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Professor Camille Bonneaud

Professor Camille Bonneaud

Director of the Environment and Sustainability Institute and Professor of Evolutionary Biology

 01326 259474

 Environment and Sustainability Institute 00.12


Environment and Sustainability Institute, University of Exeter, Penryn Campus, Penryn, Cornwall, TR10 9FE, UK


I am interested in understanding the emergence and spread of infectious diseases across wildlife and agricultural systems, as well as how to develop epidemic preparedness programmes that engage local communities to more effectively protect against animal diseases. To this end, I use a range of model systems. For example, I investigate how a pathogen of poultry jumped and evolved in a wild North American songbird, killing millions. Additionally, I study the factors driving the emergence of avian pox across the Galápagos archipelago, of leprosy in endangered West African chimpanzees, of phocine distemper in vulnerable harbour seals of the UK, as well as test the role of herring gulls in spreading anti-microbial resistant bacteria across UK landscapes. 

I am a member of the Evolution group. For further details, please see Research.


PhD., University Pierre and Marie Curie, Paris (France)
M.Sc., University Pierre and Marie Curie, Paris (France)
B.Sc. (License), University Denis Diderot, Paris (France)


EU Marie-Curie & CNRS Research Fellow, Station d’Écologie Théorique et Expérimentale (France)
Postdoctoral researcher, Department of Organismic and Evolutionary Biology, Harvard U. (USA)
Postdoctoral researcher, Center for Tropical Research, U. California, Los Angeles (USA)

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Research interests

My research interests broadly cover host-pathogen interactions in natural populations, from the mechanistic underpinnings of the co-evolutionary process, through to the conservation implications of infectious outbreaks and how to prevent them. I am particularly interested in understanding how emerging infectious pathogens jump into new host species and how they subsequently adapt to these novel host environments, as well as how host resistance/tolerance evolves following epidemic outbreak.

Research projects

Examples of projects

1. Rapid host-pathogen co-evolution following an emerging disease outbreak: Mid-1990s, Mycoplasma gallisepticum, a common bacterial pathogen of poultry, jumped into a wild North American songbird, the house finch (Haemorhous mexicanus), triggering a devastating epidemic that killed millions of birds. This epidemic is not only of one of the best documented emerging infectious outbreaks in the wild, but it is also one that was left to evolve naturally without any human intervention. As such, it offers a unique opportunity to understand how pathogens jump into new hosts, how they adapt to these novel hosts and how hosts evolve resistance and/or tolerance. We combine experimental work with molecular, immunological and microbiological approaches to study host and pathogen co-evolution over the course of this epidemic. 

Main collaborators on this project:
Prof Geoff E. Hill (Auburn University, USA)
Prof Kevin McGraw (Arizona State University, USA)
Prof Alastair Wilson (University of Exeter, UK) 
Prof Pascal Sirand-Pugnet (INRAE Bordeaux, France)
Dr Lucy Weinert (University of Cambridge, UK)
Dr Cedric Berger (Cardiff University, UK)

2. Climate change and wildlife pathogens - insights from an altitudinal gradient​: Emerging diseases are expected to increase under climate change as pathogens colonise new areas and infect new host populations. Empirical tests of these changes, however, are difficult to execute without confounding influences. We examine how climate change might affect the interaction between malaria parasites and their avian hosts in the French Pyrenees, where altitude (and hence climate), but not habitat or day length, varies significantly over short distances. We compare the prevalence, species richness and transmission seasonality of malaria parasites in four closely monitored nest-box populations of great tits (Parus major) and blue tits (Cyanistes caeruleus); these populations are located over short geographical distances (4-16km) and at various altitudes (400-1600m) of the Pyrenees Mountains. 

Main collaborators on this project:
Dr Alexis Chaine (Station d'Ecologie Experimentale de Moulis, CNRS, France)
​Prof Andy Russell (University of Exeter, UK)
Dr Joshua Lynton-Jenkins (Animal & Plant Health Agency, UK)

3. Infectious disease introduction to a remote archipelago: Populations introduced to novel pathogens are often the most vulnerable and can suffer a high cost. Such was the case when Plasmodium relictum was introduced to Hawaii, contributing to the extinction of dozens of endemic bird species. We aim to understand how pathogens become established in novel populations by studying their introduction to Darwin’s finch species in the Galápagos. This isolated archipelago has evolved a unique avifauna. However, over the past 200 years humans have facilitated the introduction of two key pathogens; avian pox (Avipoxvirus) and avian malaria (Haemosporidia). By screening for these diseases, we aim to determine their prevalence, distribution, and potential costs in these naive hosts. This effort will help reveal emerging threats to iconic bird species within a unique UNESCO world heritage site.

