DOI: 10.5061/dryad.0zpc86735
title: Data from: Carotenoid-dependent plumage coloration is associated with reduced male care in passerine birds

description: The immense diversity of plumage coloration exhibited by birds is the
 result of either pigments deposited in the feathers or microstructural
 arrangements of feather barbules. Some of the most common pigments are
 carotenoids, which produce bright yellow, orange, and red colors.
 Carotenoids differ from other pigments since birds cannot synthesize them
 de novo and must obtain them from the diet. Carotenoid pigments are
 usually associated with signaling and sexual selection, although they also
 have antioxidant properties and play a role in the immune response. Here,
 we hypothesize that carotenoid-dependent plumage coloration functions as a
 signal of a male's tendency to invest in offspring care because they
 play an important role in self-maintenance and may provide key information
 about individual quality, allowing females to obtain information about a
 male's tendency to invest in offspring care. Using phylogenetic
 comparative analyses across 349 passerine birds, we show that species that
 consume carotenoid-rich foods have more carotenoid-dependent plumage
 coloration than species with carotenoid-poor diets. In addition,
 carotenoid-dependent plumage coloration is associated with decreased male
 investment in offspring care. Our results suggest that investment in
 carotenoid-dependent plumage coloration trades off against male investment
 in offspring care and will likely have broad implications for our
 understanding of the ecological contexts that facilitate various
 evolutionary processes, such as sexual selection and signaling associated
 with plumage colors.
description: Data were collected from published sources, including both
 primary reference works (e.g., Olson and Owens 2005; Thomas et al. 2014;
 Liker et al. 2015) and compendia such as the Handbook of the Birds of the
 World Alive (del Hoyo et al. 2016; accessed between May and December
 2016).  Liker et al. (2015) compiled detailed
 information on relative investment by each sex in six components of avian
 parental care (nest building, incubation, nest guarding, chick brooding,
 chick feeding, and chick guarding (guarding and defending the brood
 post-hatching)) from reference books and published literature (references
 in Liker et al. 2015 updated in Gonzalez-Voyer et al. 2022). We focus here
 on only three components: nest building, incubation, and chick feeding,
 which are the ones for which information was available for the greatest
 number of species. We used an existing dataset (Thomas
 et al. 2014) on the presence of carotenoid-dependent plumage coloration in
 all neornithine species. Presence (1) or absence (0) of
 carotenoid-dependent coloration was scored for all species based on
 plumage colors observed in illustrations taken from the Handbook of the
 Birds of the World Alive online (del Hoyo et al. 2016), using as a
 reference a set of species for which the presence of carotenoids in
 plumage had been chemically confirmed. We selected all passerine species
 from the list and ensured that it was the male who presented
 carotenoid-dependent coloration. As we were interested not only in the
 presence of carotenoid-dependent coloration but also in the amount
 expressed on the body of the species, we developed our scoring system
 following Owens and Hartley (1998) to obtain a quantitative estimate for
 the amount of area of the body with plumage with carotenoid-dependent
 coloration. For species for which Thomas et al. (2014) had confirmed the
 presence of carotenoid coloration, we estimated the amount of
 carotenoid-dependent coloration, whereas for those species without
 carotenoid-dependent coloration according to Thomas et al. (2014), we
 assigned a value of 0. We estimated the amount of carotenoids distributed
 throughout the body of the bird because the more pigments you need, the
 more expensive the signal is likely to be, and we assume that the
 selection pressure will be stronger. We only considered the colorations
 that typically depend on these pigments: bright reds, oranges, and
 yellows. To minimize bias, we selected three volunteers with birdwatching
 experience, but uninformed about the hypotheses we wished to test, to
 score the extent of each patch covered by carotenoid-dependent plumage
 coloration. Each observer was shown illustrations from the Handbook of the
 Birds of the World of the 177 passerine species that Thomas et al. (2014)
 classified as presenting carotenoid-dependent plumage coloration and asked
 to score patches with: if the carotenoid-dependent coloration covered the
 whole patch or a part of it; 0 if it did not show carotenoid-dependent
 coloration. The observers scored eight body regions per species: head,
 nape, back, rump, throat, chest, belly, tail, and wings. In the rare
 instances where the patch observers were meant to score was not visible in
 the images, we searched descriptions and pictures of the species to verify
 if they had carotenoid-dependent coloration in the patch. We scored these
 regions because they cover the whole body of the bird. In cases where
 multiple subspecies were illustrated, we scored coloration in the nominate
 subspecies. The continuous measure for carotenoid-dependent coloration was
 the sum of the scores for all patches for each species.
