Sex-specific consequences of juvenile dispersal for survival and reproduction in an island lizard

Crain, Caroline 1 ; Wittman, Tyler 1 ; Bhave, Rachana2 1; Seears, Heidi1; Reedy, Aaron1 3; Warner, Daniel3; Cox, Robert 1

Published Sep 05, 2025 on Dryad. https://doi.org/10.5061/dryad.70rxwdc9t

Data files

Sep 05, 2025 version files 1.21 MB

dispersal.data.csv

1.16 MB
README.md

49.85 KB

Abstract

Dispersal is a costly, though potentially rewarding, behavior with important fitness consequences for juveniles. When the costs or benefits of juvenile dispersal differ between sexes, sex-biased dispersal should be favored, though such sex-specific consequences are rarely measured for multiple components of lifetime fitness in the wild. Here, we use detailed mark-recapture data from four annual cohorts of juveniles with associated estimates of juvenile survival and adult reproductive success from genetic parentage to measure natural selection on two dispersal phenotypes (binary dispersal propensity and continuous dispersal distance) in an island population of lizards (Anolis sagrei). Juvenile dispersal was consistently male-biased, with males exhibiting a higher propensity to disperse and dispersing twice as far as females. Males that dispersed had higher survival to adulthood and total fitness than males that remained philopatric, whereas fitness components did not differ between dispersing and philopatric females. Although this result indicates that the fitness benefits of dispersal are sex-specific, we found no difference in fitness components between dispersing and philopatric males after accounting for body size, which was positively correlated with dispersal. Likewise, we did not consistently detect direct selection on dispersal distance in either sex when including body size in multivariate selection analyses, nor did we find consistent correlational selection on combinations of body size and dispersal distance in either sex. We conclude that selection on juvenile dispersal propensity is sex-specific, but likely indirect and mediated through its positive association with body size in males.

Dataset DOI: 10.5061/dryad.70rxwdc9t

Description of the data and file structure

The findings of this paper are drawn from a mark-recapture and genotyping study carried out on a focal island population of brown anole lizards (Anolis sagrei) within the Guana Tolomato Matanzas Natural Estuarine Research Reserve in Florida. Although animals in this population have been tracked between 2011 and 2021 from a long-term ongoing study, this dataset primarily focuses on the individuals in the 2015-2018 cohorts, their survival, reproductive fitness, and locations at various censuses throughout the year. In this study, we evaluated the sex-specific fitness consequences of juvenile dispersal in the wild. We tested for selection on both the propensity to disperse and on the distance dispersed.

Based on the methods, the dataset provided illustrates individual identity of animals (e.g., ID, Sex, Cohort), location of capture information (zone location by age and season), fitness of individuals (S_1_1_spring, survival to adulthood; off_count_year_1, First year reproductive success; tot_off, Lifetime Reproductive success). The mother’s location through her lifetime, as determined by genetic parentage, is also provided. Dispersal variables at various timepoints are displayed as both the zone location dispersed to and from (e.g., j04_11) and as the distances between those zone locations dispersed to and from (eg., j04_11_dist).

Files and variables

File: dispersal.data.csv

Description: The following variables are present in the dataset.

