The hidden world of egg coloration: Exploring the drivers of variation in UV reflectance
Understanding how selection pressures operate at different evolutionary scales to promote diversity in various species' traits and phenotypes has long intrigued evolutionary biologists. In recent years, efforts have been made to comprehend the function of UV reflectance in avian eggs, leading t...
- Autores:
-
Mendiwelso Moreno, Maria Elisa
- Tipo de recurso:
- Trabajo de grado de pregrado
- Fecha de publicación:
- 2023
- Institución:
- Universidad de los Andes
- Repositorio:
- Séneca: repositorio Uniandes
- Idioma:
- eng
- OAI Identifier:
- oai:repositorio.uniandes.edu.co:1992/69438
- Acceso en línea:
- http://hdl.handle.net/1992/69438
- Palabra clave:
- Eggshell color
UV reflectance
Macroecological analysis
Light exposure
Biología
- Rights
- openAccess
- License
- Attribution-NonCommercial-NoDerivatives 4.0 Internacional
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dc.title.none.fl_str_mv |
The hidden world of egg coloration: Exploring the drivers of variation in UV reflectance |
dc.title.alternative.none.fl_str_mv |
El mundo oculto de la coloración de los huevos: Exploración de los factores de variación de la reflectancia UV |
title |
The hidden world of egg coloration: Exploring the drivers of variation in UV reflectance |
spellingShingle |
The hidden world of egg coloration: Exploring the drivers of variation in UV reflectance Eggshell color UV reflectance Macroecological analysis Light exposure Biología |
title_short |
The hidden world of egg coloration: Exploring the drivers of variation in UV reflectance |
title_full |
The hidden world of egg coloration: Exploring the drivers of variation in UV reflectance |
title_fullStr |
The hidden world of egg coloration: Exploring the drivers of variation in UV reflectance |
title_full_unstemmed |
The hidden world of egg coloration: Exploring the drivers of variation in UV reflectance |
title_sort |
The hidden world of egg coloration: Exploring the drivers of variation in UV reflectance |
dc.creator.fl_str_mv |
Mendiwelso Moreno, Maria Elisa |
dc.contributor.advisor.none.fl_str_mv |
Ocampo Rincón, David Cadena Ordóñez, Carlos Daniel |
dc.contributor.author.none.fl_str_mv |
Mendiwelso Moreno, Maria Elisa |
dc.contributor.researchgroup.es_CO.fl_str_mv |
Biología Evolutiva de Vertebrados (EVOLVERT) |
dc.subject.keyword.none.fl_str_mv |
Eggshell color UV reflectance Macroecological analysis Light exposure |
topic |
Eggshell color UV reflectance Macroecological analysis Light exposure Biología |
dc.subject.themes.es_CO.fl_str_mv |
Biología |
description |
Understanding how selection pressures operate at different evolutionary scales to promote diversity in various species' traits and phenotypes has long intrigued evolutionary biologists. In recent years, efforts have been made to comprehend the function of UV reflectance in avian eggs, leading to the proposal of different hypotheses. Among the most extensively studied are the UV resistance hypothesis and the egg detectability hypothesis, both of which we investigated in this study to determine to which one may explain patterns in UV reflectance and to evaluate the driving forces associated with such variation. This study is the first to take a large-scale macroecological view of eggshell UV coloration in novel data collected for over 500 avian species and analyzed using phylogenetic comparative methods. We identified the importance of brightness and the exposure of the nest in relation to the UV chroma reflected by the egg. Likewise, although we did not observe a statistically significant effect of nest type on UV reflectance, we did find patterns that, taking into account the mean UV reflectance measurements and character mapping, reveal greater support for the UV resistance hypothesis in Passeriformes and Charadriiformes, which could suggest future research. Overall, this research contributes to a deeper understanding of the mechanisms driving UV coloration in avian eggs and sheds light on the complex interplay between selection pressures and the evolution of species' traits. |
publishDate |
2023 |
dc.date.accessioned.none.fl_str_mv |
2023-08-08T19:24:58Z |
dc.date.available.none.fl_str_mv |
2023-08-08T19:24:58Z |
dc.date.issued.none.fl_str_mv |
2023-08-04 |
dc.type.es_CO.fl_str_mv |
Trabajo de grado - Pregrado |
dc.type.driver.none.fl_str_mv |
info:eu-repo/semantics/bachelorThesis |
dc.type.version.none.fl_str_mv |
info:eu-repo/semantics/acceptedVersion |
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http://purl.org/coar/resource_type/c_7a1f |
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http://purl.org/redcol/resource_type/TP |
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http://purl.org/coar/resource_type/c_7a1f |
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dc.identifier.uri.none.fl_str_mv |
http://hdl.handle.net/1992/69438 |
dc.identifier.instname.es_CO.fl_str_mv |
instname:Universidad de los Andes |
dc.identifier.reponame.es_CO.fl_str_mv |