Main collaborators on this project:
Prof Jaime Chaves (San Francisco State University, USA/University San Francisco de Quito, Ecuador) 
Dr Joshua Lynton-Jenkins (Animal & Plant Health Agency, UK)

4. Barcoding Galapagos: Nearly 200 years ago, the unique biota of Galapagos inspired one of the greatest scientific revolutions in history – Charles Darwin’s theory of evolution by natural selection. Today, this Natural World Heritage site (est.1976) and UNESCO Biosphere Reserve (est.1984) also inspires pioneering models of sustainability, conservation and eco-tourism. Such models are celebrated for their long-term solutions to existing tensions between the preservation of biodiversity and the social-economic well-being of local inhabitants. However, the Covid-19 pandemic has revealed their vulnerability to short-term perturbation. We trained and employed >80 naturalist guides to catalogue the biodiversity of Galapagos. This ‘Barcode of Life’ project will ensure that: (1) the genetic profile of Galapagos is documented so that the direct and indirect impacts of environmental perturbations can be quantified; and (2) naturalist guides receive immediate capacity building employment. Our aim is to help guide future attempts at using locally-driven research to improve the socio-economic well-being and resilience of key workers in the eco-tourism industry.

Read more:

Main collaborator on this project:
Prof Jaime Chaves (San Francisco State University, USA/University San Francisco de Quito, Ecuador)
Prof Diana Pazmino (University San Francisco de Quito, Ecuador)
Prof Carlos Mena  (University San Francisco de Quito, Ecuador)
Prof Andy Russell (University of Exeter, UK)
Dr Thomas Chaigneau (University of Exeter, UK)
Galapagos Conservation Trust
Agencia de Bioseguridad Galapagos

