Contents Page

Conference Welcome ...... 1

Important Information...... 2

Invited Speakers...... 5

Program At A Glance...... 8

Detailed Program...... 10

Monday 1st April...... 10

Tuesday 2nd April...... 11

Wednesday 3rd April...... 14

Thursday 4th April...... 17

Poster Presentations...... 20

Delegate List ...... 24

Abstracts: Invited Speakers...... 31

Abstracts: Selected Talks...... 37

Abstracts: Poster Presentations...... 99

Conference Welcome

Welcome to Churchill College in the beautiful city of Cambridge, home to Evolution Evolving. The title of this meeting endeavours to highlight both that the evolutionary process itself evolves over time and that evolutionary biology is a vibrant field of enquiry with a theoretical framework that also evolves. Evolvability is just one of several emerging themes in the relationship between development and evolution that constitute the unique focus of this conference. Additional topics include the evolutionary causes and consequences of developmental bias, plasticity, niche construction and extra-genetic inheritance. Also distinctive of Evolution Evolving is the inclusion of contributions to the history and philosophy of evolutionary biology, which we believe are underappreciated but critically important to our field. We have tried hard to ensure that the meeting features a balanced program of talks and poster sessions, empirical and theoretical work, and a good mix of presentations from researchers of all genders, career stages, and backgrounds. We hope that you enjoy three wonderful days listening to marvelous talks and discussing evolutionary science, and leave the meeting feeling both excited and inspired.

The organisers

Paul Brakefield Kevin Laland Tobias Uller University of Cambridge University of St Andrews Lund University

Katrina Falkenberg Andrew Buskell University of St Andrews University of Cambridge

1 Important Information

The Venue

A map of Churchill College can be found on page four. Larger paper maps of Churchill College and Cambridge itself are available at the registration desk.

Main Concourse (C) - registration - welcome reception - poster sessions - morning and afternoon coffee breaks

Wolfson Hall (WH) - morning and afternoon talk sessions

Fellow’s Dining Room (FDR) - afternoon talk sessions

Churchill College Dining Hall (DH) - lunches - conference dinner

Seminar Room 6 - a quiet breakout space; this room is located close to staircase 37 in East Court (denoted ‘E’ on the map) and directly opposite the Jock Colville Hall

Meals and Refreshments

Included in your registration fees are the welcome reception on Monday evening, as well as morning and afternoon teas and fork buffet lunches on Tuesday, Wednesday and Thursday. Limited drinks will be served at the poster sessions. Additional drinks can be purchased from The Buttery (College bar), which is adjacent to the main concourse. The Buttery serves light snacks and refreshments throughout the day.

Wifi

Eduroam is available on site. Alternatively, you can collect a wifi ticket from the registration desk.

2 Social Media

We encourage the use of social media throughout the conference as it is a powerful tool to share findings and engage with the scientific community at large. Presenters: if you do not wish for your talk or a particular slide to be tweeted, please indicate this with an image such as the one to the right. Social media users: please respect the wishes of presenters, and do not tweet images of posters without the presenter’s permission.

#evoevolving is the conference hashtag and you can tag us with @evoevolving.

We recommend presenters and social media users read the social media guidelines of the World Marine Mammal Conference 2019: http://smmconference.org/socialmediaguidelines

Sponsors

We are very grateful for financial support from the John Templeton Foundation and The Galton Institute.

3 4 Invited Speakers

Alexander Badyaev Ecology and Evolutionary Biology, The University of Arizona, USA Badyaev is a Professor of Evolutionary Biology. He addresses conceptual questions at the intersection of evolution, ecology, and development using a wide array of empirical and analytical approaches in field and laboratory study systems.

Renee Duckworth Ecology and Evolutionary Biology, The University of Arizona, USA The ultimate goal of Duckworth’s work is to understand the link between micro and macroevolutionary processes with specific focus on the role of behavior in ecological and evolutionary feedbacks. To achieve these goals, she integrates approaches from evolutionary and physiological ecology using both large scale field experiments and laboratory systems.

Laurel Fogarty Max Planck Institute for Evolutionary Anthropology, Germany Fogarty works on theory related to non-genetic inheritance. Most of her work has a focus on cultural inheritance, cultural evolution, and cultural niche construction. She also has a general interest in theoretical niche construction.

Jukka Jernvall Institute of Biotechnology, University of Helsinki, Finland Jernvall’s research focuses on the developmental and ecological drivers of phenotypic evolution. By integrating different scales and methodologies, his group has used the mammalian dentition to explore how morphology can inform us about evolution.

5 Alan C Love Department of Philosophy and Minnesota Center for Philosophy of Science, University of Minnesota - Twin Cities, USA Love’s research focuses on conceptual issues in biology and has concentrated on evolutionary developmental biology, developmental biology, molecular biology, and paleontology. He uses a combination of approaches to investigate a variety of philosophical questions, including conceptual change, explanatory pluralism, knowledge structure, reductionism, the nature of historical science, and interdisciplinary epistemology.

Joanna Masel Ecology and Evolutionary Biology, The University of Arizona, USA Masel is a theoretical population geneticist and molecular evolutionist. She studies a range of foundational questions in evolution, including robustness and evolvability, the de novo birth of proteins and their subsequent long-term change, and the interplay between relative and absolute forms of competition.

Armin P Moczek Department of Biology, Indiana University, USA Moczek’s interests revolve broadly around the interplay of genetic, developmental, and ecological mechanisms in the origin and diversification of novel, complex traits. His work focuses on insects, from beetles and their horns to fireflies and their bio-luminescent lanterns.

Angela Potochnik Department of Philosophy, University of Cincinnati, USA Potochnik is an Associate Professor of Philosophy and the Director of the Center for Public Engagement with Science at the University of Cincinnati. Her research addresses the nature of science and its successes, the relationships between science and the public, and methods in population biology. She is the author of Idealizations and the Aims of Science (2017) and coauthor of Recipes for Science: An Introduction to Scientific Methods and Reasoning (2018).

6 Sean Rice Department of Biological Sciences, Texas Tech University, USA Rice’s research focuses on the mathematical principles underlying evolutionary processes. He is particularly interested in the role of stochasticity in the evolution of complex behavioural and developmental systems.

Jessica Riskin Department of History, Stanford University, USA Riskin’s research interests centre upon the history of science, politics and culture during the 17th to 19th centuries. She is currently working on a set of connected research projects in the history of evolutionary theory, including one on Lamarck and another on ideas about human evolution.

7 Program At A Glance

Monday 1st April 18.00 - 19.45 Welcome reception (C)

Tuesday 2nd April 08.50 - 10.00 Conference welcome and plenary talk (WH) 10.00 - 10.30 Coffee Break (C) 10.30 - 12.10 Selected talks (WH) 12.10 - 13.30 Lunch (DH) 13.30 - 14.10 Keynote talks, two parallel sessions (WH, FDR) 14.10 - 15.10 Selected talks, two parallel sessions (WH, FDR) 15.10 - 15.40 Coffee Break (C) 15.40 - 17.20 Selected talks, two parallel sessions (WH, FDR) 17.20 - 19.00 Poster Session (C)

Wednesday 3rd April 09.00 - 10.00 Plenary talk (WH) 10.00 - 10.30 Coffee Break (C) 10.30 - 12.10 Selected talks (WH) 12.10 - 13.30 Lunch (DH) 13.30 - 14.10 Keynote talks, two parallel sessions (WH, FDR) 14.10 - 15.10 Selected talks, two parallel sessions (WH, FDR) 15.10 - 15.40 Coffee Break (C) 15.40 - 17.20 Selected talks, two parallel sessions (WH, FDR) 17.20 - 19.00 Poster Session (C)

8 Thursday 4th April 09.00 - 10.00 Plenary talk (WH) 10.00 - 10.30 Coffee Break (C) 10.30 - 12.10 Selected talks (WH) 12.10 - 13.30 Lunch (DH) 13.30 - 14.10 Keynote talks, two parallel sessions (WH, FDR) 14.10 - 15.10 Selected talks, two parallel sessions (WH, FDR) 15.10 - 15.40 Coffee Break (C) 15.40 - 17.20 Selected talks, two parallel sessions (WH, FDR) 17.20 - 18.30 Free Time 18.30 - 19.30 Plenary talk (WH) 19.45 – 23.00 Conference Dinner (DH)

WH = Wolfson Hall FDR = Fellow’s Dining Room C = Concourse DH = Churchill College Dining Hall