reponame:Repositorio Institucional Séneca |
dc.identifier.repourl.es_CO.fl_str_mv |
repourl:https://repositorio.uniandes.edu.co/ |
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http://hdl.handle.net/1992/69438 |
identifier_str_mv |
instname:Universidad de los Andes reponame:Repositorio Institucional Séneca repourl:https://repositorio.uniandes.edu.co/ |
dc.language.iso.es_CO.fl_str_mv |
eng |
language |
eng |
dc.relation.references.es_CO.fl_str_mv |
Cherry, M. I., & Bennett, T. D. (2001). Egg colour matching in an African cuckoo, as revealed by ultraviolet-visible reflectance spectrophotometry. Proceedings of the Royal Society of London. Series B: Biological Sciences, 268(1467), 565-571. https://doi.org/10.1098/rspb.2000.1414 Price-Waldman, R., & Stoddard, M. C. (2021). Avian Coloration Genetics: Recent Advances and Emerging Questions. Journal of Heredity, 112(5), 395-416. https://doi.org/10.1093/jhered/esab015 Riehl, C. (2011). Paternal investment and the "sexually selected hypothesis" for the evolution of eggshell coloration: Revisiting the assumptions. Auk, 128(1), 175-179. https://doi.org/10.1525/auk.2011.10171 Jagannath, A., Shore, R. F., Walker, L. A., Ferns, P. N., & Gosler, A. G. (2008). Eggshell pigmentation indicates pesticide contamination. Journal of Applied Ecology, 45(1), 133-140. https://doi.org/10.1111/j.1365-2664.2007.01386.x Orlowski, G., Niedzielski, P., Merta, D., Pokorny, P., & Proch, J. (2020). Quantifying the functional disparity in pigment spot-background egg colour ICP-OES-based eggshell ionome at two extremes of avian embryonic development. Scientific Reports, 10(1), 1-15. https://doi.org/10.1038/s41598-020-79040-4 Hanley, D., Cassey, P., & Doucet, S. M. (2013). Parents, predators, parasites, and the evolution of eggshell colour in open nesting birds. Evolutionary Ecology, 27(3), 593-617. https://doi.org/10.1007/s10682-012-9619-6 Dainson, M., Hauber, M. E., López, A. V., Grim, T., & Hanley, D. (2017). Does contrast between eggshell ground and spot coloration affect egg rejection?. The Science of Nature, 104, 1-9. https://doi.org/10.1007/s00114-017-1476-2 Yang, C., Wang, J., and Liang, W. (2016). Blocking of ultraviolet reflectance on bird eggs reduces nest predation by aerial predators. Journal of Ornithology, 157: 43-47. https://doi.org/10.1007/s10336-015-1243-0 Honza, M., Polaciková, L., & Procházka, P. (2007). Ultraviolet and green parts of the colour spectrum affect egg rejection in the song thrush (Turdus philomelos). Biological Journal of the Linnean Society, 92(2), 269-276. https://doi.org/10.1111/j.1095-8312.2007.00848.x Lahti, D. C., & Ardia, D. R. (2016). Shedding light on bird egg color: Pigment as parasol and the dark car effect. The American Naturalist, 187(5), 547-563. https://doi.org/10.1086/685780 Mayani-Parás, F., Kilner, R. M., Stoddard, M. C., Rodríguez, C., & Drummond, H. (2015). Behaviorally induced camouflage: a new mechanism of avian egg protection. The American Naturalist, 186(4), E91-E97. https://doi.org/10.1086/682579 Hanley, D., Doucet, S. M., & Dearborn, D. C. (2010). A blackmail hypothesis for the evolution of conspicuous egg coloration in birds. The Auk, 127(2), 453-459. https://doi.org/10.1525/auk.2009.09090 Soler, J. J., Moreno, J., Aviles, j., & Moller, A. P. (2005). Blue and green egg-color intensity is associated with parental: effort and mating system in passerines: support for: the sexual selection hypothesis. Evolution, 59(3), 636-644. https://doi.org/10.1525/auk.2009.09090 Ladouce, M., Barakat, T., Su, B. L., Deparis, O., & Mouchet, S. R. (2020). Scattering of ultraviolet light by avian eggshells. Faraday Discussions, 223, 63-80. https://doi.org/10.1039/D0FD00034E Cassey, P., Thomas, G. H., Portugal, S. J., Maurer, G., Hauber, M. E., Grim, T.Lovell, G,. Miksík, I. (2012). Why are birds' eggs colourful? Eggshell pigments co-vary with life-history and nesting ecology among British breeding non-passerine birds. Biological Journal of the Linnean Society, 106(3), 657-672. https://doi.org/10.1111/j.1095-8312.2012.01877.x Hansell, M. (2000). Bird nests and construction behaviour. Cambridge University Press. Paul, N. D., & Gwynn-Jones, D. (2003). Ecological roles of solar UV radiation: towards an integrated approach. Trends in Ecology & Evolution, 18(1), 48-55. https://doi.org/10.1016/S0169-5347(02)00014-9 Wang, J., Yang, C., Shi, H., & Liang, W. (2016). Reflectance and artificial nest experiments of reptile and bird eggs imply an adaptation of bird eggs against ultraviolet. Ecological research, 31(1), 105-110. https://doi.org/10.1007/s11284-015-1317-8 Avilés, J. M., Soler, J. J., & Pérez-Contreras, T. (2006). Dark nests and egg colour in birds: A possible functional role of ultraviolet reflectance in egg detectability. Proceedings of the Royal Society B: Biological Sciences, 273(1603), 2821-2829. https://doi.org/10.1098/rspb.2006.3674 Ödeen, A., Håstad, O., & Alström, P. (2011). Evolution of ultraviolet vision in the largest avian radiation-the passerines. BMC