Variables

Variable	Description
ID	Animal ID; Unique identifying ID number given to each captured individual.
Sex	Sex (Categorical variable, M = male, F = female)
Cohort	Hatch year; Assignment based on the year in which the individual was known to be born (e.g., a cohort 2015 individual would have hatched in the year 2015).
Tree_0_3_lsumm_corrected	The zone of capture during an individual's natal year, in the 3rd census (late summer). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
Tree_0_4_fall_corrected	The zone of capture during an individual's natal year, 4th census (fall). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
Tree_1_1_spring_corrected	The zone of capture during an individual's first reproductive year, 1st census (spring). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
Tree_1_2_esumm_corrected	The zone of capture during an individual's first reproductive year, 2nd census (early summer). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
Tree_1_3_lsumm_corrected	The zone of capture during an individual's first reproductive year, 3rd census (late summer). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
Tree_1_4_fall_corrected	The zone of capture during an individual's first reproductive year, 4th census (fall). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
Tree_2_1_spring_corrected	The zone of capture during an individual's second reproductive year, 1st census (spring). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
Tree_2_2_esumm_corrected	The zone of capture during an individual's second reproductive year, 1st census (spring). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
Tree_2_3_lsumm_corrected	The zone of capture during an individual's second reproductive year, 3rd census (late summer). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
Tree_2_4_fall_corrected	The zone of capture during an individual's second reproductive year, 4th census (fall). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
Dam_Tree_0_3_lsumm	Mother's zone of capture in her natal year, 3rd census (late summer). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
Dam_Age	The age (in years) of the mother at the time of offspring hatching. Determined based on the cohort that the mother was born into, as well as her survival data.
S_1_1_spring	Survival to the first reproductive year, first census (spring); binary ( 0 = died; 1= survived)
off_count_year_1	Total number of assigned offspring during an individual's first reproductive year
tot_off	Total number of assigned offspring over an individual's lifetime
Dam_Tree_1_2_esumm_corrected	Mother's zone of capture in her first reproductive year, 2nd census (early summer). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
Dam_Tree_1_3_lsumm_corrected	Mother's zone of capture in her first reproductive year, 3rd census (late summer). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
Dam_Tree_2_2_esumm_corrected	Mother's zone of capture in her second reproductive year, 2nd census (early summer). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
Dam_Tree_2_3_lsumm_corrected	Mother's zone of capture in her second reproductive year, 3rd census (late summer). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
Dam_Tree_3_2_esumm_corrected	Mother's zone of capture in her third reproductive year, 2nd census (early summer). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
Dam_Tree_3_3_lsumm_corrected	Mother's zone of capture in her third reproductive year, 3rd census (late summer). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
Dam_Tree_4_2_esumm_corrected	Mother's zone of capture in her fourth reproductive year, 2nd census (early summer). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
Dam_Tree_4_3_lsumm_corrected	Mother's zone of capture in her fourth reproductive year, 3rd census (late summer). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
j00_03	The presumed hatch zone followed by the zone of capture in an individual's natal year, 3rd census (late summer; e.g., H7H11). The 'presumed hatch zone' is the zone of the mother at the approximate time that the offspring egg was laid. Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
j00_03_corrected	Distance (in meters) between the presumed hatch zone and the individual's zone of capture in their natal year, 3rd census (late summer). NA values indicate cases when dispersal could not be estimated due to a missing zone location.
j00_04	The presumed hatch zone followed by the zone of capture in an individual's natal year, 4th census (fall; e.g., H7H11). The 'presumed hatch zone' is the location of the mother at the approximate time that the offspring egg was laid. Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
j00_04_corrected	Distance (in meters) between the presumed hatch zone and the individual's zone of capture in their natal year, 4th census (fall). NA values indicate cases when dispersal could not be estimated due to a missing zone location.
j00_11	The presumed hatch zone followed by the individual's zone in their 1st reproductive year, 1st census (spring; e.g., H7H11). The 'presumed hatch zone' is the location of the mother at the approximate time that the offspring egg was laid. Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
j00_11_corrected	The distance (in meters) between the presumed hatch zone and the individual's zone of capture in their 1st reproductive year, 1st census (spring). NA values indicate cases when dispersal could not be estimated due to a missing zone location.
mjtrees	The mother's zone at the time that the offspring egg was laid followed by the zone that the individual was first observed (natal year, 3rd census, late summer; e.g., H7H11). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
mjdistance	The distance (in meters) between the mother's zone at the time the offspring egg was laid and the zone that the individual was first observed (natal year, 3rd census, late summer). NA values indicate cases when dispersal could not be estimated due to a missing zone location.
Mass_0_3_lsumm	Measured body weight of individuals (in grams) during their natal year, 3rd census (late summer)
SVL_0_3_lsumm	Measured snout-vent length (mm) during an individual's natal year, 3rd census (late summer)
j03_04	The zone of capture during an individual's natal year, 3rd census (late summer) followed by the zone of capture during an individual's Location natal year, 4th census (fall; e.g., H7H11). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
j04_11	The zone of capture during an individual's natal year, 4th census (fall) followed by the zone of capture during an individual's first reproductive year 1st census (spring; e.g., H7H11). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
j11_12	The zone of capture during an individual's first reproductive year, 1st census (spring) followed by the zone of capture during an individual's first reproductive year, 2nd census (early summer; e.g., H7H11). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
j12_13	The zone of capture during an individual's first reproductive year, 2nd census (early summer) followed by the zone of capture during an individual's first reproductive year, 3rd census (late summer) (e.g., H7H11). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
j13_14	The zone of capture during an individual's first reproductive year, 3rd census (late summer) followed by the zone of capture during an individual's first reproductive year, 4th census (fall; e.g., H7H11). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
