DOI: 10.5061/dryad.0rxwdbsf5
title: Data from: Ornamentation is associated with social costs in male red-backed fairywrens (<em>Malurus melanocephalus</em>)

description: Sexually-selected ornaments typically confer reproductive benefits, but
 ornamented and unornamented male phenotypes often co-occur in the same
 population. Reduced social costs are one potential compensatory benefit of
 unornamented male phenotypes, but few studies have tested this hypothesis
 in a natural context. Here we present a field-based experiment testing
 whether ornamented plumage carries social costs in the red-backed
 fairywren (Malurus melanocephalus), a small Australian songbird that
 exhibits delayed but flexible plumage maturation. Only one-quarter of
 one-year-old male red-backed fairywrens acquire ornamented plumage, but
 young ornamented males obtain greater reproductive success than young
 males that retain brown, female-like plumage. We tested whether ornamented
 plumage is associated with social costs by implanting one-year-old males
 with testosterone to experimentally induce molt into ornamented plumage,
 then recorded the resulting social interactions. Testosterone-implanted
 males developed ornamentation and received more aggression than did
 control-implanted males without ornamentation, supporting the hypothesis
 that molt into ornamented plumage is socially costly. Moreover,
 one-year-old males who paired with a female and naturally acquired
 ornamented plumage exhibited smaller home ranges than implanted males,
 suggesting that paired one-year-old males who reduce their home ranges may
 experience fewer costly interactions with neighbors. Our results add to
 growing evidence that social costs can enforce the honesty of
 sexually-selected ornaments.
description: # Data from: Ornamentation is associated with social costs in male
 red-backed fairywrens (*Malurus melanocephalus*) File preparation date:
 October 25, 2025 Dates and locations of data collection: Location: Lake
 Samsonvale in Queensland, Australia (27° 16’S, 152° 41’E) Dates:
 June-August 2017, June-August 2018 Seasons are named for the year the
 breeding season ends in (i.e., these years correspond to the 2018 and 2019
 seasons). ## Summary of code and data ## Scripts Implant experiment
 script_pub.R - analyses for the plumage manipulation experiment - testing
 whether the aggression received by testosterone and control-implanted
 males differed. Results appear in Table 1 and Figure 1. Natural vs T males
 script_pub.R - compares aggression received by naturally-molting males and
 testosterone-implanted males. Results appear in the text. Homerange and
 molt script_pub.R - compares home range sizes between 1-year-old males of
 different implant types to naturally ornamented one-year-old males.
 Results appear in Figure 2. Territory size reduction simulations_pub.R -
 simulations to test whether reductions in home range size may be a
 strategy to reduce interactions with neighboring males. Results appear in
 Figure 3. ## Data file descriptions NA cells refer to not available data. 
 ### Birdlist_implant_males_pub.csv: Summary: Each row provides information
 on a one-year-old male in the implant experiment Columns: * Year - Year
 the one-year-old male was active in * FWnumber - Individual identifier for
 each bird * Bird - Colorbands of each bird (see below for a list of
 colors) * Sex - Sex of each bird * Current.Age - Age of the bird in years
 * Age.Exact - Whether the male's age was exact or estimated (min) *
 Implant - Implant type (C = control, T = testosterone, None = no implant)
 * Implant_date - Date the male was implanted (Year-month-day) * SG.Year -
 The male's social group ID and Year * chases - Number of chases