Evolutionary Biology, 11(1), 1-8. https://doi.org/10.1186/1471-2148-11-313 Maurer, G., Portugal, S. J., & Cassey, P. (2011). An embryo's eye view of avian eggshell pigmentation. Journal of Avian Biology, 42(6), 494-504. https://doi.org/10.1111/j.1600-048X.2011.05368.x Clements, J. F., T. S. Schulenberg, M. J. Iliff, T. A. Fredericks, J. A. Gerbracht, D. Lepage, S. M. Billerman, B. L. Sullivan, and C. L. Wood. (2021). The eBird/Clements checklist of Birds of the World: v2021. Downloaded from https://www.birds.cornell.edu/clementschecklist/download/ R Core Team (2022). R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. URL https://www.R-project.org/. D'Alba, L., Torres, R., Waterhouse, G. I., Eliason, C., Hauber, M. E., & Shawkey, M. D. (2017). What does the eggshell cuticle do? A functional comparison of avian eggshell cuticles. Physiological and Biochemical Zoology, 90(5), 588-599. https://doi.org/10.1086/693434 Maia, R., Eliason, C. M., Bitton, P. P., Doucet, S. M., & Shawkey, M. D. (2013). pavo: an R package for the analysis, visualization and organization of spectral data. Methods in Ecology and Evolution, 4(10), 906-913. Billerman, S. M., Keeney, B. K., Rodewald, P. G., & Schulenberg, T. S. (2020). Birds of the World. Cornell Laboratory of Ornithology, Ithaca, NY, USA. Simón, J. E., & Pacheco, S. (2005). On the standardization of nest descriptions of neotropical birds. Revista Brasileira de ornitologia, 13(2), 143-154. Tobias, J. A. Sheard, C., Pigot, A. L., Devenish, A. J. M., Yang, J., Sayol, F., Neate-Clegg, M. H. C., Alioravainen, N., Weeks, T. L., Barber, R. A., Walkden, P. A., MacGregor, H. E. A., Jones, S. E. I., Vincent, C., Phillips, A. G., Marples, N. M., Montaño-Centellas, F. A., Leandro-Silva, V., Claramunt, S,Schleuning, M. (2022). AVONET: Morphological, ecological and geographical data for all birds. Ecology Letters, 25, 581-597. https://doi.org/10.1111/ele.13898 Englert Duursma, D., Gallagher, R. V., Price, J. J., & Griffith, S. C. (2018). Variation in avian egg shape and nest structure is explained by climatic conditions. Scientific Reports, 8(1), 1-10. https://doi.org/10.1038/s41598-018-22436-0 Heenan, C. B. (2013). An overview of the factors influencing the morphology and thermal properties of avian nests. Avian Biology Research, 6(2), 104-118. https://doi.org/10.3184/003685013X13614670646299 Gómez, J., Ramo, C., Stevens, M., Liñán-Cembrano, G., Rendón, M. A., Troscianko, J. T., & Amat, J. A. (2018). Latitudinal variation in biophysical characteristics of avian eggshells to cope with differential effects of solar radiation. Ecology and Evolution, 8(16), 8019-8029. https://doi.org/10.1002/ece3.4335 Cherry, M. I., & Gosler, A. G. (2010). Avian eggshell coloration: new perspectives on adaptive explanations. Biological Journal of the Linnean Society, 100(4), 753-762. https://doi.org/10.1111/j.1095-8312.2010.01457.x Kilner, R. M. (2006). The evolution of egg colour and patterning in birds. Biological Reviews, 81(3), 383-406. https://doi.org/10.1017/S1464793106007044 Wegrzyn, E., Leniowski, K., Rykowska, I., & Wasiak, W. (2011). Is UV and blue-green egg colouration a signal in cavity-nesting birds?. Ethology Ecology & Evolution, 23(2), 121-139. https://doi.org/10.1080/03949370.2011.554882 Maurer, G., Portugal, S. J., Hauber, M. E., Miksík, I., Russell, D. G., & Cassey, P. (2015). First light for avian embryos: eggshell thickness and pigmentation mediate variation in development and UV exposure in wild bird eggs. Functional Ecology, 29(2), 209-218. https://doi.org/10.1111/1365-2435.12314 Schneider, C. A., Rasband, W. S., & Eliceiri, K. W. (2012). NIH Image to ImageJ: 25 years of image analysis. Nature Methods, 9(7), 671-675. doi:10.1038/nmeth.2089 Nagy, J., Hauber, M. E., Hartley, I. R., & Mainwaring, M. C. (2019). Correlated evolution of nest and egg characteristics in birds. Animal Behaviour, 158, 211-225. https://doi.org/10.1016/j.anbehav.2019.10.015 Stoddard, M. C., Yong, E. H., Akkaynak, D., Sheard, C., Tobias, J. A., & Mahadevan, L. (2017). Avian egg shape: Form, function, and evolution. Science, 356(6344), 1249-1254. DOI: 10.1126/science.aaj1945 Jetz, W., Thomas, G. H., Joy, J. B., Hartmann, K., & Mooers, A. O. (2012). The global diversity of birds in space and time. Nature, 491(7424), 444-448. https://doi.org/10.1038/nature11631 Ocampo, D., De Silva, T.N., Sheard, C., Stoddard, M.C. (2023). Evolution of nest architecture in tyrant flycatchers and allies. Philosophical Transactions B. https://doi.org/10.1098/rstb.2022.0148 Bouckaert, R., Drummond, A., Rambaut, A., Suchard, M., Vaughan, T., & Heled, Y. (2014). BEAST2: Bayesian evolutionary analysis sampling trees. Revell, L. J. (2012). Phytools: An R package for phylogenetic comparative biology (and other things). Methods in Ecology and Evolution, 3(2), 217-223 Revell, L. J. (2013). Two new graphical methods for mapping trait evolution on phylogenies. Methods in Ecology and Evolution, 4(8), 754-759. https://doi.org/10.1111/2041-210X.12066 