Guilhem Doulcier

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My portrait, thanks to Charlotte R. I'm an Evolutionary Biologist. I'm passionate about the interplays between ecological and evolutionary dynamics.

Current affiliation: Theory and Method in Biosciences (Macquarie University, Sydney, Australia), and Theoretical Biology Department (Max Planck Institute, Plön, Germany).



  1. Stability of Ecologically Scaffolded Traits During Evolutionary Transitions in Individuality. Guilhem Doulcier, Peter Takacs, Katrin Hammerschmidt, Pierrick Bourrat. BioRxiv, 2023. Code
  2. Neutral Diversity in Experimental Metapopulations. Guilhem Doulcier, Amaury Lambert. arxiv, 2023. Code
  3. Individuality Through Ecology. Pierrick Bourrat, Peter Takacs, Guilhem Doulcier, Matthew C. Nitschke, Andrew J. Black, Katrin Hammerschmidt, Paul B. Rainey. OSF, 2023.


  1. Evolutionary Transitions in Individuality and Life Cycle Closure. Guilhem Doulcier, Peter Takacs, Pierrick Bourrat. Philosophy of Science, 2023.
  2. From Fitness-Centered to Trait-Centered Explanations: What Evolutionary Transitions in Individuality Teach Us About Fitness. Peter Takacs, Guilhem Doulcier, Pierrick Bourrat. Philosophy of Science, 2023.
  3. Rapid dissemination of host-metabolism-manipulating transposon-like entities via integrative and conjugative elements. Elena Colombi, Frederic Bertels, Guilhem Doulcier, Ellen McConnell, Tatyana Pichugina, Kee Hoon Sohn, Christina Straub, Honour McCann, Paul B. Rainey. PNAS (in print), 2023.
  4. Tradeoff breaking model of evolutionary transitions in individuality and the limits of the fitness-decoupling metaphor. Pierrick Bourrat*, Guilhem Doulcier*, Caroline J. Rose, Paul B. Rainey, Katrin Hammerschmidt. eLife, 2022. Blog Preprint Code PDF
  5. Life history tradeoffs, division of labor and evolutionary transitions in individuality. Guilhem Doulcier, Katrin Hammerschmidt, Pierrick Bourrat. in The Evolution of Multicellularity, CRC Press, 2022. Preprint Code
  6. A leader cell triggers end of lag phase in populations of Pseudomonas fluorescens. Maxime Ardré, Guilhem Doulcier, Naama Brenner, Paul B. Rainey. microLife, 2022.
  7. Eco-evolutionary dynamics of nested Darwinian populations and the emergence of community-level heredity. Guilhem Doulcier, Amaury Lambert, Silvia De Monte, Paul B. Rainey. eLife, 2020. Code
  8. Taming fitness: Organism-Environment Interdependencies Preclude Long-Term Fitness Forecasting. Guilhem Doulcier, Peter Takacs, Pierrick Bourrat. BioEssays, 2020.
  9. Evolution of Collective-level Darwinian Properties. Guilhem Doulcier. in PhD Thesis, Université Paris Science et Lettres, 2019.
  10. A framework for estimating species-specific contributions to community indicators. Pierre Gausere*, Guilhem Doulcier*, Vincent Devictor, Sonia Kéfi. Ecological Indicators, 2019.
  11. Increases in aridity lead to drastic shifts in the assembly of dryland complex microbial networks. Manuel Delgado-Baquerizo, Guilhem Doulcier, David J. Eldridge, Daniel B. Stouffer, Fernando T. Maestre, Juntao Wang, Jeff R. Powell, Thomas C. Jeffries, Brajesh K. Singh. Land Degradation & Development, 2019.
  12. Experimental manipulation of selfish genetic elements links genes to microbial community function. Steven Quistad, Guilhem Doulcier, Paul B. Rainey. Phil. Trans. R. Soc. B, 2019. Preprint
  13. VConTACT: An IVirus Tool to Classify Double-Stranded DNA Viruses That Infect Archaea and Bacteria. Benjamin Bolduc, Ho Bin Jang, Guilhem Doulcier, Zhi-Qiang You, Simon Roux, Matthew B. Sullivan. PeerJ, 2017.
  14. The evolution of adhesiveness as a social adaptation. Thomas Garcia*, Guilhem Doulcier*, Silvia De Monte. eLife, 2015. Code
  15. Patterns and ecological drivers of ocean viral communities. Jennifer R. Brum, J. Cesar Ignacio-Espinoza, Simon Roux, Guilhem Doulcier, et al.. Science, 2015.


I had the pleasure to act as a teaching assistant in the Biology Department of the École normale supérieure (Paris). I collected several lecture notes in the domain of computer science, mathematics, and evolutionary theory.


I am or was involved in the development of several free and open source software.

  • Matpopmod (2021-ongoing, gitlab) : Analysis of Matrix Population Models in Python (beta)
  • Diapo (2019) : Assemble a presentation from a set of Inkscape SVGs.
  • Colgen (2019-ongoing) : Collective level genealogies analysis.
  • Dropsignal (2016-2020, gitlab): Signal processing for millifluidic experiments.
  • ZEN (2016-ongoing, gitlab): Evolutionary dynamics simulations.
  • ULM (2015-2017, gitlab): Population dynamics simulations.
  • rnetcarto (2014-2015, github): R-package for modularity optimisation of graph partition.
  • ipdt (2015): A tool to run iterated prisoner's dilemma tournaments.
  • ShapeCompare (2013): A tool to visualize and compare multiple hShape experiments results treated with the Shape Finder software.

I firmly believe that all knowledge should be free (as in free speech) and accessible to all. I would go as far as to say that it is a hard requirement for good science. I have no interest in working or collaborating using non-free software in a professional setting.


guilhem (dot) doulcier (at) normalesup (dot) org (PGP)