scored behaviour did not
 involve an interacting partner, cells are annotated as "n/a".
 "nmr" stands for "non-marked raven"; "mr"
 stands for "marked raven" (which could not be identified). *
 dur: Duration of the scored behaviour [sec]. Cells annotated as
 "n/a" indicate behaviours which were not scored for their
 duration, but for frequency only. * Latency1: Latency until the focal bird
 resumed feeding on site after the playback [sec], i.e., Feeding latency;
 only calculated once per individual and per trial. * Latency2: Latency
 until the focal bird returned to the feeding site after flying away/ on a
 tree after hearing the startle playback [sec]., i.e., Return latency; only
 calculated once per individual and per trial. * Success: Reports whether
 the scored behaviour was successful or not; only applicable for
 "steal food"; 0=no, 1=yes. Cells annotated as "n/a"
 indicate behaviours which were not scored for success. * Frequency: How
 often the scored behaviour was repeated (if applicable, i.e., amount of
 pecks during "pick/dig/walk and search"). Cells annotated as
 "n/a" indicate behaviours, which were not scored for frequency.
 * position: Location of the focal bird within the test area (only
 recordedduring location changes; not continuously). *
 sum_antipredator_freq: Sum of all scored antipredator behaviours [counts];
 calculated once per individual and per individual trial. This data is
 sorted in the column "Events_anti-predator" in Dataset S1.csv.,
 which was used for the analysis in the article. *
 sum_foraging_type1+3_dur: Sum of all scored manipulate/eat food [sec] and
 pick/dig/walk and search [sec] instances; calculated once per individual
 and per individual trial. This data is sorted in the column
 "Duration_feeding" in Dataset S1.csv., which was used for the
 analysis in the article. * sum_foraging_type8_dur: Sum of all scored
 "wait and watch" instances [sec]; calculated once per individual
 and per individual trial. This data is sorted in the column
 "Duration_watching" in Dataset S1.csv., which was used for the
 analysis in the article. #### File: Dataset_S1.csv
 **Description:** Curated dataset used for the personality analyses and the
 LMMs for stimulus type, age and sex effects. ##### Variables * Trial:
 Reports the order in which the trials were conducted; ranges from 1-10. *
 Individual_Trial: Reports the order of trials from an individual view, as
 none of the 12 individuals joined all 10 trials. I.e., The 3rd trial that
 was conducted within the scope of this study might have been the first
 trial in which individual "Gluco" partook in. Therefore it would
 be marked with "Trial=3", but "Individual_Trial=1". *
 Year: Reports the year in which the trial was conducted. * Date: Reports
 the full date on which a trial was conducted in the format: DD/MM/YY. *
 Starting_time: Time of day the experiment started. Note: Starting time for
 trial 1 was unfortunately not noted down during conduction of the test and
 is therefore annotated as "n/a". * Stimulus: Information about
 the exact stimulus that was used in a trial. * Stimulus_type: Information
 on whether the stimulus used was natural (=1) or man-made (=2). *
 Speaker_position: Reports whether the speaker was attached to a fence pole
 either ~1.5 m (=1) or ~3 m (=2) away from the camera. * Stimulus_start:
 Information about how much time passed between the start of the experiment
 until the startle noise was played [min, sec]. * Video_length [sec]: Total
 duration of the trial. * Weather: If the weather during testing was 1=sun,
 2=cloudy/foggy, 3=rainy, 4=heavily raining, 5=little snowing, 6=heavily
 snowing. * Temp: Temperature [°C] on the day the test was conducted. *
 Food: Type of food that was provided during the trial. * Phase:
 1=Pre-playback, 2=During Playback (30 sec for raptor playbacks, 20 sec for
 gunshots), 3=Post-playback * Birth_year: Year in which the focal bird
 hatched. * Age: Age [years] of the focal bird during experiment
 conduction. * Age_class: If the focal bird was Juvenile, a Subadult or an
 Adult during experiment conduction. * Age_ordered: 1= this year's
 hatchling, 2= previous year's hatchling, 3= subadult, 4= adult * Sex:
 Sex of the focal bird. * Focal_ID: Name of the focal bird. * min:
 Timepoint at which the scored behaviour started in the video. *
 Behav_Category: Which category the scored behaviour was assigned to;
 0=Self-maintenance, 1=Anti-predator, 2=Foraging, 3=Object manipulation,
 5=Affiliative, 9=Out of view, 10=Locomotion. * Behav_Type: Which behaviour
 from a specific behaviour catergory was scored; Self-maintenance 1=resting
 2=auto-preening 3=scratching/stretching/shaking 4=beak wipe 5=panting
 10=other Anti-predator 1=alert (upright and thin) 2=freeze/crowch 3=scold
 - vocalize 5=jumping jack 6=flee/retreat 10=other Foraging
 1=manipulate/eat food 2=transport/cache/retrieve 3=pick/dig/walk and
 search 6=steal food: attempt + success 8=wait and watch 10=other Object
 manipulation 1=manipulate/hold/carry object Affiliative 1=cont sit (body
 length) 2=allo-preen 3=touch/hold 4=approach/follow 5=vocalize - soft call
 10=other Out of view 1=move away 2=re-locate 10=other Locomotion
 1=walk/run/bounce/jump 2=take off 3=landing 4=flying * Description: Which
 behaviour was scored. * dur: Duration of the scored behaviour [sec]. Cells
 annotated as "n/a" indicate behaviours which were not scored for
 their duration, but for frequency only. * Feeding_Latency[sec]: Latency
 until the focal bird resumed feeding on site after the playback. *
 Return_Latency[sec]: Latency until the focal bird returned to the feeding
 site after flying away/ on a tree after hearing the startle playback
 [sec]. * Arrival_Latency[sec]: Latency from the start of the trial until
 the focal bird landed in the experimental area for the first time. *
 Frequency: How often the focal bird performed "pecks" during the
 scored behaviour. "n/a" indicates instances of missing data. *
 Events_anti-predator: Sum of all scored antipredator behaviours [counts].