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Journal articles

Lynton-Jenkins J, Brundl A, Cauchoix M, Lejeune L, Salle L, Thiney A, Russell A, Chaine A, Bonneaud C (In Press). Contrasting the seasonal and elevational prevalence of generalist avian haemosporidia in co-occurring host species. Ecology and Evolution
Bonneaud C, Herrel A, Videlier M (In Press). Intra-individual variation in exploration behaviour in a largely aquatic frog: effects of sex and personality traits. Herpetological Journal
Dowling A, Hill G, Bonneaud C (In Press). Multiple differences in pathogen-host cell interactions following a bacterial host shift. Scientific Reports
Bonneaud C, Giraudeau M, Tardy L, Staley M, Hill GE, McGraw KJ (In Press). Rapid antagonistic coevolution in an emerging pathogen and its vertebrate host. Current Biology, 1-25.
Bonneaud C, Lucy W, Bram K (In Press). Understanding the emergence of bacterial pathogens in novel hosts. Philosophical Transactions B: Biological Sciences
Ferraguti M, Magallanes S, Jiménez‐Peñuela J, Martínez‐de la Puente J, Garcia‐Longoria L, Figuerola J, Muriel J, Albayrak T, Bensch S, Bonneaud C, et al (2023). Environmental, geographical and time‐related impacts on avian malaria infections in native and introduced populations of house sparrows (<i>Passer domesticus</i>), a globally invasive species. Global Ecology and Biogeography, 32(5), 809-823. Abstract.
San-Jose LM, Bestion E, Pellerin F, Richard M, Di Gesu L, Salmona J, Winandy L, Legrand D, Bonneaud C, Guillaume O, et al (2023). Investigating the genetic basis of vertebrate dispersal combining RNA-seq, RAD-seq and quantitative genetics. Mol Ecol, 32(12), 3060-3075. Abstract.  Author URL.
Lynton‐Jenkins JG, Chaine AS, Russell AF, Bonneaud C (2023). Parasite detection and quantification in avian blood is dependent on storage medium and duration. Ecology and Evolution, 13(2). Abstract.
Lynton-Jenkins JG, Russell AF, Chaves J, Bonneaud C (2021). Avian disease surveillance on the island of San Cristobal, Galapagos. ECOLOGY AND EVOLUTION, 11(24), 18422-18433.  Author URL.
Bersacola E, Parathian H, Frazão-Moreira A, Jaló M, Sanhá A, Regalla A, Saíd AR, Quecuta Q, Camará ST, Quade SMFF, et al (2021). Developing an Evidence-Based Coexistence Strategy to Promote Human and Wildlife Health in a Biodiverse Agroforest Landscape. Frontiers in Conservation Science, 2 Abstract.
Zhang Y, Hill GE, Ge Z, Park NR, Taylor HA, Andreasen V, Tardy L, Kavazis AN, Bonneaud C, Hood WR, et al (2021). Effects of a Bacterial Infection on Mitochondrial Function and Oxidative Stress in a Songbird. Physiol Biochem Zool, 94(2), 71-82. Abstract.  Author URL.
Hockings KJ, Mubemba B, Avanzi C, Pleh K, Düx A, Bersacola E, Bessa J, Ramon M, Metzger S, Patrono LV, et al (2021). Leprosy in wild chimpanzees. Nature, 598(7882), 652-656. Abstract.
Gates D, Staley M, Tardy L, Giraudeau M, Hill G, McGraw K, Bonneaud C (2021). Levels of pathogen virulence and host resistance both shape the antibody response to an emerging bacterial disease. Scientific Reports, 11
Dobreva MP, Lynton-Jenkins JG, Chaves JA, Tokita M, Bonneaud C, Abzhanov A (2021). Sex identification in embryos and adults of Darwin’s finches. PLOS ONE, 16(3), e0237687-e0237687. Abstract.
Marden E, Abbott RJ, Austerlitz F, Ortiz-Barrientos D, Baucom RS, Bongaerts P, Bonin A, Bonneaud C, Browne L, Alex Buerkle C, et al (2021). Sharing and reporting benefits from biodiversity research. Mol Ecol, 30(5), 1103-1107.  Author URL.
Ducret V, Richards AJ, Videlier M, Scalvenzi T, Moore KA, Paszkiewicz K, Bonneaud C, Pollet N, Herrel A (2021). Transcriptomic analysis of the trade-off between endurance and burst-performance in the frog Xenopus allofraseri. BMC Genomics, 22(1). Abstract.  Author URL.
Ducret V, Videlier M, Moureaux C, Bonneaud C, Herrel A (2020). Do female frogs have higher resting metabolic rates than males? a case study with <i>Xenopus allofraseri</i>. Journal of Zoology, 312(4), 221-226. Abstract.
Bonneaud C, Longdon B (2020). Emerging pathogen evolution. EMBO reports, 21(9).