9 Detailed Program

Monday 1st April

18.00 - 19.45 Registration 18.00 - 19.45 Welcome reception

10 Tuesday 2nd April

Wolfson Hall Morning 08.50 - 09.00 Conference Welcome: Kevin Laland 09.00 - 10.00 Plenary Talk: Renee Duckworth Dynamic stability and robustness of ecological systems: a missing link to evolutionary mechanisms? The University of Arizona, USA 10.00 - 10.30 Coffee Break 10.30 - 12.10 General Session 10.30 - 10.50 Mycalesina in morphospace: how developmental bias shapes evolutionary diversification in butterflies Oskar Brattström, University of Cambridge, UK 10.50 - 11.10 Fighting about frequency Karen Kovaka, Virginia Tech, USA 11.10 - 11.30 On leaders and followers: the evolution of phenotypic plasticity can lead the evolution of genetic evolvability more easily than vice versa Miguel Brun-Usan, University of Southampton, UK 11.30 - 11.50 Bird nests evolve flexibly in response to egg characteristics Catherine Sheard, University of St Andrews, UK 11.50 - 12.10 Adaptive evolution NOT driven by natural selection (?!?). On individual responses, inclusive inheritance, and fleshing out the second term of that famous Price equation Pim Edelaar, University Pablo de Olavide, Spain 12.10 - 13.30 Lunch

11 Wolfson Hall Afternoon 13.30 - 14.10 Keynote Talk: Armin P Moczek On the origins of novelty and diversity in development and evolution: case studies on horned beetles Indiana University, USA 14.10 - 17.20 Themed Session: Developmental Plasticity 14.10 - 14.30 DNA methylation lends plasticity to the coloration of an African cichlid Sebastian Alvarado, Queens College CUNY, USA 14.30 - 14.50 Unexplored maternal effect mediators, thyroid hormones, in an eco-evolutionary context Suvi Ruuskanen, University of Turku, Finland 14.50 - 15.10 Phenotypic plasticity facilitates colonization of savannas in Mycalesina butterflies Sridhar Halali, University of Cambridge, UK 15.10 - 15.40 Coffee Break 15.40 - 16.00 Does thermal plasticity align with local adaptation? – An interspecific comparison of wing shape and size in sepsid flies Patrick Rohner, University of Zurich, Switzerland 16.00 - 16.20 Macroevolutionary dynamics of phenological plasticity Stephen De Lisle, Lund University, Sweden 16.20 - 16.40 Developmental regulation and evolution of nutrition-responsive growth: transcriptomic underpinnings of plasticity in horned beetles Sofia Casasa, Indiana University, USA 16.40 - 17.00 Involvement of maternal yolk thyroid hormones in the evolution of life history traits in birds: a comparative analysis Bin-Yan Hsu, University of Turku, Finland 17.00 - 17.20 Phenotypic divergence across spadefoot toad species mirrors within-species consequences and mechanisms of developmental plasticity Christoph Liedtke, Doñana Biological Station, CSIC, Spain 17.20 - 19.00 Poster Session

12 Fellow’s Dining Room Afternoon 13.30 - 14.10 Keynote Talk: Angela Potochnik Causal patterns and how our theories change University of Cincinnati, USA 14.10 - 17.20 Themed Session: Philosophy of Biology 14.10 - 14.30 What the history of biology tells us about the Extended Evolutionary Synthesis Jan Baedke, Ruhr University Bochum, Germany Daniel J Nicholson, Konrad Lorenz Institute for Evolution and Cognition Research, Austria 14.30 - 14.50 Towards a novel unifying theory of the genotype concept David Ricote Hernández, Centro Andaluz de Biología del Desarrollo, CSIC-Universidad Pablo de Olavide-Junta de Andalucía, Spain 14.50 - 15.10 Rethinking cybernetics in contemporary theoretical biology Flavia Fabris, Egenis, the Centre for the Study of Life Sciences, University of Exeter, UK 15.10 - 15.40 Coffee Break 15.40 - 16.00 The Extended Evolutionary Synthesis debate: some epistemological and sociological dimensions Alejandro Fábregas-Tejeda and Francisco Vergara-Silva, Instituto de Biología, Universidad Nacional Autónoma de México, Mexico 16.00 - 16.20 Beyond standard views of reproduction: pregnant females as individuals and their role in evolution Laura Nuño de la Rosa and Arantza Etxeberria, Complutense University of Madrid, Spain 16.20 - 16.40 Evolution of multicellularity: cheating done right Walter Veit, University of Bristol, UK 16.40 - 17.00 The explanatory robustness of developmental types: a propensity view Cristina Villegas, Complutense University of Madrid, Spain 17.00 - 17.20 The cultural evolution of cultural evolution Jonathan Birch, London School of Economics & Political Science, UK 17.20 - 19.00 Poster Session

13 Wednesday 3rd April

Wolfson Hall Morning 09.00 - 10.00 Plenary Talk: Alan C Love Structuring knowledge in evolutionary biology University of Minnesota - Twin Cities, USA 10.00 - 10.30 Coffee Break 10.30 - 12.10 General Session 10.30 - 10.50 Evolution by natural induction Richard Watson, University of Southampton, UK 10.50 - 11.10 Generalized pathways to specific outcomes: stress links maternal environment and adaptive offspring trait variation Ahva Potticary, The University of Arizona, USA 11.10 - 11.30 Evolution of migratory behavior by barnacle geese: the roles of experience, memory and social learning Thomas Oudman, NIOZ Netherlands Institute for Sea Research, The Netherlands, and University of St Andrews, UK 11.30 - 11.50 Theory-based ecology and niche construction Liz Pásztor, Biological Institute, Eötvös Loránd University, Hungary 11.50 - 12.10 The Hot Spring Hypothesis for the origin of life: implications for niche construction theory in the first major transition in evolution from non-life to Life Bruce Damer, The Biota Institute, USA, and UC Santa Cruz, USA 12.10 - 13.30 Lunch

14 Wolfson Hall Afternoon 13.30 - 14.10 Keynote Talk: Jukka Jernvall To walk or not to walk the line of a developmental bias University of Helsinki, Finland 14.10 - 16.00 Themed Session: Developmental Bias 14.10 - 14.30 Developmental bias and morphological convergence in Anolis lizards Nathalie Feiner, Lund University, Sweden 14.30 - 14.50 Measuring developmental bias: quantitative genetics meets developmental biology Lisandro Milocco, University of Helsinki, Finland 14.50 - 15.10 The evolutionary potential of environmentally induced phenotypes Reinder Radersma, Lund University, Sweden 15.10 - 15.40 Coffee Break 15.40 - 16.00 Developmental interactions established during speciation leave a lasting trace in descendent populations Laura Mears, University of Southampton, UK 16.00 - 17.20 Themed Session: Evolutionary Innovation 16.00 - 16.20 What kind of evolutionary novelty (if any) could be language? Salvatore Ivan Amato and Giuliana Pulvirenti, University of Messina, Italy 16.20 - 16.40 It’s a horn, a wing, a helmet: the role of wing serial homologs in insect innovations Yonggang Hu, Indiana University, USA 16.40 - 17.00 Beyond Darwin: how every individual adapts to its internal perturbations? Yoav Soen, Weizmann Institute of Science, Israel 17.00 - 17.20 How the beetle head got its horns: integrating a novel trait into a conserved trait complex David M Linz, Indiana University, USA 17.20 - 19.00 Poster Session

15 Fellow’s Dining Room Afternoon 13.30 - 14.10 Keynote Talk: Laurel Fogarty The inheritance and heritability of niche constructed traits Max Planck Institute for Evolutionary Anthropology, Germany 14.10 - 16.20 Themed Session: Niche Construction 14.10 - 14.30 Niche construction affects the consistency and strength of natural selection Andrew Clark, University of St Andrews, UK 14.30 - 14.50 Linking coral morphological traits and reef fish community structure Luisa Fontoura, Macquarie University, Australia 14.50 - 15.10 Microbial community structure and function driven by infaunal niche construction in intertidal sediments Adam Wyness, University of St Andrews, UK 15.10 - 15.40 Coffee Break 15.40 - 16.00 Keeping close neighbours, corals are spatially clustered within and across species Maria Dornelas, University of St Andrews, UK 16.00 - 16.20 Holed-up: nesting in a cavity as an evolutionary driver of parental care in fishes and birds Karina Vanadzina, University of St Andrews, UK 16.20 - 17.20 Themed Session: Cultural Evolution 16.20 - 16.40 Trait specialisation, innovation and the evolution of culture in changing environments Dominik Deffner, Max Planck Institute for Evolutionary Anthropology, Germany 16.40 - 17.00 A New Caledonian crow's-eye-view on how to develop probing intelligence: the evolution of tool-use and tool-making as embodied, distributed and socially situated perception action processes Joanna Fairlie, Independent Scholar 17.00 - 17.20 Generalized Darwinism revisited: how a new synthesis changes our view on cultural evolution Karim Baraghith, Heinrich-Heine University, Germany 17.20 - 19.00 Poster Session

16 Thursday 4th April

Wolfson Hall Morning 09.00 - 10.00 Plenary Talk: Alexander Badyaev Control theory in evolution The University of Arizona, USA 10.00 - 10.30 Coffee Break 10.30 - 12.10 General Session 10.30 - 10.50 Stress-induced mutagenesis systems and the adaptive immunity Bartlomiej Swiatczak, University of Science and Technology of China, China 10.50 - 11.10 Phenotypic integration as both developmental bias and developmental facilitation of evolutionary diversification: empirical examples from teeth, toes, and torsos Kathryn Kavanagh, University of Massachusetts Dartmouth, USA 11.10 - 11.30 Can plasticity create irreversible constraints? Alfredo Rago, University of Southampton, UK 11.30 - 11.50 Cultural evolution, revolution and transmission of humpback whale song Ellen C Garland, University of St Andrews, UK 11.50 - 12.10 Niche construction: resolving the process debate Tim Lewens, University of Cambridge, UK 12.10 - 13.30 Lunch