 To assess the carotenoid content in the diet of each species, we
 developed a scoring system following Olson and Owens (2005), who scored
 carotenoid intake based on a coarse-scale index that ranks seven diet
 categories based on their carotenoid content: 1 for seeds, nuts, wood; 2
 for nectar, pollen, sap, exudates, lerps; 3 for vertebrates; 4 for
 invertebrates; 5 for foliage, flowers, fungi; 6 for fruit; and 7 for
 algae, diatoms. We collected diet data from verbal descriptions in the
 Handbook of the Birds of the World Alive (del Hoyo et al. 2016). Since
 most species do not feed solely on a single group, we assigned relative
 values based on the importance of each group in their diet as described in
 HBW Alive.  We used body mass as an estimate of the
 body size of the males of each species. We included body size in our
 analyses because it is strongly correlated with life history traits and
 could reflect possible restrictions for the development of plumage
 coloration in birds, such as metabolic rate and the ingestion of food with
 carotenoids. Body mass data were obtained from Liker et al. (2015) for
 almost all species. For species lacking data, we extracted it from Dale et
 al. (2015) and the HBW Alive. Log-transformed values of body mass were
 used in the statistical analysis. We obtained the
 species’ breeding distribution from del Hoyo et al. (2016) for
 latitude.
description: # Data from: Carotenoid-dependent plumage coloration is associated with
 reduced male care in passerine birds * Authors: Veronica Rincon Rubio *
 Dataset Contributors: Veronica Rincon Rubio, Alejandro Gonzalez Voyer,
 Andras Liker, Tamas Szekely * Date created: 2016-05-20 * Date modified:
 2023-03-12 * Suggested Citations: * Dataset citation: Rincn-Rubio, V. A.,
 Szkely, T., Liker, A., & Gonzalez-Voyer, A. 2023. Data from:
 Carotenoid-dependent plumage coloration is associated with reduced male
 care in passerine birds. Behavioral Ecology.
 [https://doi.org/10.5061/dryad.0zpc86735](https://doi.org/10.5061/dryad.0zpc86735) ## Contact Information * Name: Alejandro Gonzalez Voyer * Affiliations: Departamento de Ecologa Evolutiva, Instituto de Ecologa, Universidad Nacional Autnoma de Mxico, Circuito Exterior AP 70-275, Mxico, DF 04510, Mexico * ORCID ID: [http://orcid.org/0000-0002-5072-1688](http://orcid.org/0000-0002-5072-1688) * Email: [alejandro.gonzalez@iecologia.unam.mx](mailto:alejandro.gonzalez@iecologia.unam.mx) * Address: e-mail preferred * Alternative Contact: * Name: Veronica A. Rincon Rubio * Affiliations: Departamento de Ecologa Evolutiva, Instituto de Ecologa, Universidad Nacional Autnoma de Mxico, Circuito Exterior AP 70-275, Mxico, DF 04510, Mexico; Departamento de Ecologa Evolutiva, Instituto de Ecologa, Universidad Nacional Autnoma de Mxico, Circuito Exterior AP 70-275, Mxico, DF 04510, Mexico * Email: [veronica.rincon@ciencias.unam.mx](mailto:veronica.rincon@ciencias.unam.mx) * Address: e-mail preferred * Contributor ORCID IDs: * Tamas Szekely: [https://orcid.org/0000-0001-9985-859X](https://orcid.org/0000-0001-9985-859X) * Andras Liker: [https://orcid.org/0000-0001-8545-4869](https://orcid.org/0000-0001-8545-4869) #### Data and File Overview Table of Contents #### Summary Metrics * File count: 2 * Total file size: 63 KB * File formats: .csv; .tre #### Table of Contents * FinalDataF.csv * FinalDataF.numbers * ConsensusTree.tre #### Setup * Unpacking instructions: n/a * Recommended software/tools: RStudio 1.2.5033; R version 4.1.2 #### Details for: FinalDataF.csv This dataset contains a combination of information collated from published studies and data that was collected anew. The new data includes an estimate of the amount of the body of a given male passerine which is covered by carotenoid-dependent plumage coloration as well as the estimated amount of carotenoids obtained from the diet. Details are available in the published article and in the Dryad website. * Description: A comma-delimited file containing the species scientific name, the weight of average males, whether males present carotenoid-dependent plumage coloration or not, an estimate of the proportion of the body covered by such coloration, an amount of carotenoids obtained from the diet, the distribution, and an estimate of the frequency of polygamy