Bonneaud C, Tardy L, Hill GE, McGraw KJ, Wilson AJ, Giraudeau M (2020). Experimental evidence for stabilizing selection on virulence in a bacterial pathogen. Evolution Letters, 4(6), 491-501. Abstract.
Hockings KJ, Mubemba B, Avanzi C, Pleh K, Düx A, Bersacola E, Bessa J, Ramon M, Metzger S, Patrono LV, et al (2020). Leprosy in wild chimpanzees. Abstract.
Dobreva MP, Lynton-Jenkins JG, Chaves JA, Tokita M, Bonneaud C, Abzhanov A (2020). Sex identification in embryos and adults of Darwin’s finches. Abstract.
Padilla P, Ducret V, Bonneaud C, Courant J, Herrel A (2019). Acclimation temperature effects on locomotor traits in adult aquatic anurans (X. tropicalis and X. laevis) from different latitudes: possible implications for climate change. Conserv Physiol, 7(1). Abstract.  Author URL.
Tardy L, Giraudeau M, Hill GE, McGraw KJ, Bonneaud C (2019). Contrasting evolution of virulence and replication rate in an emerging bacterial pathogen. Proc Natl Acad Sci U S A, 116(34), 16927-16932. Abstract.  Author URL.
Bonneaud C, Tardy L, Giraudeau M, Hill GE, McGraw KJ, Wilson AJ (2019). Evolution of both host resistance and tolerance to an emerging bacterial pathogen. EVOLUTION LETTERS, 3(5), 544-554.  Author URL.
Giraudeau M, Heidinger B, Bonneaud C, Sepp T (2019). Telomere shortening as a mechanism of long-term cost of infectious diseases in natural animal populations. Biol Lett, 15(5). Abstract.  Author URL.
Staley M, Hill GE, Josefson CC, Armbruster JW, Bonneaud C (2018). Bacterial Pathogen Emergence Requires More than Direct Contact with a Novel Passerine Host. Infect Immun, 86(3). Abstract.  Author URL.
Staley M, Bonneaud C, McGraw KJ, Vleck CM, Hill GE (2018). Detection of Mycoplasma gallisepticum in House Finches ( Haemorhous mexicanus) from Arizona. Avian Dis, 62(1), 14-17. Abstract.  Author URL.
Gates DE, Valletta JJ, Bonneaud C, Recker M (2018). Quantitative host resistance drives the evolution of increased virulence in an emerging pathogen. J Evol Biol, 31(11), 1704-1714. Abstract.  Author URL.
Bonneaud C, Staley M, McGraw K, Vleck C, Hill G (2017). Detection of Mycoplasma gallisepticum in House Finches (Haemorhous mexicanus) from Arizona. Avian Diseases
Bonneaud C, Sepil I, Wilfert L, Calsbeek R (2017). Plasmodium Infections in Natural Populations of Anolis sagrei Reflect Tolerance Rather Than Susceptibility. Integrative and Comparative Biology, 57(2), 352-361.
Bonneaud C, Wilson R, Seebacher F (2016). Immune-challenged fish up-regulate their metabolic scope to support locomotion. PLoS One
Bonneaud C, Marnocha E, Herrel A, Vanhooydonck B, Irschick DJ, Smith TB (2015). Developmental plasticity affects sexual size dimorphism in an anole lizard. Functional Ecology, 30(2), 235-243. Abstract.
Staley M, Bonneaud C (2015). Immune responses of wild birds to emerging infectious diseases. Parasite Immunol, 37(5), 242-254. Abstract.  Author URL.
Balenger SL, Bonneaud C, Sefick SA, Edwards SV, Hill GE (2015). Plumage color and pathogen-induced gene expression in a wild bird. BEHAVIORAL ECOLOGY, 26(4), 1100-1110.  Author URL.
Videlier M, Cornette R, Bonneaud C, Herrel A (2015). Sexual differences in exploration behavior in Xenopus tropicalis?. J Exp Biol, 218(Pt 11), 1733-1739. Abstract.  Author URL.
Richards AJ, Herrel A, Bonneaud C (2015). htsint: a Python library for sequencing pipelines that combines data through gene set generation. BMC Bioinformatics, 16 Abstract.  Author URL.
Videlier M, Bonneaud C, Cornette R, Herrel A (2014). Exploration syndromes in the frog <i>Xenopus</i> (<i>Silurana</i>) <i>tropicalis</i>: correlations with morphology and performance?. JOURNAL OF ZOOLOGY, 294(3), 206-213.  Author URL.
Careau V, Biro PA, Bonneaud C, Fokam EB, Herrel A (2014). Individual variation in thermal performance curves: swimming burst speed and jumping endurance in wild-caught tropical clawed frogs. Oecologia, 175(2), 471-480. Abstract.  Author URL.
Herrel A, Vasilopoulou-Kampitsi M, Bonneaud C (2014). Jumping performance in the highly aquatic frog, Xenopus tropicalis: sex-specific relationships between morphology and performance. PeerJ, 2 Abstract.  Author URL.