17 Wolfson Hall Afternoon 13.30 - 14.10 Keynote Talk: Joanna Masel Mutational and pre-mutational shaping of adaptation The University of Arizona, USA 14.10 - 17.20 Themed Session: Evolvability 14.10 - 14.30 Comprehensive transcriptomics tracks changes in gene expression and coding sequences during C4 evolution Lamiaa A Munshi, The University of Sheffield, UK 14.30 - 14.50 Computational complexity is an ultimate constraint on evolution Artem Kaznatcheev, University of Oxford, UK 14.50 - 15.10 Population-level variation in sperm-mediated paternal effects in threespined sticklebacks Jennifer Hellmann, University of Illinois Urbana-Champaign, USA 15.10 - 15.40 Coffee Break 15.40 - 16.00 The effect of parental care on population evolvability Sonia Pascoal, University of Cambridge, UK 16.00 - 16.20 The thrifty watchmaker Michael Lachmann, Santa Fe Institute, USA 16.20 - 16.40 Variable phenotypic plasticity and its consequences for evolutionary potential Franziska Brunner, University of Liverpool, UK 16.40 - 17.00 Comparative genomics of convergent evolution in the emergence of complex traits Claudio L Flores Martinez, University of Hamburg, Germany 17.00 - 17.20 Epigenetic potential in native and introduced populations of house sparrows (Passer domesticus) Haley Hanson, University of South Florida, USA 17.20 - 18.30 Free time 18.30 - 19.30 Plenary Talk: Jessica Riskin The restless machinery of life Stanford University, USA 19.45 - 23.00 Conference Dinner

18 Fellow’s Dining Room Afternoon 13.30 - 14.10 Keynote Talk: Sean Rice How to make general theories of evolution applicable to complex special cases Texas Tech University, USA 14.10 - 16.00 Themed Session: Mathematical Models 14.10 - 14.30 Why does natural selection favour the transgenerational inheritance of social adversity? Bram Kuijper, University of Exeter, UK 14.30 - 14.50 Trait mean-variance relationships impact speed of adaptation Rebecca Hoyle, University of Southampton, UK 14.50 - 15.10 The influence of multi-level demographic and ecological feedbacks in the evolution of cooperation Antonio MM Rodrigues, University of Cambridge, UK 15.10 - 15.40 Coffee Break 15.40 - 16.00 The evolution of teaching and anti-teaching James Allen, University of Cambridge, UK 16.00 - 17.20 Themed Session: Inclusive Inheritance 16.00 - 16.20 Evolution of small RNA production under fluctuating environmental conditions Willian TAF Silva, Uppsala University, Sweden 16.20 - 16.40 Inheritance of acquired traits - implications for evolution? Sophie Juliane Veigl, University of Vienna, Austria 16.40 - 17.00 Egg development plasticity and the transmission of maternal effects in Daphnia Stewart Plaistow, University of Liverpool, UK 17.00 - 17.20 Environmental heterogeneity mediates the evolution of trans- generational effects on fitness Martin Lind, Uppsala University, Sweden

19 Poster Presentations

P01 Expanding the conceptual scope of niche construction theory Bendik Hellem Aaby, KU Leuven, Belgium P02 Could a major evolutionary transition have happened in human evolution? Paulo C Abrantes, University of Brasília, Brazil P03 Dialectics of evolution; epistemology versus practice Ozan A Altinok, WWU Münster, Germany P04 Overprinting and the evolvability of novel protein-coding genes Zachary Ardern, Technical University of Munich, Germany P05 Morphological divergence in Drosophila ananassae due to nutritional variations through changes in allometric relationship and trait size Bhumika, Banaras Hindu University, India P06 Coral-built structural complexity enhances coral recruitment: niche construction in action? Viviana Brambilla, University of St Andrews, UK P07 When are more experiments necessary? Using evolutionary considerations in investigating animal minds Simon Brown, Columbia University, USA P08 The evolution of unicellularity in a multicellular world Charlie Cornwallis, Lund University, Sweden P09 Enactive evolution Amanda Corris, University of Cincinnati, USA P10 Behemoth: Henri Bergson's and Gilles Deleuze's vitalism, a proto evolutionary biology Jack Robert Edmunds-Coopey, University of Durham, UK P11 Effects of phenotypic robustness on adaption and evolvability Giuseppe Fusco, University of Padova, Italy P12 Spadefoot toad tadpoles incur maintenance costs of developmental plasticity in the form of increased oxidative stress Ivan Gomez-Mestre, Doñana Biological Station, CSIC, Spain P13 The evolution of barriers to exploitative behaviors Jonathan R Goodman, University of Cambridge, UK

20 P14 Three lessons of Bergson's creative evolution in the wake of transhuman evolution Daina Habdankaite, Vilnius University, Lithuania, and Turin University, Italy P15 Evolution of phenotypic complexity: a developmental perspective Pascal Felix Hagolani, University of Helsinki, Finland P16 Timing of maternal exposure to toxic cyanobacteria and offspring fitness in Daphnia magna: implications for the evolution of anticipatory maternal effects Alexander Hegg, Lund University, Sweden P17 Studying the evolutionary process with genetic algorithms Edith Invernizzi, University of St Andrews, UK P18 The evolutionary origin of our moral beliefs Man-Him Ip, University of Birmingham, UK P19 Genetic assimilation in the fossil record: phenotypic plasticity and accommodation in Cambrian arthropods Illiam Jackson, Lund University, Sweden P20 Extended cognitive systems, meaning, and evolutionary transitions Hilton F Japyassú, Universidade Federal da Bahia, Brazil P21 Mechanisms of hormone mediated parent-offspring conflict Neeraj Kumar, University of Groningen, The Netherlands, and Max Planck Institute for Ornithology, Germany P22 The impact of individual immune experience on niche construction in red flour beetles, Tribolium castaneum Lai Ka Lo, University of Münster, Germany P23 Trait-based theory leads to generality across successional systems Jessica Haghkerdar, University of St Andrews, UK P24 From anisogamy to intrasexual competition Ivain Martinossi, Uppsala University, Sweden P25 Diet-based developmental plasticity in a freshwater isopod Blake Matthews, Eawag, Switzerland P26 Global human niche construction: an evolutionary trap? Andra Meneganzin, Università degli Studi di Padova, Italy P27 A weapons-testes tradeoff in males extends to female traits Christine Miller, University of Florida, USA

21 P28 Assessing adaptive and non-adaptive evolution of the glycolytic enzyme triosephosphate isomerase Ricardo Muñiz Trejo, National Autonomous University of Mexico, Mexico P29 The evolution of hierarchical structure: separating the causes and consequences of evolvability Frederick J Nash, University of Southampton, UK P30 Spandrels and trait delimitation: no such thing as “architectural constraint” Mark E Olson, Universidad Nacional Autónoma de México, Mexico P31 With a little help from my friends: the role of the microbiota in dung beetle diversification Erik Parker, Indiana University, USA P32 Transposable elements: the unexpected junk that shaped genomes and challenged evolutionary biology Valentina Peona, Uppsala University, Sweden P33 Weismann, Crick, and others: flows of information in evolution Arnaud Pocheville, CNRS and Université Paul Sabatier, France P34 Delineating the boundaries of adaptation by tracing histories of environmental induction Ahva Potticary, The University of Arizona, USA P35 The interaction of learning and evolution can enable adaptive behaviours that are not presupposed in the learning ability David Prosser, University of Southampton, UK P36 Naturalising the language faculty: the role of structures and developmental processes Giuliana Pulvirenti, University of Messina, Italy P37 Teleology and evolution Rasmus Sandnes Haukedal, Durham University, UK P38 The evolution of adaptive epigenetic inheritance Zuzana Sekajova, Uppsala University, Sweden P39 Unique evolutionary properties of plastic traits: an empirical study of erosion of phenotypic plasticity under relaxed selection Helen Spence-Jones, University of St Andrews, UK P40 A culturally driven plasticity hypothesis for the evolution of human life course Francesco Suman, Independent Scholar

22 P41 Unique, essential, and unknown: the enigma of taxonomically restricted, essential genes of unknown function Change Tan, University of Missouri, USA P42 The evolution of relatedness and evolutionary transitions in individuality Christoph Thies, University of Southampton, UK P43 Deconstructing reef complexity: relating coral diversity, rugosity and fractal dimension Damaris Torres-Pulliza, Macquarie University, Australia P44 Cultural evolution and neuroscience: a pathway to integration Ryutaro Uchiyama, London School of Economics, UK P45 Lost (and found) in translation: circulation of EES-related concepts in two scientific communities in Mexico Francisco Vergara-Silva and Alejandro Fábregas-Tejeda, Universidad Nacional Autónoma de México, Mexico P46 Testing the flexible stem hypothesis in stickleback fish: is ancestral morphological plasticity adaptive? Matthew Wund, The College of New Jersey, USA P47 Quantifying niche constructing traits in reef corals Kyle Zawada, University of St Andrews, UK P48 Two senses of biological possibility Jason Zinser, University of Wisconsin - Stevens Point, USA P49 Plasticity in mouthpart length in response to developmental diet in leaf-footed bugs Sara Zlotnik, University of Florida, USA P50 Evolution and development of polyphenisms: a mechanistic model based on bistable switches Roman Zug, Lund University, Sweden