Orme, D., Freckleton, R., Thomas, G. Petzoldt, T., Fritz, S., Isaac, N., & Pearse, W. (2018). caper: Comparative Analyses of Phylogenetics and Evolution in R. R package version 1.0.1 Burnham, K. P., Anderson, D. R., & Huyvaert, K. P. (2011). AIC model selection and multimodel inference in behavioral ecology: Some background, observations, and comparisons. Behavioral Ecology and Sociobiology, 65(1), 23-35. https://doi.org/10.1007/s00265-010-1029-6 Barton, K. (2019). MuMIn: Multi-model inference. R package version 1.43.15. Retrieved from http://r-forge.r-project.org/projects/mumin/. Burnham, K. P., & Anderson, D. R. (2002). A practical information-theoretic approach. In Model selection and multimodel inference (2nd ed., p. 2). Springer. D'Alba, L., Goldenberg, J., Nallapaneni, A., Parkinson, D. Y., Zhu, C., Vanthournout, B., & Shawkey, M. D. (2021). Evolution of eggshell structure in relation to nesting ecology in non-avian reptiles. Journal of morphology, 282(7), 1066-1079. https://doi.org/10.1002/jmor.21347 Wisocki, P. A., Kennelly, P., Rojas Rivera, I., Cassey, P., Burkey, M. L., & Hanley, D. (2020). The global distribution of avian eggshell colours suggest a thermoregulatory benefit of darker pigmentation. Nature Ecology & Evolution, 4(1), 148-155. https://doi.org/10.1038/s41559-019-1003-2 Hanley, D., Grim, T., Cassey, P., & Hauber, M. E. (2015). Not so colourful after all: eggshell pigments constrain avian eggshell colour space. Biology letters, 11(5), 20150087. https://doi.org/10.1098/rsbl.2015.0087 Lee, W. S., Kwon, Y. S., & Yoo, J. C. (2010). Egg survival is related to the colour matching of eggs to nest background in Black-tailed Gulls. Journal of Ornithology, 151, 765-770. https://doi.org/10.1007/s10336-010-0508-x Yang, C., Møller, A. P., & Liang, W. (2022). Light matters: Nest illumination alters egg rejection behavior in a cavity-nesting bird. Avian Research, 13, 100016. https://doi.org/10.1016/j.avrs.2022.100016 Maziarz, M., & Wesolowski, T. (2014). Does darkness limit the use of tree cavities for nesting by birds?. Journal of Ornithology, 155, 793-799. https://doi.org/10.1007/s10336-014-1069-1 Moreno, J., & Osorno, J. L. (2003). Avian egg colour and sexual selection: does eggshell pigmentation reflect female condition and genetic quality?. Ecology Letters, 6(9), 803-806. https://doi.org/10.1046/j.1461-0248.2003.00505.x Lahti, D. C. (2008). Population differentiation and rapid evolution of egg color in accordance with solar radiation. The Auk, 125(4), 796-802. https://doi.org/10.1525/auk.2008.07033 L'Herpiniere, K. L., Tims, A. R., Englert Duursma, D., & Griffith, S. C. (2021). The evolution of egg colour and patterning in Australian songbirds. Evolution, 75(12), 3132-3141. https://doi.org/10.1111/evo.14375 Gosler, A. G., Higham, J. P., & James Reynolds, S. (2005). Why are birds' eggs speckled?. Ecology Letters, 8(10), 1105-1113. https://doi.org/10.1111/j.1461-0248.2005.00816.x Cassey, P., Thomas, G. H., Portugal, S. J., Maurer, G., Hauber, M. E., Grim, T., & Miksík, I. (2012). Why are birds' eggs colourful? Eggshell pigments co-vary with life-history and nesting ecology among British breeding non-passerine birds. Biological Journal of the Linnean Society, 106(3), 657-672. https://doi.org/10.1111/j.1095-8312.2012.01877.x Yang, C., Wang, L., Hsu, Y.-C., Antonov, A., Moksnes, A., Røskaft, E., Liang, W. & Stokke, B. G. (2013) UV reflectance as a cue in egg discrimination in two Prinia species exploited differently by brood parasites in Taiwan. The Ibis 155, 571-575. https://doi.org/10.1111/ibi.12043 Honza, M., Capek, M., Mikulica, O., & Samas, P. (2022). Ultraviolet coloration of avian parasitic egg does not cue egg rejection in the common redstart host. Journal of Ornithology, 163(4), 903-909. https://doi.org/10.1007/s10336-022-01991-4 Ericson PG, Anderson CL, Britton T, Elzanowski A, Johansson US et al., (2006). Diversification of Neoaves: Integration of molecular sequence data and fossils. Biol. Lett. 2: 543-547. https://doi.org/10.1098/rsbl.2006.0523 Gómez, J., Pereira, A. I., Pérez-Hurtado, A., Castro, M., Ramo, C., & Amat, J. A. (2016). A trade-off between overheating and camouflage on shorebird eggshell colouration. Journal of Avian Biology, 47(3), 346-353. https://doi.org/10.1111/jav.00736 Kinoshita, S., Yoshioka, S., & Miyazaki, J. (2008). Physics of structural colors. Reports on Progress in Physics, 71(7), 076401. DOI 10.1088/0034-4885/71/7/076401 Honza, M., Sulc, M., & Cherry, M. I. (2014). Does nest luminosity play a role in recognition of parasitic eggs in domed nests? A case study of the red bishop. Naturwissenschaften, 101, 1009-1015. https://doi.org/10.1007/s00114-014-1240-9 Xiao, H., Hu, Y., Lang, Z., Fang, B., Guo, W., Zhang, Q. I et al (2017). How much do we know about the breeding biology of bird species in the world?. Journal of Avian Biology, 48(4), 513-518. https://doi.org/10.1111/jav.00934 Wiemann, J., Yang, T. R., & Norell, M. A. (2018). Dinosaur egg colour had a single evolutionary origin. Nature, 563(7732), 555-558. https://doi.org/10.1038/s41586-018-0646-5 Marki, P. Z., Fabre, P. H., Jønsson, K. A., Rahbek, C., Fjeldså, J., & Kennedy, J. D. (2015). Breeding system evolution influenced the geographic expansion and diversification of the core Corvoidea (Aves: Passeriformes). Evolution, 69(7), 1874-1924. https://doi.org/10.1111/evo.12695 Carvalho, C. B., Macedo, R. H., & Graves, J. A. (2006). Breeding strategies of a socially monogamous neotropical passerine: extra-pair fertilizations, behavior, and morphology. The Condor, 108(3), 579-590. https://doi.org/10.1093/condor/108.3.579 |