 received * attacks - Number of attacks recieved * CandA - Sum of chases
 and attacks * Breeder - Whether the male was a breeder paired with a
 female (1) or a helper (0) during the breeding season * nest - Did we find
 a nest for the male * sired - Number of chicks the male sired during the
 breeding season (within-pair and extra-pair young) ###
 1yo_edgelist_pub.csv: Summary: Edgelist showing all observed interactions
 between one-year-old male red-backed fairywrens and other red-backed
 fairywrens (RBFWs) Columns: * bird.year - Color bands of the 1-year-old
 male RBFW and the year he was 1yo * sp.ID - Sampling point ID, usually
 collected every 5 minutes, shows all birds associating in a location at a
 time * Bird.x - Color bands of the RBFW the one-year-old male was seen
 associating with (see below for a list of colors) * Date - Date of the
 observation (Year-month-day) * Time - Time of the observation (Australian
 Eastern Standard Time AEST) * Molt.x - Percent Red-black plumage exhibited
 by Bird.x * Sex - Sex of Bird.x * Current.Age - Age of Bird.x in years *
 Ageexact. - Whether Bird.x's age was exact or estimated (min) *
 Social.Group.x - Bird.x's social group ID * Bird.y - Color bands of
 the 1-year-old male RBFW (see below for a list of colors) * Molt.y -
 Percent Red-black plumage exhibited by Bird.y * Social.Group.y -
 Bird.y's social group ID * sg.same - Whether Bird.x and Bird.y were
 in the same social group * Year - Year (season) the observation was
 collected * CandA - Whether Bird.x directed any aggression towards Bird.y
 * Implant - Bird.y's implant type (C = control, T = testosterone,
 None = no implant) * sg.year - Bird.y's social group and year ###
 1yo_edgelist_t_vs_natural_pub.csv Summary: Edgelist showing all observed
 interactions between testosterone-implanted and naturally-molting
 one-year-old male red-backed fairywrens and other red-backed fairywrens
 (RBFWs) Columns: * bird.year - Color bands of the 1-year-old male RBFW and
 the year he was 1yo * sp.ID - Sampling point ID, usually collected every 5
 minutes, shows all birds associating in a location at a time * Bird.x -
 Color bands of the RBFW the one-year-old male was seen associating with
 (see below for a list of colors) * Date - Date of the observation
 (Year-month-day) * Time - Time of the observation (Australian Eastern
 Standard Time AEST) * Molt.x - Percent Red-black plumage exhibited by
 Bird.x * Sex - Sex of Bird.x * Current.Age - Age of Bird.x in years *
 Ageexact. - Whether Bird.x's age was exact or estimated (min) *
 Social.Group.x - Bird.x's social group ID * Bird.y - Color bands of
 the 1-year-old male RBFW (see below for a list of colors) * Molt.y -
 Percent Red-black plumage exhibited by Bird.y * Social.Group.y -
 Bird.y's social group ID * sg.same - Whether Bird.x and Bird.y were
 in the same social group * Year - Year (season) the observation was
 collected * CandA - Whether Bird.x directed any aggression towards Bird.y
 * Implant - Bird.y's implant type (C = control, T = testosterone,
 None = no implant) * sg.year - Bird.y's social group and year ###
 1yo_gps_18_pub.csv Summary: Waypoint locations for one-year-old males
 collected during obsevations of non-breeding social groups Columns: * WP -
 Waypoint ID * Sighting - Sampling point ID, usually collected every 5
 minutes * Observation - Observation ID, observations were usually 15-60
 minutes, with sightings collected every 5 minutes * Date - Date of the
 sighting (Year-month-day) * Time - Time of the sighting (Australian
 Eastern Standard Time AEST) * Bird - Color bands of the 1-year-old male
 the waypoint is associated with (see below for a list of colors) * lat -