23 Delegate List

Aaby, Bendik Hellem KU Leuven P01 Abrantes, Paulo C University of Brasília P02 Allen, James University of Cambridge FDR, 4th Altinok, Ozan A WWU Münster P03 Alvarado, Sebastian Queens College CUNY WH, 2nd Amato, Salvatore Ivan University of Messina WH, 3rd Ardern, Zachary Technical University of Munich P04 Armfield, Roxanne University of Cambridge Arnold, Kevin John Templeton Foundation Badyaev, Alex The University of Arizona WH, 4th Baedke, Jan Ruhr University Bochum FDR, 2nd Banzhaf, Wolfgang Michigan State University Baraghith, Karim Heinrich Heine University Düsseldorf FDR, 3rd Beaudoin, Chris University of Cambridge Berkemeier, Francisco University College London Bhumika Banaras Hindu University P05 Birch, Jonathan London School of Economics FDR, 2nd Bocedi, Greta University of Aberdeen Brakefield, Paul University of Cambridge Brambilla, Viviana University of St Andrews P06 Brattström, Oskar University of Cambridge WH, 2nd Brown, Simon Columbia University P07 Brun-Usan, Miguel University of Southampton WH, 2nd Brunner, Franziska University of Liverpool WH, 4th Buskell, Andrew University of Cambridge

Campo Bes, Israel University of Valencia Cano, Hugo University College London Casasa, Sofia Indiana University WH, 2nd Catherall, Andrew University of Cambridge Chauvet, Adrien The University of Sheffield

24 Clark, Andrew University of St Andrews FDR, 3rd Collins-Attwood, Mairenn University of Cambridge Cornwallis, Charlie Lund University P08 Corris, Amanda University of Cincinnati P09 Damer, Bruce The Biota Institute, UC Santa Cruz WH, 3rd De Lisle, Stephen Lund University WH, 2nd De Sanctis, Bianca University of Cambridge Deffner, Dominik Max Planck Institute for Evolutionary FDR, 3rd Anthropology Dobreva, Mariya Imperial College London Dornelas, Maria University of St Andrews FDR, 3rd Duckworth, Renee The University of Arizona WH, 2nd Edelaar, Pim University Pablo de Olavide WH, 2nd Edmunds-Coopey, Jack Robert University of Durham P10 Etxeberria, Arantza University of the Basque Country FDR, 2nd Fábregas-Tejeda, Alejandro Universidad Nacional Autónoma de FDR, 2nd México Fabris, Flavia University of Exeter FDR, 2nd Fairlie, Joanna Independent Scholar FDR, 3rd Falkenberg, Katrina J University of St Andrews

Falzone, Alessandra University of Messina

Faria, Vitor G University of Basel Feiner, Nathalie Lund University WH, 3rd

Feldman, Marcus Stanford University Fjågesun, Tonje University of Cambridge Flores Martinez, Claudio L University of Hamburg WH, 4th Fogarty, Laurel Max Planck Institute for Evolutionary FDR, 3rd Anthropology Fontoura, Luisa Macquarie University FDR, 3rd Fusco, Giuseppe University of Padova P11

Gallardo Pinedo, Adrian University of Valencia Garland, Ellen C University of St Andrews WH, 4th

George, Alison New Scientist 25 Gomez-Mestre, Ivan Doñana Biological Station, CSIC P12 Goodman, Jonathan R University of Cambridge P13

Goymer, Patrick Nature Groothuis, Ton University of Groningen Habdankaite, Daina Vilnius University, Turin University P14 Haghkerdar, Jessica University of St Andrews P23 Hagolani, Pascal Felix University of Helsinki P15 Halali, Sridhar University of Cambridge WH, 2nd Halina, Marta University of Cambridge Halle, Stefan Friedrich Schiller University Jena Hanson, Haley University of South Florida WH, 4th Hegg, Alexander Lund University P16

Helanterä, Heikki University of Oulu Hellmann, Jennifer University of Illinois Urbana- WH, 4th Champaign Henderson, Tara University of Cambridge Hoyle, Rebecca University of Southampton FDR, 4th Hsu, Bin-Yan University of Turku WH, 2nd Hu, Yonggang Indiana University WH, 3rd Invernizzi, Edith University of St Andrews P17 Ip, Man-Him University of Birmingham P18 Jackson, Illiam Lund University P19 Japyassú, Hilton F Universidade Federal da Bahia P20 Jernvall, Jukka University of Helsinki WH, 3rd Johnstone, Rufus University of Cambridge Kavanagh, Kathryn University of Massachusetts WH, 4th Dartmouth Kaznatcheev, Artem University of Oxford WH, 4th Kilner, Rebecca University of Cambridge Kovaka, Karen Virginia Tech WH, 2nd Kozielska, Magdalena University of Groningen Kuijper, Bram University of Exeter - Penryn Campus FDR, 4th

26 Kumar, Neeraj University of Groningen, P21 Max Planck Institute for Ornithology Lachmann, Michael Santa Fe Institute WH, 4th Laland, Kevin N University of St Andrews

Lazarus, John Newcastle University Leitao, Alex University of Cambridge Lewens, Tim University of Cambridge WH, 4th Liedtke, Christoph Doñana Biological Station, CSIC WH, 2nd Lind, Martin Uppsala University FDR, 4th Linz, David M Indiana University WH, 3rd

Livnat, Adi University of Haifa Lo, Lai Ka University of Münster P22 Love, Alan C University of Minnesota - Twin Cities WH, 3rd

Madin, Joshua S University of Hawaii

Maeso, Ignacio Andalusian Centre for Developmental Biology Mahen, Beeshman University of Cambridge Marconi, Aleksandra University of Cambridge Martin, Marty University of South Florida Martinez, Pedro Universitat de Barcelona Martinossi, Ivain Uppsala University P24 Masel, Joanna The University of Arizona WH, 4th Matthews, Blake Eawag P25 Mears, Laura University of Southampton WH, 3rd Méndez Salinas, Emiliano University of Groningen Meneganzin, Andra Università degli Studi di Padova P26 Merleau-Ponty, Noémie University of Cambridge Miller, Christine University of Florida P27 Milocco, Lisandro University of Helsinki WH, 3rd Moczek, Armin P Indiana University WH, 2nd Muñiz Trejo, Ricardo National Autonomous University of P28 Mexico Munshi, Lamiaa A The University of Sheffield WH, 4th 27 Nash, Frederick J University of Southampton P29

Nelson, Paul Biola University Nicholson, Daniel J Konrad Lorenz Institute for Evolution FDR, 2nd and Cognition Research Nuño de la Rosa, Laura Complutense University of Madrid FDR, 2nd Odling-Smee, John Mansfield College - Oxford Odling-Smee, Ros University of Oxford - Retiree Olson, Mark E Universidad Nacional Autónoma de P30 México Oudman, Thomas NIOZ Netherlands Institute for Sea WH, 3rd Research, University of St Andrews Padilla Iglesias, Cecilia University of Cambridge Page, Karen M University College London Pagnotta, Murillo University of St Andrews Parker, Erik Indiana University P31 Pascoal, Sonia University of Cambridge WH, 4th Pásztor, Liz Eötvös University WH, 3rd Paternò, Orazioluca Università degli Studi di Torino Paterson, David University of St Andrews Peona, Valentina Uppsala University P32

Perera, Roly Alan Turing Institute

Picao-Osorio, Joao Institute of Biology of the École Normale Supérieure Plaistow, Stewart University of Liverpool FDR, 4th Pocheville, Arnaud CNRS, Université Paul Sabatier P33 Potochnik, Angela University of Cincinnati FDR, 2nd Potticary, Ahva The University of Arizona WH, 3rd, P34

Pretelli, Ilaria Max Planck Institute for Evolutionary Anthropology Prosser, David University of Southampton P35 Pulvirenti, Giuliana University of Messina WH, 3rd, P36 Radersma, Reinder Lund University WH, 3rd Rago, Alfredo University of Southampton WH, 4th 28

Ratikainen, Irja Norwegian University of Science and Technology Renton, Jessie University College London

Retzlaff, Nancy MPI for Mathematics in the Sciences, Leipzig University Rice, Sean Texas Tech University FDR, 4th Ricote Hernández, David Andalusian Centre for Developmental FDR, 2nd Biology Riederer, Jana University of Groningen Riskin, Jessica Stanford University WH, 4th