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Attribution-NonCommercial-NoDerivatives 4.0 Internacionalhttp://creativecommons.org/licenses/by-nc-nd/4.0/info:eu-repo/semantics/openAccesshttp://purl.org/coar/access_right/c_abf2Ocampo Rincón, Davidc8acf79f-0ca3-4226-b22e-1d1089143f0b600Cadena Ordóñez, Carlos Daniel8308cacd-eb61-4f56-9496-b997382da70d600Mendiwelso Moreno, Maria Elisaf1453d65-d195-4250-8271-d0c8b47a5217600Biología Evolutiva de Vertebrados (EVOLVERT)2023-08-08T19:24:58Z2023-08-08T19:24:58Z2023-08-04http://hdl.handle.net/1992/69438instname:Universidad de los Andesreponame:Repositorio Institucional Sénecarepourl:https://repositorio.uniandes.edu.co/Understanding how selection pressures operate at different evolutionary scales to promote diversity in various species' traits and phenotypes has long intrigued evolutionary biologists. In recent years, efforts have been made to comprehend the function of UV reflectance in avian eggs, leading to the proposal of different hypotheses. Among the most extensively studied are the UV resistance hypothesis and the egg detectability hypothesis, both of which we investigated in this study to determine to which one may explain patterns in UV reflectance and to evaluate the driving forces associated with such variation. This study is the first to take a large-scale macroecological view of eggshell UV coloration in novel data collected for over 500 avian species and analyzed using phylogenetic comparative methods. We identified the importance of brightness and the exposure of the nest in relation to the UV chroma reflected by the egg. Likewise, although we did not observe a statistically significant effect of nest type on UV reflectance, we did find patterns that, taking into account the mean UV reflectance measurements and character mapping, reveal greater support for the UV resistance hypothesis in Passeriformes and Charadriiformes, which could suggest future research. Overall, this research contributes to a deeper understanding of the mechanisms driving UV coloration in avian eggs and sheds light on the complex interplay between selection pressures and the evolution of species' traits.Entender cómo operan las presiones de selección a diferentes escalas evolutivas para promover la diversidad en los rasgos y fenotipos de diversas especies ha intrigado durante mucho tiempo a los biólogos evolutivos. En los últimos años, se han hecho esfuerzos para comprender la función de la reflectancia UV en los huevos de las aves, lo que ha llevado a proponer diferentes hipótesis. Entre las más ampliamente estudiadas se encuentran la hipótesis de la resistencia a los rayos UV y la hipótesis de la detectabilidad del huevo, las cuales investigamos en este estudio para determinar cuál de ellas podría explicar patrones en la reflectancia UV y evaluar las fuerzas impulsoras asociadas a dicha variación. Este estudio es el primero que adopta una visión macroecológica a gran escala sobre la coloración UV de la cáscara de los huevos usando datos nuevos de más de 500 especies de aves analizados con métodos filogenéticos comparativos. Identificamos la importancia del brillo y la exposición del nido en relación con el croma UV reflejado por el huevo. Asimismo, aunque no observamos un efecto estadísticamente significativo del tipo de nido sobre la reflectancia UV, si observamos patrones que, tomando en cuenta la media de las medidas de la reflectancia UV y el mapeo de caracteres, indican que hay un mayor apoyo a la hipótesis de la resistencia UV, particularmente en Passeriformes y Charadriiformes, lo que abre la puerta para futuras investigaciones. En general, esta investigación contribuye a una comprensión más profunda de los mecanismos que impulsan la coloración UV en los huevos de las aves y arroja luz sobre la compleja interacción entre las presiones de selección y la evolución de los rasgos de las especies.Neotropical Ornithological SocietyBiólogoPregrado30 páginasapplication/pdfengUniversidad de los AndesBiologíaFacultad de CienciasDepartamento de Ciencias BiológicasThe hidden world of egg coloration: Exploring the drivers of variation in UV reflectanceEl mundo oculto de la coloración de los huevos: Exploración de los factores de variación de la reflectancia UVTrabajo de grado - Pregradoinfo:eu-repo/semantics/bachelorThesisinfo:eu-repo/semantics/acceptedVersionhttp://purl.org/coar/resource_type/c_7a1fTexthttp://purl.org/redcol/resource_type/TPEggshell colorUV reflectanceMacroecological