j14_21	The zone of capture during an individual's first reproductive year, 4th census (fall) followed by the zone of capture during an individual's second reproductive year, 1st census (spring; e.g., H7H11). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
j21_22	The zone of capture during an individual's second reproductive year, 1st census (spring) followed by the zone of capture during an individual's second reproductive year, 2nd census (early summer; e.g., H7H11). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
j22_23	The zone of capture during an individual's second reproductive year, 2nd census (early summer) followed by the zone of capture during an individual's second reproductive year, 3rd census (late summer; e.g., H7H11). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
j23_24	The zone of capture during an individual's second reproductive year, 3rd census (late summer) followed by the zone of capture during an individual's second reproductive year, 4th census (fall; e.g., H7H11). Unique ID codes for each zone (e.g., H72F) refer to the study island (H), the tree number (1-100), and areas of small shrubs and vegetation (M/Y) or open grass, mud, and sand (F) adjacent to a particular tree. Zone IDs with an underscore character indicate areas located between two numbered trees (e.g., H25_H26). NA values indicate a missing zone location.
j03_04_dist	Distance (in meters) between the zone of capture during the natal year, 3rd census (late summer) and the zone of capture during the natal year, 4th census (fall). NA values indicate cases when dispersal could not be estimated due to a missing zone location.
j04_11_dist	Distance (in meters) between the zone of capture during the natal year, 4th census (fall) and the zone of capture during the first reproductive year, 1st census (spring). NA values indicate cases when dispersal could not be estimated due to a missing zone location.
j11_12_dist	Distance (in meters) between the zone of capture during the first reproductive year, 1st census (spring) and the zone of capture during the first reproductive year, 2nd census (early summer). NA values indicate cases when dispersal could not be estimated due to a missing zone location.
j12_13_dist	Distance (in meters) between the zone of capture during the first reproductive year, 2nd census (early summer) and the zone of capture during the first reproductive year, 3rd census (late summer). NA values indicate cases when dispersal could not be estimated due to a missing zone location.
j13_14_dist	Distance (in meters) between the zone of capture during the first reproductive year, 3rd census (late summer) and the zone of capture during the first reproductive year, 4th census (fall). NA values indicate cases when dispersal could not be estimated due to a missing zone location.
j14_21_dist	Distance (in meters) between the zone of capture during the first reproductive year, 4th census (fall) and the zone of capture during the second reproductive year, 1st census (spring). NA values indicate cases when dispersal could not be estimated due to a missing zone location.
j21_22_dist	Distance (in meters) between the zone of capture during the second reproductive year, 1st census (spring) and the zone of capture during the second reproductive year, 2nd census (early summer). NA values indicate cases when dispersal could not be estimated due to a missing zone location.
j22_23_dist	Distance (in meters) between the zone of capture during the second reproductive year, 2nd census (early summer) and the zone of capture during the second reproductive year, 3rd census (late summer). NA values indicate cases when dispersal could not be estimated due to a missing zone location.
j23_24_dist	Distance (in meters) between the zone of capture during the second reproductive year, 3rd census (late summer) and the zone of capture during the second reproductive year, 4th census (fall). NA values indicate cases when dispersal could not be estimated due to a missing zone location.
sdisp0	Binary dispersal from the zone where an individual was first observed (natal year, 3rd census, late summer) to the zone they were captured in during their natal year, 4th census (fall); 0 meter threshold; (0 = philopatric, whereby an individual was captured in the exact same zone; 1 = disperser; whereby an individual was captured in any other zone). NA values indicate cases when dispersal could not be estimated due to a missing zone location.
disp0	Binary dispersal from the individual's presumed hatch zone to the zone of capture in their natal year 4th census (fall); 0 meter threshold; (philopatric = an individual was captured in the exact same zone; disperser = an individual was captured in any other zone). NA values indicate cases when dispersal could not be estimated due to a missing zone location.
disp5	Binary dispersal from the individual's presumed hatch zone to the zone of capture in their natal year 4th census (fall); 5 meter threshold; (philopatric = an individual was captured within 5 meters of their original zone; disperser = an individual was captured more than 5 meters from their original zone). NA values indicate cases when dispersal could not be estimated due to a missing zone location.
disp10	Binary dispersal from the individual's presumed hatch zone to the zone of capture in their natal year 4th census (fall); 10 meter threshold; (philopatric = an individual was captured within 10 meters of their original zone; disperser = an individual was captured more than 10 meters from their original zone). NA values indicate cases when dispersal could not be estimated due to a missing zone location.
mjdisp0	Binary distance between the zone of a mother at the time the offspring egg was laid and the zone where an individual was first observed (natal year, 3rd census, late summer); 0 meter threshold; (0=mother/offspring captured in the same zone; 1=mother/offspring captured in different zones). NA values indicate cases when distance could not be estimated due to a missing zone location.
mjdisp5	Binary distance between the zone of a mother at the time the offspring egg was laid and the zone where an individual was first observed (natal year, 3rd census, late summer); 5 meter threshold; (0=mother/offspring captured in zones within 5 meters of one another; 1=mother/offspring captured in zones more than 5 meters of one another). NA values indicate cases when distance could not be estimated due to a missing zone location.
mjdisp10	Binary distance between the zone of a mother at the time the offspring egg was laid and the zone where an individual was first observed (natal year, 3rd census, late summer); 10 meter threshold; (0=mother/offspring captured in zones within 10 meters of one another; 1=mother/offspring captured in zones more than 10 meters of one another). NA values indicate cases when distance could not be estimated due to a missing zone location.
SVL_0_4_fall	Measured snout-vent length (mm) during an individual's natal year, 4th census, fall
SVL_1_1_spring	Measured snout-vent length (mm) during an individual's 1st reproductive year, 1st census, spring
jdisp0	Binary dispersal from the individual's presumed hatch zone to the zone of capture in their natal year, 4th census (fall); (0 = philopatric, whereby an individual was captured in the exact same zone; 1 = disperser; whereby an individual was captured in any other zone). NA values indicate cases when dispersal could not be estimated due to a missing zone location.
Dam	Mother's ID; The unique identifying ID number given to the individual's mother.