Robin, Nicolas University of Teacher Education St Gallen Rodrigues, Antonio MM University of Cambridge FDR, 4th Rohner, Patrick University of Zurich WH, 2nd Ruuskanen, Suvi University of Turku WH, 2nd Sandnes Haukedal, Rasmus Durham University P37 Santos, Emília University of Cambridge Scherrer, Raphaël University of Groningen Sekajova, Zuzana Uppsala University P38 Sheard, Catherine University of St Andrews WH, 2nd Silva, Willian TAF Uppsala University FDR, 4th Soen, Yoav Weizmann Institute of Science WH, 3rd Spence-Jones, Helen University of St Andrews P39

Stocks, Miranda IST Austria Suman, Francesco Independent Scholar P40 Swiatczak, Bartlomiej University of Science and Technology WH, 4th of China Tan, Change University of Missouri P41 Thies, Christoph University of Southampton P42 Torres-Pulliza, Damaris Macquarie University P43 Toyokawa, Wataru University of Konstanz Tschol, Maximilian University of Aberdeen

Tucker, Roger Sonocent Ltd 29 Uchiyama, Ryutaro London School of Economics P44

Uller, Tobias Lund University van den Engel, Peter Independent Scholar van Eldijk, Timo University of Gronignen Vanadzina, Karina University of St Andrews FDR, 3rd Vasilyeva, Zinaida Technical University of Munich Veigl, Sophie Juliane University of Vienna FDR, 4th Veit, Walter University of Bristol FDR, 2nd Vergara-Silva, Francisco Universidad Nacional Autónoma de FDR, 2nd, P45 México Villegas, Cristina Complutense University of Madrid FDR, 2nd

Vinterstare, Jerker Lund University Vykuntam, Vinit University of Cambridge Wason, Paul John Templeton Foundation Watson, Richard A University of Southampton WH, 3rd While, Geoff University of Tasmania

White, Roger Memorial University of Newfoundland Wilkins, Adam S Humboldt University Berlin

Winder, Isabelle Bangor University Wund, Matthew The College of New Jersey P46 Wyness, Adam University of St Andrews FDR, 3rd Yen, Eugenie University of Cambridge Zawada, Kyle University of St Andrews P47 Zinser, Jason University of Wisconsin - Stevens Point P48 Zlotnik, Sara University of Florida P49 Zug, Roman Lund University P50

WH = talk in Wolfson Hall, date FDR = talk in Fellow’s Dining Room, date P = Poster

30 Abstracts: Invited Speakers

Dynamic stability and robustness of ecological systems: a missing link to evolutionary mechanisms? Renee Duckworth The University of Arizona, USA

Ecosystems are often viewed as self-organizing systems that remain stable in their outputs. Their pattern of dynamic stability is remarkably similar to patterns at the macroevolutionary scale in species’ stasis. Yet, it is unclear whether the types of dynamics that produce stability in ecosystems are causally linked to patterns of species’ stasis, and, if so, what are the mechanisms underlying both? I suggest that ecological communities are in a state of dynamic stability which is maintained by the active responses of organisms to environmental change through processes such as niche construction, adaptive plasticity and habitat selection. As such, persistent evolutionary change may only occur when there is a major disruption of the ecological community as a whole. To show how such dynamics might be studied and how a body of work can be reinterpreted with this novel framework, I draw on my work on the population and community dynamics of cavity nesting birds. I use this example to illustrate how a deeper understanding of the mechanisms underlying dynamic stability may be crucial to predicting the two most common outcomes of environmental change: whether populations will go extinct or evolve.

31 On the origins of novelty and diversity in development and evolution: case studies on horned beetles Armin Moczek Indiana University, USA

Understanding how development and ecology shape organismal evolution is a central goal of evolutionary developmental biology. In my presentation I highlight a class of traits and organisms that are emerging as new models in evo devo and eco devo research: beetle horns and horned beetles. Horned beetles are morphologically diverse, ecologically rich, and developmentally and genetically increasingly accessible. Recent studies have begun to take advantage of these attributes and are starting to link the microevolution of horned beetle development to the macroevolution of novel, complex traits, and to identify the genetic, developmental, and ecological mechanisms, and the interactions between them, that mediate organismal innovation and diversification in natural populations. Here, I review the most significant recent findings and their contributions to current frontiers in evolutionary developmental biology. In particular, I explore the role of developmental mechanisms in biasing and facilitating innovation in evolution and the significance of host-symbiont interactions and niche construction in ecological radiations.

Causal patterns and how our theories change Angela Potochnik University of Cincinnati, USA

In evolutionary biology, attention to causes—or mechanism—is common, as is debate about which causes are important. In this talk, I motivate the idea that scientists aim to identify not just causes, but what I call causal patterns. Focus on a given causal pattern is motivated by a combination of causal facts and the nature of the research priorities. This framing changes how we should think about theory development, including the development of evolutionary theory. Theory change can be motivated in two different ways: by the discovery that a causal pattern fails to obtain, or by a shift in research priorities toward a focus on different causal patterns. On the other hand, this view also predicts the maintenance of multiple accounts of the same phenomena, accounts that may seem incompatible but that capture different causal patterns, of interest to different research programs. I develop these points with attention to the relationship between developmental biology and evolutionary theory. 32 Structuring knowledge in evolutionary biology Alan C Love University of Minnesota - Twin Cities, USA

Ongoing debates about the possibility and status of an extended evolutionary synthesis often involve disagreements about the nature and significance of particular phenomena (e.g., developmental plasticity) or mechanisms (e.g., phenotypic accommodation). However, assumptions regarding how to structure knowledge about phenomena and mechanisms in evolutionary biology remain largely implicit in these conversations. I argue that greater attention to these assumptions—and the expectations they foster— has the potential to reveal the conceptual basis of some existing controversies about theoretical frameworks in evolutionary biology and provide novel ways of structuring our evolving knowledge for future interdisciplinary inquiry.

To walk or not to walk the line of a developmental bias Jukka Jernvall University of Helsinki, Finland

Developmental processes can influence, or even bias phenotypic variation available for natural selection. While developmental biases are perhaps generally acknowledged, explanations of the specific impacts of development on evolution are easily little more than ‘just so stories’. To provide more disciplined predictions about the effects of development on evolution, developmental models or rules can be used. But what does it mean when the data does not obey a developmental rule? I will discuss a development-based ‘inhibitory cascade’ rule that predicts the evolution of mammalian tooth proportions. Whereas many mammalian species appear to vary as predicted by the inhibitory cascade rule, there are also exceptions. These exceptions can be argued to disprove the inhibitory cascade, but the exceptions themselves should also be examined critically. I will show how experimentally breaking the inhibitory cascade uncovers the processes underlying the rules, and how evolutionary exceptions may not be that exceptional after all.

33 The inheritance and heritability of niche constructed traits Laurel Fogarty, Michael J Wade Max Planck Institute for Evolutionary Anthropology, Germany

A central tenet of niche construction theory is that organisms, through their own actions, can alter the selective pressures to which they are subject and, in turn, change the rates and perhaps trajectories of their species’ evolution. However, the ways in which niche construction changes not just the selection on a trait but the structure of the inheritance of that trait are poorly understood. Here we describe a quantitative genetic model of niche construction that describes the inheritance patterns for a niche- constructing trait and an associated niche-construction-sensitive trait. We show how the response to selection can differ from that which would be expected in the absence of niche construction as a result of changes to associated breeding values. We compare these models to existing and closely related models of indirect genetic effects and discuss the implications for our understanding of the heritability of niche constructing traits and the importance of niche construction in evolutionary theory in general.

Control theory in evolution Alexander Badyaev The University of Arizona, USA

With a few notable exceptions from neurobiology, cytology, and physiology, there is little cross-talk between evolutionary theory and control theory of dynamic systems. Yet, both fields are intensely focused on essentially an identical set of problems – on reconciling stability and change, continuity and discreetness, innovability and complexity, and inherency and contingency. Both fields aim to resolve these problems by predicting the distribution and kinds of “controls” that either change or regulate complex systems. As a result, we now have a set of parallel solutions to essentially the same problems but from largely non-overlapping starting points, backgrounds, and motivations. I will review these solutions, their empirical manifestations, and the insights they provide into some of the most long-standing issues in evolutionary biology, including the most enigmatic of them all – the relationship between form and function in development and inheritance.

34 Mutational and pre-mutational shaping of adaptation Joanna Masel The University of Arizona, USA

Evolvability depends on both the quantity and quality of heritable variants. For polygenic traits, the quality of the mutational neighborhood is a more important determinant of a population’s evolvability than the spread of the population across a neutral genotype network. A first approximation of mutational quality is the ratio between the two modes of the distribution of fitness effects, which tend either to reduce fitness to zero, or have weak effects, but are rarely in between. If developmental errors in the present mimic the effects of future mutations, selection can act pre- mutationally to make the relative frequencies of these two modes more favorable. Data on the cryptic sequences beyond stop codons suggests that highly expressed cryptic sequences have experienced more such pre-mutational shaping. Evolutionary capacitance to exploit such pre-screened sources of variation can either evolve through capacitance widgets such as the yeast [PSI+] prion, or can emerge non-adaptively. Capacitance makes the crossing of fitness valleys far more likely. Epimutations differ from ordinary mutations in their rates – they are typically not just higher, but also lack a strong excess of loss of function over gain of function – but are otherwise analyzable by conventional theoretical population genetic approaches. The relative availability of different kinds of beneficial mutation is known to shape adaptation when the product of the beneficial mutation rate and census population size UN<1. When UN>1, clonal interference is expected to make differences in the selection coefficient s much more important than differences in U. However, the phenomenon of mutation-driven adaptation re-emerges when U>s.