Cheatsazan H, de Almedia APLG, Russell AF, Bonneaud C (2013). Experimental evidence for a cost of resistance to the fungal pathogen, Batrachochytrium dendrobatidis, for the palmate newt, Lissotriton helveticus. BMC Ecol, 13 Abstract.  Author URL.
Richards AJ, Kosinski AS, Bonneaud C, Legrand D, Owzark K (2013). lpEdit: an editor to facilitate reproducible analysis via literate programming. Proceedings of the 9th Python in Science conference, 80-84.
Ewen JG, Bensch S, Blackburn TM, Bonneaud C, Brown R, Cassey P, Clarke RH, Pérez-Tris J (2012). Establishment of exotic parasites: the origins and characteristics of an avian malaria community in an isolated island avifauna. Ecology Letters, 15(10), 1112-1119. Abstract.
Ewen JG, Bensch S, Blackburn TM, Bonneaud C, Brown R, Cassey P, Clarke RH, Pérez-Tris J (2012). Establishment of exotic parasites: the origins and characteristics of an avian malaria community in an isolated island avifauna. Ecol Lett, 15(10), 1112-1119. Abstract.  Author URL.
Bonneaud C, Balenger SL, Hill GE, Russell AF (2012). Experimental evidence for distinct costs of pathogenesis and immunity against a natural pathogen in a wild bird. Molecular Ecology, 21(19), 4787-4796. Abstract.
Bonneaud C, Balenger SL, Hill GE, Russell AF (2012). Experimental evidence for distinct costs of pathogenesis and immunity against a natural pathogen in a wild bird. Mol Ecol, 21(19), 4787-4796. Abstract.  Author URL.
Bonneaud C, Balenger SL, Zhang J, Edwards SV, Hill GE (2012). Innate immunity and the evolution of resistance to an emerging infectious disease in a wild bird. Mol Ecol, 21(11), 2628-2639. Abstract.  Author URL.
Herrel A, Gonwouo L, Fokam E, Ngundy W, Bonneaud C (2012). Inter-sexual differences in body shape and locomotor performance in the aquatic frog, Xenopus tropicalis. Journal of Zoology(287), 311-316.
Herrel A, Bonneaud C (2012). Temperature dependence of locomotor performance in the tropical clawed frog, Xenopus tropicalis. J Exp Biol, 215(Pt 14), 2465-2470. Abstract.  Author URL.
Herrel A, Bonneaud C (2012). Trade-offs between burst performance and maximal exertion capacity in a wild amphibian, Xenopus tropicalis. J Exp Biol, 215(Pt 17), 3106-3111. Abstract.  Author URL.
Delaney NF, Balenger S, Bonneaud C, Marx CJ, Hill GE, Ferguson-Noel N, Tsai P, Rodrigo A, Edwards SV (2012). Ultrafast evolution and loss of CRISPRs following a host shift in a novel wildlife pathogen, Mycoplasma gallisepticum. PLoS Genet, 8(2). Abstract.  Author URL.
James RS, Tallis J, Herrel A, Bonneaud C (2012). Warmer is better: thermal sensitivity of both maximal and sustained power output in the iliotibialis muscle isolated from adult Xenopus tropicalis. J Exp Biol, 215(Pt 3), 552-558. Abstract.  Author URL.
Marzal A, Ricklefs RE, Valkiūnas G, Albayrak T, Arriero E, Bonneaud C, Czirják GA, Ewen J, Hellgren O, Hořáková D, et al (2011). Diversity, loss, and gain of malaria parasites in a globally invasive bird. PLoS One, 6(7). Abstract.  Author URL.
Njabo KY, Cornel AJ, Bonneaud C, Toffelmier E, Sehgal RNM, Valkiūnas G, Russell AF, Smith TB (2011). Nonspecific patterns of vector, host and avian malaria parasite associations in a central African rainforest. Mol Ecol, 20(5), 1049-1061. Abstract.  Author URL.
Bonneaud C, Balenger SL, Russell AF, Zhang J, Hill GE, Edwards SV (2011). Rapid evolution of disease resistance is accompanied by functional changes in gene expression in a wild bird. Proc Natl Acad Sci U S A, 108(19), 7866-7871. Abstract.  Author URL.
Sehgal RNM, Buermann W, Harrigan RJ, Bonneaud C, Loiseau C, Chasar A, Sepil I, Valkiūnas G, Iezhova T, Saatchi S, et al (2011). Spatially explicit predictions of blood parasites in a widely distributed African rainforest bird. Proc Biol Sci, 278(1708), 1025-1033. Abstract.  Author URL.
Bonneaud C, Sinsheimer JS, Richard M, Chastel O, Sorci G (2009). Mhc polymorphisms fail to explain the heritability of phytohaemagglutinin-induced skin swelling in a wild passerine. Biol Lett, 5(6), 784-787. Abstract.  Author URL.