 * Events_anti-predator_corrected: Sum of all scored antipredator
 behaviours [counts], divided by the total video length. *
 Duration_feeding: Sum of all scored manipulate/eat food [sec] and
 pick/dig/walk and search [sec] instances. *
 Duration_Feeding[sec]_corrected: Sum of all scored manipulate/eat food
 [sec] and pick/dig/walk and search [sec] instances, divided by the total
 video length. * Duration_watching: Sum of all scored "wait and
 watch" instances [sec]. * Duration_watching[per_trail_sec]_corrected:
 Sum of all scored "wait and watch" instances [sec], divided by
 the total video length. * Duration_present_total: Trial duration [sec]
 minus "out of view" events and "Arrival_Latency" for
 the focal bird until the end of the trial or until "take off"
 and "move away" without returning occurred. *
 Duration_present_total_corrected: Trial duration [sec] minus "out of
 view" events and "Arrival_Latency" for the focal bird until
 the end of the trial or until "take off" and "move
 away" without returning occurred, divided by the total video length.
 * Duration_locomotion: Sum of the durations of all instances of Locomotion
 behaviour for the focal bird during this trial [sec]. *
 Duration_locomotion[per_trail_sec]_corrected: Sum of the durations of all
 instances of Locomotion behaviour for the focal bird during this trial
 [sec], divided by the total video length. *
 Duration_present_afterStimulus: Sum of the durations of all scored
 behaviours [sec] - excluding "out of view" events - after
 "Stimulus_start" and until the end of the trial or until
 "take off" and "move away" without returning occurred.
 * Duration_present_afterStimulus_corrected: Sum of the durations of all
 scored behaviours [sec] - excluding "out of view" events - after
 "Stimulus_start" and until the end of the trial or until
 "take off" and "move away" without returning occurred,
 divided by the total video length. #### File: ICC_arrival.csv
 **Description:** Data used for ICC calculations. It was extracted from the
 dataset S1 and only contains two columns with 12 rows. The first column is
 the measured value of the first trial an individual participated in;
 Second column contains the value (e.g., Feeding latency) of their second
 trial. Values are ordered after bird ID (i.e. Bird 1' latency is
 always in row 1; Bird 2 in row 2 etc.). ##### Variables * Trial1: Latency
 from the start of the trial until the focal bird landed in the
 experimental area for the first time [sec]; scored in the first individual
 trial. * Trial2: Latency from the start of the trial until the focal bird
 landed in the experimental area for the first time [sec]; scored in the
 second individual trial. #### File: ICC_counts_antipred_corrected.csv
 **Description:** Data used for ICC calculations. It was extracted from the
 dataset S1 and only contains two columns with 12 rows. The first column is
 the measured value of the first trial an individual participated in;
 Second column contains the value (e.g., Feeding latency) of their second
 trial. Values are ordered after bird ID (i.e. Bird 1' latency is
 always in row 1; Bird 2 in row 2 etc.). ##### Variables * Trial1: Sum of
 all scored antipredator behaviours [counts], divided by the total video
 length; scored in the first individual trial. * Trial2: Sum of all scored
 antipredator behaviours [counts], divided by the total video length;
 scored in the second individual trial. #### File:
 ICC_dur_watching_corrected.csv **Description:** Data used for ICC
 calculations. It was extracted from the dataset S1 and only contains two
 columns with 12 rows. The first column is the measured value of the first
 trial an individual participated in; Second column contains the value
 (e.g., Feeding latency) of their second trial. Values are ordered after
 bird ID (i.e. Bird 1' latency is always in row 1; Bird 2 in row 2
 etc.). ##### Variables * Trial1: Sum of all scored "wait and
 watch" instances [sec], divided by the total video length; scored in
 the first individual trial. * Trial2: Sum of all scored "wait and
 watch" instances [sec], divided by the total video length; scored in
 the second individual trial. #### File: ICC_feeding_corrected.csv
 **Description:** Data used for ICC calculations. It was extracted from the
 dataset S1 and only contains two columns with 12 rows. The first column is
 the measured value of the first trial an individual participated in;
 Second column contains the value (e.g., Feeding latency) of their second
 trial. Values are ordered after bird ID (i.e. Bird 1' latency is
 always in row 1; Bird 2 in row 2 etc.). ##### Variables * Trial1: Sum of
 all scored manipulate/eat food [sec] and pick/dig/walk and search [sec]
 instances, divided by the total video length; scored in the first
 individual trial. * Trial2: Sum of all scored manipulate/eat food [sec]
 and pick/dig/walk and search [sec] instances, divided by the total video
 length; scored in the second individual trial. #### File:
 ICC_feeding_latency.csv **Description:** Data used for ICC calculations.