35 How to make general theories of evolution applicable to complex special cases Sean Rice Texas Tech University, USA

Much of evolutionary theory consists of special case models; built around simplifying assumptions that make them tractable, but limit their applicability outside of the particular systems for which they were built. We also have some very general results, such as the Price equation, that are mathematically simple and in principle hold true for all evolving systems. In practice, however, it is often difficult to apply these results to particular systems of interest, since it is not clear how the details of complex biological systems translate into the few terms of the general equation. I will show how new mathematical tools allow us to build truly general theories -- requiring no simplifying assumptions -- that are nonetheless applicable to the biological details of particular systems. I will illustrate this approach by showing how it gives insight into the evolution of complex cooperation, and helps us to understand large scale patterns in the evolution of multicellularity.

The restless machinery of life Jessica Riskin Stanford University, USA

Ever since the beginnings of modern science in the 17th century, philosophers and scientists have used clockwork and related machinery as a metaphor to describe the universe and the natural world. The machinery model generally makes one think of something rigid, rote, constrained, something that ticks and whirs along without changing. However, hidden within this centuries-old clockwork-machinery metaphor is a tradition of people who have understood artificial machinery, and nature, very differently: as restless, responsive, and dynamic. These renegades include the German philosopher Leibniz who first described living things as “organisms,” and who said living organisms resembled clocks in their “restlessness”; the French naturalist Lamarck who was the first to present a theory of what we now call evolution; and the Austrian pioneer in quantum physics, Erwin Schrödinger, whose landmark 1944 manifesto What is Life was foundational to the new field of molecular biology. All three compared living beings to clockwork machinery but all three specified that they meant restless clocks. The talk will explore the history of restless machinery as a model for living things.

36 Abstracts: Selected Talks

Mycalesina in morphospace: how developmental bias shape evolutionary diversification in butterflies Oskar Brattström, Kwaku Aduse-Poku, Erik van Bergen, Vernon French, Paul M Brakefield University of Cambridge, UK

The developmental mechanisms that generate morphology can in theory bias the independent evolution of traits sharing ontogenetic pathways, making certain evolutionary changes more likely than others. The eyespots commonly found on butterfly wings each have concentric rings of differing colours and these classic examples of serially repeated pattern elements have been a focus for evo-devo research. Previous work on a model species of Mycalesina butterfly, Bicyclus anynana, has provided insights into the developmental control of individual eyespots using experimental evolution. These studies showed that the relative size of a eyespots on the same wing surface is highly flexible, whereas they are resistant to diverging in colour-composition, presumably due to shared developmental process. Here we survey the diversity in eyespot colour composition, across an extensive phylogeny of Mycalesina butterflies. Results are largely consistent with those from B. anynana, with the notable exception of a diverse clade on Madagascar. Thus, whereas each species generally has eyespots of similar colour-composition, consistent with a major role for developmental bias in shaping evolutionary diversification, most Malagasy Heteropsis species have gained independent control of eyespot colour-composition enabling a wider exploration of morphospace. We then investigated the formation of this novel phenotype by manipulating eyespot development in early pupal wings of H. iboina. The results show that the bias has been broken through a novel difference in the response of different areas of wing tissue to a conserved eyespot patterning signal. Our study demonstrates how development can potentially constrain the evolutionary independence of traits, but that at a macro evolutionary level the constraint can be released by an innovative developmental change allowing rapid morphological change and facilitating evolutionary diversification. Future directions include more detailed studies of how the bias was broken in the early Heteropsis radiation, and whether the bias found in most Mycalesina might be adaptive.

37 Fighting about frequency Karen Kovaka Virginia Tech, USA

Many controversies in biology, including many of those at the heart of the Extended Evolutionary Synthesis, have two distinctive features. First, they are not controversies about whether some hypothesis is right or wrong. Rather, they are controversies about how often a certain pattern or process occurs. Second, these controversies are rarely resolved in favor of one side or another. They are more likely to fizzle out or be abandoned. Examples of historical controversies that have these two features include disputes about (a) punctuated equilibrium versus gradualism, (b) sympatric versus allopatric speciation, (c) genetic drift versus natural selection, and (d) genetic versus extra-genetic inheritance. I provide an account of what justifies these kinds of controversies, and of how they contribute to science. This is an important project, because many criticisms of the Extended Evolutionary Synthesis rely on skepticism about the value of and justification for such controversies. More specifically, I argue for three claims: 1. There are serious questions about the value of controversies with the two features I described above. How can we rationally justify scientific controversies that rarely reach a resolution? Why should any scientist participate in such a controversy if they can know in advance, with high probability, how the controversy will end? 2. The areas of research encompassed by the Extended Evolutionary Synthesis include many controversies that have the two features I described above. 3. Despite the serious questions raised in (1), these kinds of controversies are rationally justified. Engaging in these controversies improves the quality of the explanations that biologists are able to give about the phenomena central to the controversies."

38 On leaders and followers: the evolution of phenotypic plasticity can lead the evolution of genetic evolvability more easily than vice versa Miguel Brun-Usan, Alfredo Rago, Richard A Watson University of Southampton, UK

In conventional evolutionary theory, phenotypic plasticity is often thought to be a genetically encoded feature which can be evolved autonomously. However, since both the effect of environmental conditions on phenotypes (the ExP map underlying plasticity) and the effect of genetic mutations on phenotypes (the GxP map underlying evolvability) depend on development, changes in one may affect the other. But which is the leader and which is the follower? By means of computational modelling, we show that it is much easier for the evolution of plasticity to lead the evolution of genetic evolvability than vice versa. Two complementary reasons explain this observed asymmetry: First, genetic and environmental inputs affect development in different mechanistic ways, and thus the GxP and the ExP maps, although related, do not exhibit identical dynamical nor variational properties. Second, the evolution of a GxP map is more limited than the ExP map. In the GxP map, selection on phenotypes is strong and selection on GxP maps is relatively weak. In contrast, the evolution of an ExP map does not suffer these limitations: an individual can be selected according to its ability to produce a distribution of phenotypes under different environmental conditions. We demonstrate that this asymmetry is consistent under different modelling strategies, parameter combinations and experimental setups, thus suggesting that phenotype-first scenarios may be the norm rather than the exception in biological evolution. Put in simpler words: phenotypes are more often the leaders, and genotypes are more often the followers."

39 Bird nests evolve flexibly in response to egg characteristics Catherine Sheard, Sally E Street, Mary C Stoddard, Camille Troisi, Andrew Clark, Susan D Healy, Kevin N Laland University of St Andrews, UK

Cultural evolution allows humans to dramatically modify their own selective environment (e.g. building shelter), a process known as ‘niche construction’. A classic non-human example of niche construction is avian nest building, which recent evidence in both the lab and the wild has shown can be socially transmitted. Here, we explore the co-evolution of avian nest building and egg traits. We compiled a database of nest structure and location for 8,231 species of birds, which we combined with a new database of egg colouration for 4,762 species and previously-published information on the egg shape of 1,400 species (Stoddard et al. 2017). Using Bayesian phylogenetic comparative methods, we show that nest structure and location is correlated with egg size, shape, colouration, and patterning. In many cases, however, it is the egg trait that evolves first, followed by a corresponding change in the nest characteristic. On a global scale, these egg traits are related to a variety of intrinsic and extrinsic factors. It has been previously shown that egg size is tightly correlated with body mass and that egg shape is related to adaptations for flight ability (Stoddard et al. 2017); we additionally demonstrate that egg colour and patterning is correlated with environmental variables and body mass, finding no support for traditional hypotheses related to brood parasitism or sexual selection. Taken together, this evidence runs against the conventional view of nests as highly conserved constraints on avian reproduction and instead suggests that nest construction is a flexible behavioural trait that can quickly respond to developmental sources of selection, including properties of the eggs.

Stoddard, M. C., E. H. Yong, D. Akkaynak, C. Sheard, J. A. Tobias, & L. Mahadevan. (2018) Avian egg shape: Form, function, and evolution. Science 356: 1249-1257.