analysisLight exposureBiologíaCherry, M. I., & Bennett, T. D. (2001). Egg colour matching in an African cuckoo, as revealed by ultraviolet-visible reflectance spectrophotometry. Proceedings of the Royal Society of London. Series B: Biological Sciences, 268(1467), 565-571. https://doi.org/10.1098/rspb.2000.1414Price-Waldman, R., & Stoddard, M. C. (2021). Avian Coloration Genetics: Recent Advances and Emerging Questions. Journal of Heredity, 112(5), 395-416. https://doi.org/10.1093/jhered/esab015Riehl, C. (2011). Paternal investment and the "sexually selected hypothesis" for the evolution of eggshell coloration: Revisiting the assumptions. Auk, 128(1), 175-179. https://doi.org/10.1525/auk.2011.10171Jagannath, A., Shore, R. F., Walker, L. A., Ferns, P. N., & Gosler, A. G. (2008). Eggshell pigmentation indicates pesticide contamination. Journal of Applied Ecology, 45(1), 133-140. https://doi.org/10.1111/j.1365-2664.2007.01386.xOrlowski, G., Niedzielski, P., Merta, D., Pokorny, P., & Proch, J. (2020). Quantifying the functional disparity in pigment spot-background egg colour ICP-OES-based eggshell ionome at two extremes of avian embryonic development. Scientific Reports, 10(1), 1-15. https://doi.org/10.1038/s41598-020-79040-4Hanley, D., Cassey, P., & Doucet, S. M. (2013). Parents, predators, parasites, and the evolution of eggshell colour in open nesting birds. Evolutionary Ecology, 27(3), 593-617. https://doi.org/10.1007/s10682-012-9619-6Dainson, M., Hauber, M. E., López, A. V., Grim, T., & Hanley, D. (2017). Does contrast between eggshell ground and spot coloration affect egg rejection?. The Science of Nature, 104, 1-9. https://doi.org/10.1007/s00114-017-1476-2Yang, C., Wang, J., and Liang, W. (2016). Blocking of ultraviolet reflectance on bird eggs reduces nest predation by aerial predators. Journal of Ornithology, 157: 43-47. https://doi.org/10.1007/s10336-015-1243-0Honza, M., Polaciková, L., & Procházka, P. (2007). Ultraviolet and green parts of the colour spectrum affect egg rejection in the song thrush (Turdus philomelos). Biological Journal of the Linnean Society, 92(2), 269-276. https://doi.org/10.1111/j.1095-8312.2007.00848.xLahti, D. C., & Ardia, D. R. (2016). Shedding light on bird egg color: Pigment as parasol and the dark car effect. The American Naturalist, 187(5), 547-563. https://doi.org/10.1086/685780Mayani-Parás, F., Kilner, R. M., Stoddard, M. C., Rodríguez, C., & Drummond, H. (2015). Behaviorally induced camouflage: a new mechanism of avian egg protection. The American Naturalist, 186(4), E91-E97. https://doi.org/10.1086/682579Hanley, D., Doucet, S. M., & Dearborn, D. C. (2010). A blackmail hypothesis for the evolution of conspicuous egg coloration in birds. The Auk, 127(2), 453-459. https://doi.org/10.1525/auk.2009.09090Soler, J. J., Moreno, J., Aviles, j., & Moller, A. P. (2005). Blue and green egg-color intensity is associated with parental: effort and mating system in passerines: support for: the sexual selection hypothesis. Evolution, 59(3), 636-644. https://doi.org/10.1525/auk.2009.09090Ladouce, M., Barakat, T., Su, B. L., Deparis, O., & Mouchet, S. R. (2020). Scattering of ultraviolet light by avian eggshells. Faraday Discussions, 223, 63-80. https://doi.org/10.1039/D0FD00034ECassey, P., Thomas, G. H., Portugal, S. J., Maurer, G., Hauber, M. E., Grim, T.Lovell, G,. Miksík, I. (2012). Why are birds' eggs colourful? Eggshell pigments co-vary with life-history and nesting ecology among British breeding non-passerine birds. Biological Journal of the Linnean Society, 106(3), 657-672. https://doi.org/10.1111/j.1095-8312.2012.01877.xHansell, M. (2000). Bird nests and construction behaviour. Cambridge University Press.Paul, N. D., & Gwynn-Jones, D. (2003). Ecological roles of solar UV radiation: towards an integrated approach. Trends in Ecology & Evolution, 18(1), 48-55. https://doi.org/10.1016/S0169-5347(02)00014-9Wang, J., Yang, C., Shi, H., & Liang, W. (2016). Reflectance and artificial nest experiments of reptile and bird eggs imply an adaptation of bird eggs against ultraviolet. Ecological research, 31(1), 105-110. https://doi.org/10.1007/s11284-015-1317-8Avilés, J. M., Soler, J. J., & Pérez-Contreras, T. (2006). Dark nests and egg colour in birds: A possible functional role of ultraviolet reflectance in egg detectability. Proceedings of the Royal Society B: Biological Sciences, 273(1603), 2821-2829. https://doi.org/10.1098/rspb.2006.3674Ödeen, A., Håstad, O., & Alström, P. (2011). Evolution of ultraviolet vision in the largest avian radiation-the passerines. BMC Evolutionary Biology, 11(1), 1-8. https://doi.org/10.1186/1471-2148-11-313Maurer, G., Portugal, S. J., & Cassey, P. (2011). An embryo's eye view of avian eggshell pigmentation. Journal of Avian Biology, 42(6), 494-504. https://doi.org/10.1111/j.1600-048X.2011.05368.xClements, J. F., T. S. Schulenberg, M. J. Iliff, T. A. Fredericks, J. A. Gerbracht, D. Lepage, S. M. Billerman, B. L. Sullivan, and C. L. Wood. (2021). The eBird/Clements checklist of Birds of the World: v2021. Downloaded from https://www.birds.cornell.edu/clementschecklist/download/R Core Team (2022). R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. URL https://www.R-project.org/.D'Alba, L., Torres, R., Waterhouse, G. I., Eliason, C., Hauber, M. E., & Shawkey, M. D. (2017). What does the eggshell cuticle do? A functional comparison of avian eggshell cuticles. Physiological and Biochemical Zoology, 90(5), 588-599. https://doi.org/10.1086/693434Maia, R., Eliason, C. M., Bitton, P. P., Doucet, S. M., & Shawkey, M. D. (2013). pavo: an R package for the analysis, visualization and organization of spectral data. Methods in Ecology and Evolution, 4(10), 906-913.Billerman, S. M., Keeney, B. K., Rodewald, P. G., & Schulenberg, T. S. (2020). Birds of the World. Cornell Laboratory of Ornithology, Ithaca, NY, USA.Simón, J. E., & Pacheco, S. (2005). On the standardization of nest descriptions of neotropical birds. Revista Brasileira de ornitologia, 13(2), 143-154.Tobias, J. A. Sheard, C., Pigot, A. L., Devenish, A. J. M., Yang, J., Sayol, F., Neate-Clegg, M. H. C., Alioravainen, N., Weeks, T. L., Barber, R. A., Walkden, P. A., MacGregor, H. E. A., Jones, S. E. I., Vincent, C., Phillips, A. G., Marples, N. M., Montaño-Centellas, F. A., Leandro-Silva, V., Claramunt, S,Schleuning, M. (2022). AVONET: Morphological, ecological and geographical data for all birds. Ecology Letters, 25, 581-597. https://doi.org/10.1111/ele.13898Englert Duursma, D., Gallagher, R. V., Price, J. J., & Griffith, S. C. (2018). Variation in avian egg shape and nest structure is explained by climatic conditions. Scientific Reports, 8(1), 1-10. https://doi.org/10.1038/s41598-018-22436-0Heenan, C. B. (2013). An overview of the factors influencing the morphology and thermal properties of avian nests. Avian Biology Research, 6(2), 104-118. https://doi.org/10.3184/003685013X13614670646299Gómez, J., Ramo, C., Stevens, M., Liñán-Cembrano, G., Rendón, M. A., Troscianko, J. T., & Amat, J. A. (2018). Latitudinal variation in biophysical characteristics of avian eggshells to cope with differential effects of solar radiation. Ecology and Evolution, 8(16), 8019-8029. https://doi.org/10.1002/ece3.4335Cherry, M. I., & Gosler, A. G. (2010). Avian eggshell coloration: new perspectives on adaptive explanations. Biological Journal of the Linnean Society, 100(4), 753-762. https://doi.org/10.1111/j.1095-8312.2010.01457.xKilner, R. M. (2006). The evolution of egg colour and patterning in birds. Biological Reviews, 81(3), 383-406. https://doi.org/10.1017/S1464793106007044Wegrzyn, E., Leniowski, K., Rykowska, I., & Wasiak, W. (2011). Is UV and blue-green egg colouration a signal in cavity-nesting birds?. Ethology Ecology & Evolution, 23(2), 121-139. https://doi.org/10.1080/03949370.2011.554882Maurer, G., Portugal, S. J., Hauber, M. E., Miksík, I., Russell, D. G., & Cassey, P. (2015). First light for avian embryos: eggshell thickness and pigmentation mediate variation in development and UV exposure in wild bird eggs. Functional Ecology, 29(2), 209-218. https://doi.org/10.1111/1365-2435.12314Schneider, C. A., Rasband, W. S., & Eliceiri, K. W. (2012). NIH Image to ImageJ: 25 years of image analysis. Nature Methods, 9(7), 671-675. doi:10.1038/nmeth.2089Nagy, J., Hauber, M. E., Hartley, I. R., & Mainwaring, M. C. (2019). Correlated evolution of nest and egg characteristics in birds. Animal Behaviour, 158, 211-225. https://doi.org/10.1016/j.anbehav.2019.10.015Stoddard, M. C., Yong, E. H., Akkaynak, D., Sheard, C., Tobias, J. A., & Mahadevan, L. (2017). Avian egg shape: Form, function, and evolution. Science, 356(6344), 1249-1254. DOI: 10.1126/science.aaj1945Jetz, W., Thomas, G. H., Joy, J. B., Hartmann, K., & Mooers, A. O. (2012). The global diversity of birds in space and time. Nature, 491(7424), 444-448. https://doi.org/10.1038/nature11631Ocampo, D., De Silva, T.N., Sheard, C., Stoddard, M.C. (2023). Evolution of nest architecture in tyrant flycatchers and allies. Philosophical Transactions B. https://doi.org/10.1098/rstb.2022.0148Bouckaert, R., Drummond, A., Rambaut, A., Suchard, M., Vaughan, T., & Heled, Y. (2014). BEAST2: Bayesian evolutionary analysis sampling trees.Revell, L. J. (2012). Phytools: An R package for phylogenetic comparative biology (and other things). Methods in Ecology and Evolution, 3(2), 217-223Revell, L. J. (2013). Two new graphical methods for mapping trait evolution on phylogenies. Methods in Ecology and Evolution, 4(8), 754-759. https://doi.org/10.1111/2041-210X.12066Orme, D., Freckleton, R., Thomas, G. Petzoldt, T., Fritz, S., Isaac, N., & Pearse, W. (2018). caper: Comparative Analyses of Phylogenetics and Evolution in R. R package version 1.0.1Burnham, K. P., Anderson, D. R., & Huyvaert, K. P. (2011). AIC model selection and multimodel inference in behavioral ecology: Some background, observations, and comparisons. Behavioral Ecology and Sociobiology, 65(1), 23-35. https://doi.org/10.1007/s00265-010-1029-6Barton, K. (2019). MuMIn: Multi-model inference. R package version 1.43.15. Retrieved