 It was extracted from the dataset S1 and only contains two columns with 12
 rows. The first column is the measured value of the first trial an
 individual participated in; Second column contains the value (e.g.,
 Feeding latency) of their second trial. Values are ordered after bird ID
 (i.e. Bird 1' latency is always in row 1; Bird 2 in row 2 etc.).
 ##### Variables * Trial1: Latency until the focal bird resumed feeding on
 site after the playback [sec]; scored in the first individual trial. *
 Trial2: Latency until the focal bird resumed feeding on site after the
 playback [sec]; scored in the second individual trial. #### File:
 ICC_presence_after_stimulus_corrected.csv **Description:** Data used for
 ICC calculations. It was extracted from the dataset S1 and only contains
 two columns with 12 rows. The first column is the measured value of the
 first trial an individual participated in; Second column contains the
 value (e.g., Feeding latency) of their second trial. Values are ordered
 after bird ID (i.e. Bird 1' latency is always in row 1; Bird 2 in row
 2 etc.). ##### Variables * Trial1: Sum of the durations of all scored
 behaviours [sec] - excluding "out of view" events - after
 "Stimulus_start" and until the end of the trial or until
 "take off" and "move away" without returning occurred,
 divided by the total video length; scored in the the first individual
 trial. * Trial2: Sum of the durations of all scored behaviours [sec] -
 excluding "out of view" events - after
 "Stimulus_start" and until the end of the trial or until
 "take off" and "move away" without returning occurred,
 divided by the total video length; scored in the the second individual
 trial. #### File: ICC_locomotion_corrected.csv **Description:** Data used
 to calculate ICC (extracted from Dataset S2) ##### Variables * Trial1: Sum
 of the durations of all instances of Locomotion behaviour [sec], divided
 by the total video length; scored in the first individual trial. * Trial2:
 Sum of the durations of all instances of Locomotion behaviour [sec],
 divided by the total video length; scored in the second individual trial.
 #### File: ICC_presence_corrected.csv **Description:** Data used for ICC
 calculations. It was extracted from the dataset S1 and only contains two
 columns with 12 rows. The first column is the measured value of the first
 trial an individual participated in; Second column contains the value
 (e.g., Feeding latency) of their second trial. Values are ordered after
 bird ID (i.e. Bird 1' latency is always in row 1; Bird 2 in row 2
 etc.). ##### Variables * Trial1: Trial duration [sec] minus "out of
 view" events and "Arrival_Latency" for the focal bird until
 the end of the trial or until "take off" and "move
 away" without returning occurred, divided by the total video length;
 scored in the the first individual trial. * Trial2: rial duration [sec]
 minus "out of view" events and "Arrival_Latency" for
 the focal bird until the end of the trial or until "take off"
 and "move away" without returning occurred, divided by the total
 video length; scored in the second individual trial. #### File:
 ICC_return_latency.csv **Description:** Data used for ICC calculations. It
 was extracted from the dataset S1 and only contains two columns with 12
 rows. The first column is the measured value of the first trial an
 individual participated in; Second column contains the value (e.g.,
 Feeding latency) of their second trial. Values are ordered after bird ID
 (i.e. Bird 1' latency is always in row 1; Bird 2 in row 2 etc.).
 ##### Variables * Trial1: Latency until the focal bird returned to the
 feeding site after flying away/ on a tree after hearing the startle
 playback [sec]; scored in the first individual trial. * Trial2: Latency
 until the focal bird returned to the feeding site after flying away/ on a
 tree after hearing the startle playback [sec]; scored in the the second
 individual trial. #### File: Bootstrap_antipredator.csv
 **Description:** Data used to calculate bootstrap-coupled estimation plots
 (extracted from dataset S1). ##### Variables * Stimulus type: Reports if
 the stimulus was natural (=Bird of prey calls) or man-made (Gunshots). *
 Frequency [counts/sec]: Sum of all scored antipredator behaviours
 [counts], divided by the total video length. #### File:
 Bootstrap_feeding_latency.csv **Description:** Data used to calculate
 bootstrap-coupled estimation plots (extracted from dataset S1). #####
 Variables * Age: Reports if the focal bird belongs to the
 "Juveniles" or "Subdadults and adults" age group. *
 Latency: Latency until the focal bird resumed feeding on site after the
 playback [sec]. #### File: Text_S1.txt **Description:**  Full R script
 used for the analysis of all results within the paper. #### ##
 Code/software The full code can be found in the file "Text S1".
 During the analyses for this paper, it was run in the statistical software
 R version 4.2.2 (R Core Team, 2019) within RStudio environment (version
 R-3.6.1).