40 Adaptive evolution NOT driven by natural selection (?!?). On individual responses, inclusive inheritance, and fleshing out the second term of that famous Price equation. Pim Edelaar University Pablo de Olavide, Spain

Evolutionary adaptation is typically accredited to natural selection only. However, natural selection can only adapt populations, and by itself has little to offer to locally maladapted individuals. Because of this, a large array of flexible individual responses to environmental variation have evolved that can help individuals increase their fitness. By means of a simple yet apparently comprehensive classification framework, I will derive how these can be grouped into three distinct classes (as appears to be confirmed from the social and economic sciences): adjustment of the phenotype (e.g. plasticity), adjustment of the environment (e.g. niche construction in the strict sense), and selection of the environment (e.g. habitat choice). I will then explore to what extent, and under what conditions, these flexible responses can additionally and independently drive adaptive evolution. I will do so by using the Price equation for evolutionary change as a context, focussing on the rather neglected second term describing parent-offspring resemblance, and touching upon discussions surrounding the Extended Evolutionary Synthesis, the Extended Phenotype, and inclusive inheritance. I think this view can help to shift our evolutionary thinking from an organism- and gene-centred position towards that of the evolution of the organism-environment interaction via genetic and alternative hereditary mean, while remaining consistent with basic evolutionary theory.

41 DNA methylation lends plasticity to the coloration of an African cichlid Sebastian Alvarado Queens College CUNY USA

Coloration is a salient trait across the animal kingdom that can allow an individual to become cryptic, conspicuous, or social. While some developmental patterns in pigmentation are static, others are dynamic to changes in their ambient environment. For example, seasonal variation in the Arctic allows mammalian species to gain and lose a white winter coat. Despite a great deal of study in developmental pigmentation patterns, little is known about how environmental cues shape the molecular mechanisms that allow an animal to change color. It stands to reason that the underlying mechanisms of a plastic trait must also be plastic. One approach to understanding these processes is through the lens of epigenetic modification and DNA methylation. DNA methylation of cytosine residues in gene promoters is a reversible modification that silences gene function in vertebrates. Since DNA methylation is involved in programming various cellular functions, it is likely that it facilitates molecular changes as pigment-bearing cells (chromatophores) change their composition during animal color changes. In this study, we used an African cichlid model system (Astatotilapia burtoni) with discrete reversible color morphs (blue and yellow) to dissect the underlying molecular processes that lend plasticity to animal coloration. We describe dynamic changes of DNA methylation within candidate regulators of endothelin signaling to demonstrate control over the function of yellow chromatophores. Our findings suggest that epigenetic processes such as DNA methylation lend plasticity to coloration, which is an important hallmark driving selection in Lake Tanganyika. Furthermore, since genetic diversity does not account for the phenotypic diversity seen in Lake Tanganyika, we propose that DNA methylation may contribute to the processes that have led to the adaptive radiation of cichlids in East African Great Lakes.

42 Unexplored maternal effect mediators, thyroid hormones, in an eco-evolutionary context Suvi Ruuskanen, Bin-Yan Hsu, Tom Sarraude, Antoine Stier University of Turku, Finland

Over the past decades, studies in maternal androgens and glucocorticoids have demonstrated that maternal hormones are key mediators of maternal effects, constituting an important source of transgenerational phenotypic variation. Nevertheless, maternal thyroid hormones (THs), despite their well-known importance on development in humans, have been largely neglected in ecological and evolutionary context. We studied environmental and genetic variation of these maternal effect mediators (biologically active T3, and prohormone T4), and associations between maternal THs and offspring development, phenotypic variation and fitness via experimental manipulation. Maternal yolk THs showed variation in relation to key maternal extrinsic and intrinsic factors (temperature and laying order), and T3 was moderately heritable. Experimental egg TH manipulations in five avian species revealed that maternal yolk THs influence hatching success, growth, physiology (metabolic rates, oxidative status) and aging markers, but the direction of the effects was highly heterogeneous across species. Context-dependent effects may be one explanation for such discrepancy, and is currently being tested. While no effect of elevated maternal yolk THs on subadult survival was detected, effects on reproductive success have not yet been assessed. However, many of the traits affected by maternal yolk THs are of known importance to fitness, pending selection on these maternal effect mediators. These results emphasize that maternal THs can be additional players of inclusive inheritance and their role e.g. in adaptive developmental programming should be further investigated.

43 Phenotypic plasticity facilitates colonization of savannas in Mycalesina butterflies Sridhar Halali, Hisham Ali, Paul M Brakefield, Oskar Brattström University of Cambridge, UK

Phenotypic plasticity has been considered to facilitate persistence of populations in new environments and the colonization of novel ecological niches. We test this hypothesis using butterflies of the genus Bicyclus in which many species show adaptive developmental plasticity with respect to seasonal changes. The rapid diversification of Bicyclus during the Miocene period has been linked to colonisation of expanding savannah grasslands. However, colonizing savannahs would likely have required a suite of adaptations to cope with seasonally fluctuating resource availability compared to the more stable ancestral forest habitats. Here we test two hypotheses: Firstly, savannah species will show a higher degree of plasticity than forest species. Secondly, plasticity may have facilitated the colonization of novel savannah habitats. We derived a plasticity index for 100 Bicyclus species by measuring the differences in eyespot size between wet and dry season morph in over 1000 museum specimens. Furthermore, we quantified the niche of each species using Environmental Niche Modelling based on museum location data and seven climate variables. The plasticity index shows a moderate phylogenetic signal (Pagel’s λ, 0.64) following an Ornstein-Uhlenbeck model of trait evolution. Savannah species exhibited a higher degree of plasticity, but forest species had higher variance (PGLS, p<0.01). Three seasonally linked climatic variables explained 70% of the variation, and correlated significantly with the plasticity index (PGLS, p<0.01). Phenotypic plasticity can modulate optimal season-specific trait expression in these butterflies which is necessary for survival, especially in the savannah. Also, the plasticity index is expected to evolve in response to habitat-specific adaptive optima. An observed higher variance among forest species is partly explained by phylogenetic inertia coupled with low cue reliability in more stable environments. Highly plastic species generally occupy extreme niches suggesting that plasticity may have been an important trait in facilitating colonization of savannas and facilitated diversification of Bicyclus.

44 Does thermal plasticity align with local adaptation? – An interspecific comparison of wing shape and size in sepsid flies Patrick T Rohner, Wolf U Blanckenhorn, David Berger University of Zurich, Switzerland

Although genetic and plastic responses are sometimes considered as unrelated processes, their phenotypic effects may often align because genetic adaptation is expected to mirror phenotypic plasticity if adaptive, but to run counter to it when maladaptive. The magnitude and direction of this alignment has further consequences for both the tempo and mode of adaptation. To better understand the interplay between phenotypic plasticity and genetic change in mediating adaptive phenotypic variation to climate variability, we quantified genetic latitudinal variation and thermal plasticity in wing loading and wing shape in two closely related and widespread sepsid flies. Common garden rearing of 16 geographical populations reared across multiple temperatures revealed that wing loading decreases with latitude in both species. Having evolved convergently, this pattern could be driven by selection for increased dispersal capacity in the cold. However, although allometry, sexual dimorphism, thermal plasticity and latitudinal differentiation in wing shape all show similar patterns in the two species, the relationship between the plastic and genetic response differed between them. While latitudinal differentiation (south to north) mirrored thermal plasticity (hot to cold) in Sepsis punctum, there was no relationship in Sepsis fulgens. While this suggests that thermal plasticity in wing shape may have contributed to mediate local adaptation in S. punctum, it also demonstrates that genetic wing shape differentiation and its relation to thermal plasticity may be complex and idiosyncratic, even among ecologically similar and closely related species. Hence, genetic responses can, but do not necessarily, align with phenotypic plasticity induced by changing environmental selection pressures. We discuss the potential role of plasticity in biasing adaptive genetic change and illustrate the usefulness of comparing replicated genetic and plastic responses of complex phenotypes in response to analogous selection pressures in revealing adaptive trait variation.

45 Macroevolutionary dynamics of phenological plasticity Stephen De Lisle, Erik Svensson Lund University, Sweden

Theory predicts that adaptive plasticity in fitness-related traits may play a key role in establishment in novel environments, persistence in changing environments, and allopatric speciation. Yet testing these hypotheses is difficult, especially at the macroevolutionary level, due to the inherent difficulty of measuring plasticity. We exploit recent methodical advances to estimate the strength of plasticity and the environmental dependence of selection, the two key parameters linking plasticity and local adaptation, from observational data from a largescale database of over one million individual records of adult damselflies and dragonflies and corresponding spring temperatures. Our aim is to elucidate the role that temperature-induced plasticity in timing of metamorphosis (phenological plasticity), a key life history transition, plays in macroevolutionary diversification. We first provide a validation of the methods using an independent dataset of phenotype and fitness from a single damselfly species, showing a striking correspondence between the observational estimate of the environmental dependence of selection and the direct estimate. We then use the observational approach to test two theoretically-motivated predictions: 1) the strength of plasticity should coevolve with the environmental dependency of selection and 2) the contribution of plasticity to local adaptation should be highest in extreme/recently colonized environments. Our analysis of over one million records from 49 species supports both predictions. First, we find correlated evolution of the strength of plasticity and the environmental dependence of selection. Second, we show that the contribution of phenotypic plasticity to within-species local adaptation increases during the recent, post-glaciation colonization of Northern regions. Our results suggest that phonological plasticity may have played an important role in temperate diversification in these insects.