 Latitude * lon - Longitude * Bird.year - Color bands and year (season) *
 Implant - The one-year-old male's implant type (C = control, T =
 testosterone, None = no implant) ### 1yo_gps_19_pub.csv Summary: Waypoint
 locations for one-year-old males collected during obsevations of
 non-breeding social groups Columns: * WP - Waypoint ID * Sighting -
 Sampling point ID, usually collected every 5 minutes * Observation -
 Observation ID, observations were usually 15-60 minutes, with sightings
 collected every 5 minutes * Date - Date of the sighting (Year-month-day) *
 Time - Time of the sighting (Australian Eastern Standard Time AEST) * Bird
 - Color bands of the 1-year-old male the waypoint is associated with (see
 below for a list of colors) * lat - Latitude * lon - Longitude * Bird.year
 - Color bands and year (season) * Implant - The one-year-old male's
 implant type (C = control, T = testosterone, None = no implant) ###
 1yo_homerange_males_pub.csv Summary: Data for the one-year-old males used
 in the home range analysis (Implant males and naturally ornamented males)
 Columns: * Bird - The one-year-old male's color bands (see below for
 a list of colors) * Year - Year the one-year-old male was active in *
 SG.Year - The one-year-old male's social group and year * Sex - Sex *
 Current.Age - Age in years * Age.Exact - Whether the male's age was
 exact or estimated (min) * Implant - The one-year-old male's implant
 type (C = control, T = testosterone, None = no implant, * Natural =
 naturally ornamented) * Bird.year - Bird and year ### akde18_list_pub.csv
 Summary: Home range size data for one-year-old males, generated in
 Homerange and molt script_pub.R Columns: * Bird - The one-year-old
 male's color bands (see below for a list of colors) * Year - Year the
 one-year-old male was active in * SG.Year - The one-year-old male's
 social group and year * Sex - Sex * Current.Age - Age in years * Age.Exact
 - Whether the male's age was exact or estimated (min) * Implant - The
 one-year-old male's implant type (C = control, T = testosterone, None
 = no implant, * Natural = naturally ornamented) * Bird.year - Bird and
 year * akde - Auto-correlated Kernel Density Estimate - home range size in
 hectares for each 1-year-old male ### akde19_list_pub.csv Summary: Home
 range size data for one-year-old males, generated in Homerange and molt
 script_pub.R Columns: * Bird - The one-year-old male's color bands
 (see below for a list of colors) * Year - Year the one-year-old male was
 active in * SG.Year - The one-year-old male's social group and year *
 Sex - Sex * Current.Age - Age in years * Age.Exact - Whether the
 male's age was exact or estimated (min) * Implant - The one-year-old
 male's implant type (C = control, T = testosterone, None = no
 implant, * Natural = naturally ornamented) * Bird.year - Bird and year *
 akde - Auto-correlated Kernel Density Estimate - home range size in
 hectares for each 1-year-old male ### associations18_pub.csv Summary: All
 non-breeding associations among red-backed fairywrens in the 2018 season
 Columns: * Sighting - Sampling point ID, usually collected every 5
 minutes, birds with the same sampling point ID were seen assocating at
 that point in time * Observation - Observation ID, observations were
 usually 15-60 minutes, with sightings collected every 5 minutes * Date -
 Date of the sighting (month/day/year) * Time - Time of the sighting
 (Australian Eastern Standard Time AEST) * WP - Waypoint ID * Bird -
 Bird's color bands (see below for a list of colors) ###
 associations19_pub.csv Summary: All non-breeding associations among
 red-backed fairywrens in the 2019 season Columns: * Sighting - Sampling