Code/software

We used RStudio to wrangle and analyze data. The following is the complete list of packages we used in analyzing and plotting the data. The Rcode used to analyze the dataset to reproduce analyses and figures in the paper is available upon request. Requests may be sent to the Corresponding Author: C. Ashlyn Crain at the following email address - nuu6ck@virginia.edu

readxl: Wickham H, Bryan J (2022). _readxl: Read Excel Files_. R package version 1.4.1,

<https://CRAN.R-project.org/package=readxl>.
ggplot2: H. Wickham. ggplot2: Elegant Graphics for Data Analysis. Springer-Verlag New York, 2016.
psych: Revelle, W. (2022) psych: Procedures for Personality and Psychological Research,

Northwestern University, Evanston, Illinois, USA,

https://CRAN.R-project.org/package=psych Version = 2.2.9.
lmerTest: Kuznetsova A, Brockhoff PB, Christensen RHB (2017). “lmerTest Package: Tests in Linear

Mixed Effects Models.” Journal of Statistical Software, 82(13), 1-26.

doi:10.18637/jss.v082.i13. https://doi.org/10.18637/jss.v082.i13.
lme4: Douglas Bates, Martin Maechler, Ben Bolker, Steve Walker (2015). Fitting Linear

Mixed-Effects Models Using lme4. Journal of Statistical Software, 67(1), 1-48.

doi:10.18637/jss.v067.i01.
patchwork: Pedersen T (2023). _patchwork: The Composer of Plots_. R package version 1.1.3,

https://CRAN.R-project.org/package=patchwork.
DHARMa: Hartig F (2022). DHARMa: Residual Diagnostics for Hierarchical (Multi-Level / Mixed)

Regression Models. R package version 0.4.6, https://CRAN.R-project.org/package=DHARMa.
emmeans: Lenth R (2023). emmeans: Estimated Marginal Means, aka Least-Squares Means_. R package

version 1.8.5, https://CRAN.R-project.org/package=emmeans.
readr: Wickham H, Hester J, Bryan J (2023). readr: Read Rectangular Text Data. R package

version 2.1.4, https://CRAN.R-project.org/package=readr.
MASS: Venables, W. N. & Ripley, B. D. (2002) Modern Applied Statistics with S. Fourth Edition.

Springer, New York. ISBN 0-387-95457-0
fields: Douglas Nychka, Reinhard Furrer, John Paige, Stephan Sain (2021). “fields: Tools for

spatial data.” R package version 15.2, https://github.com/dnychka/fieldsRPackage.
dplyr: Wickham H, François R, Henry L, Müller K, Vaughan D (2023). dplyr: A Grammar of Data

Manipulation. R package version 1.1.2, https://CRAN.R-project.org/package=dplyr.
EnvStats: Millard SP (2013). EnvStats: An R Package for Environmental Statistics. Springer, New

York. ISBN 978-1-4614-8455-4, https://www.springer.com.
rgl: Murdoch D, Adler D (2023). rgl: 3D Visualization Using OpenGL. R package version

1.2.1, https://CRAN.R-project.org/package=rgl.
tidyverse: Wickham H, Averick M, Bryan J, Chang W, McGowan LD, François R, Grolemund G, Hayes A,

Henry L, Hester J, Kuhn M, Pedersen TL, Miller E, Bache SM, Müller K, Ooms J, Robinson

D, Seidel DP, Spinu V, Takahashi K, Vaughan D, Wilke C, Woo K, Yutani H (2019). “Welcome

to the tidyverse.” Journal of Open Source Software, 4(43), 1686. doi:10.21105/joss.01686. https://doi.org/10.21105/joss.01686.
npreg: Helwig NE (2022). npreg: Nonparametric Regression via Smoothing Splines. R package

version 1.0-9, https://CRAN.R-project.org/package=npreg.
ggformula: Kaplan D, Pruim R (2023). ggformula: Formula Interface to the Grammar of Graphics. R

package version 0.10.4, https://CRAN.R-project.org/package=ggformula.
car: John Fox and Sanford Weisberg (2019). An {R} Companion to Applied Regression, Third

Edition. Thousand Oaks CA: Sage. URL: https://socialsciences.mcmaster.ca/jfox/Books/Companion/

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Data was collected as a part of this study