in the species. * Format(s): .csv * Size(s): 30 KB * Dimensions: 392 rows by 11 columns * Variables: *Species: Scientific name of each species according to the nomenclature of Jetz et al. 2012 *Family: Family to which each species corresponds according to Thomas et al. 2014 *Thomas: Plumage pigmentation of modern birds coded in reference to colors produced by carotenoid pigments. Character state [0] indicates an absence of carotenoid-consistent colors in plumge. Character state [1] indicates the presence of carotenoid-consistent colors in plumage accordinf to Thomas et al. 2014. *ConfirmedDelhey: Confirmed species with presence or absence of carotenoids according to Delhey et al. 2023. Character state [1] indicates consistent between Thomas et al. 2014 and Delhey et al. 2023. *Mean: Average participation of males of each species in three care components: nest building, incubation, and chick feeding. This mean was calculated according to the relative participation of the males in each component following the Liker et al. 2015 score. *CP2: Mean scores of relative participation by males scored by Liker et al. (2015). The activities are: nest building, incubation, and chick feeding. *Size: Log-transformed values of body mass. Body mass (in g) data were obtained from Liker et al. (2015) for almost all species. For species lacking data, we extracted it from Dale et al. (2015) and the HBW Alive. *Diet: Mean score of carotenoid content from the relative importance of each diet category multiplied by the amount of carotenoids contained according to Olson and Owens (2005). *Distribution: Categorizationon based on whether they presented a tropical distribution (23 N to 23 S), a non-tropical distribution (north of 23 N, or south of 23 S), or the distribution spanned both tropical and non-tropical areas (species breeding in non-tropical and tropical areas). We obtained the species breeding distribution from del Hoyo et al. (2016) *Scrpoligamia: Polygamy frequency data for males according to Liker et al. (2015) who scored the overall incidence of social polygamy for each sex on a scale from 0 to 4, with 0 corresponding to no (or very rare) polygamy (&lt;0.1% of individuals), 1 to rare polygamy (0.11%), 2 to uncommon polygamy (15%), 3 to moderate polygamy (520%), and 4 to common polygamy (&gt;20%). * Missing data codes: "NA" #### Details for: Details for: ConsensusTree.tre * Description: Maximum clade credibility tree from 1000 phylogenetic trees randomly selected from 10,000 phylogenies from the pseudo-posterior distribution from the avian phylogeny of Jetz et al. (2012; available at birdtree.org). * Format(s): .tre * Size(s): 33 KB * Dimensions: 391 tips; 390 nodes; mean of branch lengths: 7.43 67.54 Thomas Delhey consistency in the presence of carotenoid coloration of the species * Chrysomus_ruficapillus * Dendroica_caerulescens * Dendroica_cerulea * Nectarinia_amethystina * Ptilonorhynchus_violaceus * Sayornis_nigricans **Sources** Dale J, Dey CJ, Delhey K, Kempenaers B, Valcu M. 2015. Data from: The effects of life history and sexual selection on male and female plumage colouration. Nature. 527(7578):367370. doi:10.1038/nature15509. Delhey K, Valcu M, Dale J, Kempenaers B. 2023. Data from: The evolution of carotenoid-based plumage colours in passerine birds. J Anim Ecol. 92:(1):6677. del Hoyo J, Elliott A, Sargatal J, Christie DA, de Juana E. 2016. Handbook of the birds of the world alive. Barcelona: Lynx Edicions. [http://www.hbw.com/](http://www.hbw.com/). Accessed between May and December 2016. Liker A, Freckleton RP, Reme V, Szkely T. 2015. Data from: Sex differences in parental care: gametic investment, sexual selection, and social environment. Evolution. 69(11):28622875. doi:10.1111/evo.12786. Olson VA, Owens IPF. 2005. Data from: Interspecific variation in the use of carotenoid-based coloration in birds: diet, life history and phylogeny. J Evol Biol. 18(6):15341546. doi:10.1111/j.1420-9101.2005.00940.x. Thomas Daniel B., McGraw Kevin J., Butler Michael W., Carrano Matthew T., Madden Odile and James Helen F. 2014. Data from: Ancient origins and multiple appearances of carotenoid-pigmented feathers in birdsProc. R. Soc. B.2812014080620140806 [http://doi.org/10.1098/rspb.2014.0806](http://doi.org/10.1098/rspb.2014.0806)