 point ID, usually collected every 5 minutes, birds with the same sampling
 point ID were seen assocating at that point in time * Observation -
 Observation ID, observations were usually 15-60 minutes, with sightings
 collected every 5 minutes * Date - Date of the sighting (month/day/year) *
 Time - Time of the sighting (Australian Eastern Standard Time AEST) * WP -
 Waypoint ID * Bird - Bird's color bands (see below for a list of
 colors) ### Ages_and_Sexes_2018_pub.csv Summary: Age and sex data for all
 red-backed fairywrens in the 2018 season network data (associations)
 Columns: * Year - Year/Season * FWNo - Individual identifier for each bird
 * Bird - Bird's color bands (see below for a list of colors) * Sex -
 Sex * Current.Age - Age of the bird in the 2018 season in years * Ageexact
 - Whether the bird's age was exact or estimated (min) ###
 Ages_and_Sexes_2019_pub.csv Summary: Age and sex data for all red-backed
 fairywrens in the 2019 season network data (associations) Columns: * Year
 - Year/Season * FWNo - Individual identifier for each bird * Bird -
 Bird's color bands (see below for a list of colors) * Sex - Sex *
 Current.Age - Age of the bird in the 2019 season in years * Ageexact -
 Whether the bird's age was exact or estimated (min) ###
 birdlist18_pub.csv Summary: List of birds in the social network data
 (associations) in the 2018 season and their social group identities
 Columns: * Bird - The bird's color bands (see below for a list of
 colors) * Social.Group - Social group the bird was a part of ###
 birdlist19_pub.csv Summary: List of birds in the social network data
 (associations) in the 2019 season and their social group identities
 Columns: * Bird - The bird's color bands (see below for a list of
 colors) * Social.Group - Social group the bird was a part of ### List of
 color band acronyms: * B = Blue * G = Green (light green) * H = Hot pink *
 I = Ivy (dark green) * L = Light blue * R = Royal purple * V = Black/white
 striped * W = White * Y = Yellow * Z = Purple/white striped ## R Session
 info: R version 4.0.4 (2021-02-15) Platform: x86_64-apple-darwin17.0
 (64-bit) Running under: macOS 13.7.8 Matrix products: default LAPACK:
 /Library/Frameworks/R.framework/Versions/4.0/Resources/lib/libRlapack.dylib locale: \[1] en_US.UTF-8/en_US.UTF-8/en_US.UTF-8/C/en_US.UTF-8/en_US.UTF-8 attached base packages: \[1] stats graphics grDevices utils datasets methods base other attached packages: \[1] emmeans_1.5.5-1 ggdist_3.2.1 asnipe_1.1.15 sp_1.4-5 lmerTest_3.1-3 ctmm_0.6.0 reshape2_1.4.4 plotrix_3.8-1\ \[9] glmmTMB_1.0.2.1 car_3.0-10 carData_3.0-4 DHARMa_0.4.1 lme4_1.1-27 Matrix_1.3-2 cowplot_1.1.1 ggplot2_3.5.1\ \[17] dplyr_1.0.7 here_1.0.1 loaded via a namespace (and not attached): \[1] Rcpp_1.0.8.3 mvtnorm_1.1-1 lattice_0.20-41 zoo_1.8-9 assertthat_0.2.1 rprojroot_2.0.2\ \[7] foreach_1.5.1 utf8_1.2.2 R6_2.5.1 cellranger_1.1.0 plyr_1.8.6 coda_0.19-4\ \[13] pillar_1.9.0 rlang_1.1.0 curl_4.3 multcomp_1.4-16 readxl_1.3.1 rstudioapi_0.13\ \[19] minqa_1.2.4 data.table_1.14.0 raster_3.4-13 nloptr_1.2.2.2 splines_4.0.4 stringr_1.5.0\ \[25] foreign_0.8-81 TMB_1.7.20 munsell_0.5.0 numDeriv_2016.8-1.1 compiler_4.0.4 pkgconfig_2.0.3\ \[31] tidyselect_1.2.0 tibble_3.1.4 rio_0.5.26 codetools_0.2-18 fansi_0.5.0 withr_2.5.2\ \[37] MASS_7.3-53.1 distributional_0.3.2 grid_4.0.4 nlme_3.1-152 xtable_1.8-4 gtable_0.3.6\ \[43] lifecycle_1.0.3 DBI_1.1.1 magrittr_2.0.3 scales_1.3.0 zip_2.1.1 estimability_1.3\ \[49] cli_3.6.1 stringi_1.7.4 farver_2.1.0 ellipsis_0.3.2 generics_0.1.3 vctrs_0.6.2\ \[55] boot_1.3-27 sandwich_3.0-0 openxlsx_4.2.3 TH.data_1.0-10 iterators_1.0.13 tools_4.0.4\ \[61] forcats_0.5.1 glue_1.6.2 hms_1.0.0 abind_1.4-5 survival_3.5-5 colorspace_2.0-2\ \[67] haven_2.3.1