from http://r-forge.r-project.org/projects/mumin/.Burnham, K. P., & Anderson, D. R. (2002). A practical information-theoretic approach. In Model selection and multimodel inference (2nd ed., p. 2). Springer.D'Alba, L., Goldenberg, J., Nallapaneni, A., Parkinson, D. Y., Zhu, C., Vanthournout, B., & Shawkey, M. D. (2021). Evolution of eggshell structure in relation to nesting ecology in non-avian reptiles. Journal of morphology, 282(7), 1066-1079. https://doi.org/10.1002/jmor.21347Wisocki, P. A., Kennelly, P., Rojas Rivera, I., Cassey, P., Burkey, M. L., & Hanley, D. (2020). The global distribution of avian eggshell colours suggest a thermoregulatory benefit of darker pigmentation. Nature Ecology & Evolution, 4(1), 148-155. https://doi.org/10.1038/s41559-019-1003-2Hanley, D., Grim, T., Cassey, P., & Hauber, M. E. (2015). Not so colourful after all: eggshell pigments constrain avian eggshell colour space. Biology letters, 11(5), 20150087. https://doi.org/10.1098/rsbl.2015.0087Lee, W. S., Kwon, Y. S., & Yoo, J. C. (2010). Egg survival is related to the colour matching of eggs to nest background in Black-tailed Gulls. Journal of Ornithology, 151, 765-770. https://doi.org/10.1007/s10336-010-0508-xYang, C., Møller, A. P., & Liang, W. (2022). Light matters: Nest illumination alters egg rejection behavior in a cavity-nesting bird. Avian Research, 13, 100016. https://doi.org/10.1016/j.avrs.2022.100016Maziarz, M., & Wesolowski, T. (2014). Does darkness limit the use of tree cavities for nesting by birds?. Journal of Ornithology, 155, 793-799. https://doi.org/10.1007/s10336-014-1069-1Moreno, J., & Osorno, J. L. (2003). Avian egg colour and sexual selection: does eggshell pigmentation reflect female condition and genetic quality?. Ecology Letters, 6(9), 803-806. https://doi.org/10.1046/j.1461-0248.2003.00505.xLahti, D. C. (2008). Population differentiation and rapid evolution of egg color in accordance with solar radiation. The Auk, 125(4), 796-802. https://doi.org/10.1525/auk.2008.07033L'Herpiniere, K. L., Tims, A. R., Englert Duursma, D., & Griffith, S. C. (2021). The evolution of egg colour and patterning in Australian songbirds. Evolution, 75(12), 3132-3141. https://doi.org/10.1111/evo.14375Gosler, A. G., Higham, J. P., & James Reynolds, S. (2005). Why are birds' eggs speckled?. Ecology Letters, 8(10), 1105-1113. https://doi.org/10.1111/j.1461-0248.2005.00816.xCassey, P., Thomas, G. H., Portugal, S. J., Maurer, G., Hauber, M. E., Grim, T., & Miksík, I. (2012). Why are birds' eggs colourful? Eggshell pigments co-vary with life-history and nesting ecology among British breeding non-passerine birds. Biological Journal of the Linnean Society, 106(3), 657-672. https://doi.org/10.1111/j.1095-8312.2012.01877.xYang, C., Wang, L., Hsu, Y.-C., Antonov, A., Moksnes, A., Røskaft, E., Liang, W. & Stokke, B. G. (2013) UV reflectance as a cue in egg discrimination in two Prinia species exploited differently by brood parasites in Taiwan. The Ibis 155, 571-575. https://doi.org/10.1111/ibi.12043Honza, M., Capek, M., Mikulica, O., & Samas, P. (2022). Ultraviolet coloration of avian parasitic egg does not cue egg rejection in the common redstart host. Journal of Ornithology, 163(4), 903-909. https://doi.org/10.1007/s10336-022-01991-4Ericson PG, Anderson CL, Britton T, Elzanowski A, Johansson US et al., (2006). Diversification of Neoaves: Integration of molecular sequence data and fossils. Biol. Lett. 2: 543-547. https://doi.org/10.1098/rsbl.2006.0523Gómez, J., Pereira, A. I., Pérez-Hurtado, A., Castro, M., Ramo, C., & Amat, J. A. (2016). A trade-off between overheating and camouflage on shorebird eggshell colouration. Journal of Avian Biology, 47(3), 346-353. https://doi.org/10.1111/jav.00736Kinoshita, S., Yoshioka, S., & Miyazaki, J. (2008). Physics of structural colors. Reports on Progress in Physics, 71(7), 076401. DOI 10.1088/0034-4885/71/7/076401Honza, M., Sulc, M., & Cherry, M. I. (2014). Does nest luminosity play a role in recognition of parasitic eggs in domed nests? A case study of the red bishop. Naturwissenschaften, 101, 1009-1015. https://doi.org/10.1007/s00114-014-1240-9Xiao, H., Hu, Y., Lang, Z., Fang, B., Guo, W., Zhang, Q. I et al (2017). How much do we know about the breeding biology of bird species in the world?. Journal of Avian Biology, 48(4), 513-518. https://doi.org/10.1111/jav.00934Wiemann, J., Yang, T. R., & Norell, M. A. (2018). Dinosaur egg colour had a single evolutionary origin. Nature, 563(7732), 555-558. https://doi.org/10.1038/s41586-018-0646-5Marki, P. Z., Fabre, P. H., Jønsson, K. A., Rahbek, C., Fjeldså, J., & Kennedy, J. D. (2015). Breeding system evolution influenced the geographic expansion and diversification of the core Corvoidea (Aves: Passeriformes). Evolution, 69(7), 1874-1924. https://doi.org/10.1111/evo.12695Carvalho, C. B., Macedo, R. H., & Graves, J. A. (2006). Breeding strategies of a socially monogamous neotropical passerine: extra-pair fertilizations, behavior, and morphology. 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