Bonneaud C, Sepil I, Mila B, Buermann W, Pollinger J, Sehgal R, Valkiunas G, Iezhova TA, Saatchi S, Smith TB, et al (2009). The prevalence of avian Plasmodium is higher in undisturbed tropical forests of Cameroon. Journal of Tropical Ecology(25), 439-447.
Calsbeek R, Bonneaud C, Smith TB (2008). Differential fitness effects of immunocompetence and neighbourhood density in alternative female lizard morphs. J Anim Ecol, 77(1), 103-109. Abstract.  Author URL.
Bonneaud C, Burnside J, Edwards SV (2008). High-speed developments in avian genomics. Bioscience(58), 587-595.
Calsbeek R, Bonneaud C (2008). Postcopulatory fertilization bias as a form of cryptic sexual selection. Evolution, 62(5), 1137-1148. Abstract.  Author URL.
Calsbeek R, Bonneaud C, Pradhu S, Manoukis N, Smith TB (2007). Multiple paternity and sperm storage lead to increased genetic diversity in Anolis lizard. Evolutionary Ecology Research(9), 495-503.
Pérez-Tris J, Hellgren O, Krizanauskiene A, Waldenström J, Secondi J, Bonneaud C, Fjeldså J, Hasselquist D, Bensch S (2007). Within-host speciation of malaria parasites. PLoS One, 2(2). Abstract.  Author URL.
Bonneaud C, Chastel O, Federici P, Westerdahl H, Sorci G (2006). Complex Mhc-based mate choice in a wild passerine. Proc Biol Sci, 273(1590), 1111-1116. Abstract.  Author URL.
Bonneaud C, Pérez-Tris J, Federici P, Chastel O, Sorci G (2006). Major histocompatibility alleles associated with local resistance to malaria in a passerine. EVOLUTION, 60(2), 383-389.  Author URL.
Bonneaud C, Pérez-Tris J, Federici P, Chastel O, Sorci G (2006). Major histocompatibility alleles associated with local resistance to malaria in a passerine. Evolution, 60(2), 383-389. Abstract.  Author URL.
Bonneaud C, Richard M, Faivre B, Westerdahl H, Sorci G (2005). An Mhc class I allele associated to the expression of T-dependent immune response in the house sparrow. Immunogenetics, 57(10), 782-789. Abstract.  Author URL.
Bonneaud C, Sorci G, Morin V, Westerdahl H, Zoorob R, Wittzell H (2004). Diversity of Mhc class I and IIB genes in house sparrows (Passer domesticus). Immunogenetics, 55(12), 855-865. Abstract.  Author URL.
Bonneaud C, Mazuc J, Chastel O, Westerdahl H, Sorci G (2004). Terminal investment induced by immune challenge and fitness traits associated with major histocompatibility complex in the house sparrow. Evolution, 58(12), 2823-2830. Abstract.  Author URL.
Bonneaud C, Mazuc J, Gonzalez G, Haussy C, Chastel O, Faivre B, Sorci G (2003). Assessing the cost of mounting an immune response. Am Nat, 161(3), 367-379. Abstract.  Author URL.
Mazuc J, Bonneaud C, Chastel O, Sorci G (2003). Maternal testosterone in eggs and social context in the house sparrow (Passer domesticus). Ecology Letters(6), 1084-1090.
Mazuc J, Bonneaud C, Chastel O, Sorci G (2003). Social environment affects female and egg testosterone levels in the house sparrow (<i>Passer domesticus</i>). ECOLOGY LETTERS, 6(12), 1084-1090.  Author URL.


Zhang Y, Hill GE, Ge Z, Park N, Taylor H, Andreasen V, Kavazis AN, Bonneaud C, Hood WR (2019). Effects of Mycoplasma gallisepticum on mitochondrial function and oxidative stress in house finch.  Author URL.
Bonneaud C, Richards A, Herrel A, Seebacher F, Wilson R (2017). Using multi-level transcriptomics and metabolic measures to investigate the trade-off between performance and immunity.  Author URL.
Staley M, Bonneaud C, Hill GE (2016). Insights into the role of host access in limiting disease emergence in a natural host-pathogen system.  Author URL.
Staley M, Bonneaud C, Giradeau M, Mcgraw KJ, Hill GE (2015). The epidemiology of Mycoplasma gallisepticum infections in Arizona house finches (Haemorhous mexicanus).  Author URL.
Staley M, Bonneaud C, Hill GE (2014). Are more ornamented birds better at combatting disease? Assessing the links between condition and ornamentation.  Author URL.
Herrel A, Videlier M, Cornette R, Bonneaud C (2014). Should I stay or should I go: exploration behavior in the frog Xenopus tropicalis.  Author URL.

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