46 Developmental regulation and evolution of nutrition-responsive growth: transcriptomic underpinnings of plasticity in horned beetles Sofia Casasa, Eduardo E Zattara, Armin P Moczek Indiana University, USA

Plastic responses to nutritional variation are ubiquitous features of developmental systems, yet our understanding of the mechanisms that regulate nutritional plasticity as well as bias its diversification is limited. We used comparative RNAseq analyses of Onthophagus horned beetles to better understand the transcriptomic mechanisms underlying the development and evolution of nutrition-sensitive plasticity of horns, male-biased weapons used in competition over mating opportunities. We investigated three Onthophagus species whose nutritional plasticity in horn formation ranged from the ancestral, moderately polyphenic state to the derived exaggerated polyphenic state, or alternatively, the secondary loss of nutrition responsiveness. We find that (i) the regulation of polyphenic development is partly shared across species regardless of the degree of nutrition-sensitivity, (ii) that diverse genes have been recruited into the evolution, and in particular the exaggeration, of nutrition-responsiveness, and (iii) that secondary loss of plasticity is paralleled by extensive loss of gene expression plasticity. Our results suggest that nutrition-mediated plasticity in morphology is underlain by a corresponding genome-wide magnitude of gene expression plasticity, and that both novel as well as pre-existing gene regulatory networks contribute to the elaboration of nutrition responsive development. By utilizing a comparative, unbiased, genome-wide assessment our work provides comprehensive insights into the regulation and evolution of nutrition-responsive development.

47 Involvement of maternal yolk thyroid hormones in the evolution of life history traits in birds: a comparative analysis Bin-Yan Hsu, Veli-Matti Pakanen, Tapio Eeva, Blandine Doligez, Ton GG Groothuis, Erkki Korpimäki, Toni Laaksonen, Asmoro Lelono, Pat Monaghan, Tom Sarraude, Barbara Tschirren, Robert L Thomson, Jere Tolvanen, Rodrigo A Vásquez, Suvi Ruuskanen University of Turku, Finland

As mediators of prenatal maternal effects, maternal hormones can bring about additional influence on evolutionary processes above and beyond genomic inheritance. Maternal thyroid hormones (THs), well-known for the essentiality on embryonic brain development, may harbour great evolutionary significance but our understanding to this is extremely limited. Lately, a few experimental studies in birds demonstrated that elevated maternal yolk THs within the natural variation influenced offspring development and growth. However, these studies showed striking differences between species, despite the essentially similar experimental designs. This thus calls for comparative studies to investigate the evolutionary relationship between maternal THs and life histories. In this study, we analysed maternal yolk THs from un-incubated eggs from 34 bird species across 17 families and 6 orders, representing a wide range of body size and life histories. Our phylogenetic comparative analysis suggested clear links between maternal yolk THs across a few main life-history continuums. First, migratory species deposited higher concentrations of THs in the egg yolks than residential species. Because of the regulatory roles of THs in seasonal activities, this association suggests the involvement of maternal THs in the evolution of migration or the circannual rhythm. Second, precocial species deposit larger amounts of T3, the bioactive TH, in the egg yolks than altricial species after controlling for species’ body masses. As the development of thyroid gland starts much earlier in precocial species than in altricial species, this association suggests the importance of maternal T3 to support the earlier development of the hypothalamic-pituitary-thyroid axis. In conclusion, our data suggested important involvement of maternal THs in the evolution of life histories. The patterns we identified should provide important basis for further experimental work.

48 Phenotypic divergence across spadefoot toad species mirrors within-species consequences and mechanisms of developmental plasticity Christoph Liedtke, Ivan Gomez-Mestre Doñana Biological Station, CSIC, Spain

Phenotypic divergence across derived lineages sometimes mirrors phenotypic plasticity in the common ancestor, suggesting that environmentally induced phenotypic changes may precede changes in genotype (i.e. ‘flexible stem hypothesis’) that can evolve into species divergence through genetic accommodation. Pelobatoids, the spadefoot toads, stand out as a good system for studying the role of plasticity in evolution because this group shows extensive developmental rate variation within and between species with predictable consequences on adult phenotype. Whereas Pelobates cultripes has retained the highly plastic developmental rate plesiomorphic of this group, this plasticity has been greatly reduced in Scaphiopus couchii as a consequence of adaptation to ephemeral breeding ponds. As a result of this adaptive process Scaphiopus couchii now presents several features that resemble an accelerated version of Pelobates cultripes, from overall morphology to fat storages to metabolic rate and endocrine regulation of development. To understand how the transcriptional regulation may have evolved between these divergent species, we have also characterized the gene regulatory network underlying developmental rate in both P. cultripes and S. couchii. We have investigated whether the same underlying transcriptional profile is conserved across species, and whether it varies between species in its environmental sensitivity. Spadefoot toads provide a good example of how canalization of ancestrally plastic traits can contribute to adaptive divergence and evolutionary novelty.

49 What the history of biology tells us about the Extended Evolutionary Synthesis Jan Baedke1, Daniel J Nicholson2 1Ruhr University Bochum, Germany; 2Konrad Lorenz Institute for Evolution and Cognition Research, Austria

Recently, there has been a growing number of calls for expanding evolutionary theory. Advocates of a so-called Extended Evolutionary Synthesis (EES) have argued that phenomena studied in evo-devo, epigenetics, and niche construction theory should be more seriously integrated into standard evolutionary theory. This includes the conviction that evolutionary investigations should focus less on genes and more on organisms and their reciprocal interactions with their environment. Unfortunately, this recent debate lacks a historical understanding of what kind of expansion the EES actually is (or could be) and on which theoretical traditions it draws upon. So far, attempts to historically contextualize the EES have focused primarily on showing how it differs from the Modern Synthesis (MS). We show that this historical perspective is limited. We argue that while the EES introduces a battery of concepts, theories, and explanatory standards to evolutionary biology compared to the MS, many of these are actually quite old. In fact, the theoretical core of the EES is not ‘post-MS’ but rather ‘pre- MS’. That is to say, the EES reintroduces claims into biology regarding the relationship between genes, organisms, and their environment that were already clearly articulated and vigorously defended by early twentieth-century biologists, especially organicists, dialectical materialists and (German) holistic biologists. We discuss six recurring theoretical claims: (1) ‘genes neither control nor determine development’; (2) ‘genes are not the exclusive repositories of inheritance’; (3) ‘organisms act as agents in their environments’; (4) ‘organisms and environments reciprocally constitute each other’; (5) ‘organisms extend into one another’; and (6) ‘organisms form together new collective individuals’. If none of these claims are new, then why did they not become incorporated into evolutionary theory much earlier? In the final part of our talk we shall attempt to answer this crucial question.

50 Towards a novel unifying theory of the genotype concept David Ricote Hernández, Ignacio Maeso Centro Andaluz de Biología del Desarrollo, CSIC, Universidad Pablo de Olavide, Spain

Understanding the Genotype-Phenotype (G-P) interdependence is fundamental for all biological fields. Surprisingly, discussions about the limits of the G-P dualism have been sidestepped by mainstream scientific discussions, despite the fact that both experimental practice (from mendelian genetics to -omics) and theoretical background (from phenomenology to information theory) have changed profoundly since Johannsen first coined the term genotype in 1911. Indeed, multiple unresolved contradictions between the original G-P formulation and its current use are showing that explicit theoretical reevaluations are urgently needed. In particular, the so-called extragenetic inheritance phenomena have not been possible to accommodate to the genotype concept due to the current lack of a neat distinction between genotype and genome, excluding inherited epigenetic configurations as fundamental and integral parts of the genotypical identity of organisms. Here, we provide a novel proposal towards a unifying theory of genotype based on 4 objective and experimentally verifiable criteria (self-templating, translatability, plasticity and inheritability) that are able to account for all potential sources of biological inheritance as genotypic constitutions, from molecular structures to social behaviour. Our proposal follows a different path than previous related frameworks (such as the replicator concept) because it does not rely on information to delineate the boundary between genotype and phenotype, but on the evolvability conferred by a growing number of nested and interrelated genotypical systems. By doing so, we provide new solutions to several long-standing problems, from the periodic resurgences of lamarckian phantoms to controversies on biological levels and units of selection.

51 Rethinking cybernetics in contemporary theoretical biology Flavia Fabris Egenis, the Centre for the Study of Life Sciences, University of Exeter, UK

In recent years, the contributions of cybernetics to the development of evolutionary developmental (evo-devo) biology have increasingly been recognised. The particular theories and models developed during the flourishing of cybernetics in the early 20th century laid the foundation for the systems approach, which is nowadays widely and fruitfully employed in molecular biology, genetics, genomics, immunology, developmental biology, and ecology. Nevertheless, no philosopher or biologist seems to know what cybernetics is, and often what they think they know they dislike: cybernetics is often identified with a reductive ‘machine conception’ of the organism and an engineering view of biology. However, once we understand what cybernetics is really about, we see such conceptions are mistaken and moreover that a cybernetic perspective can shed significant light on major discussions in current biology and its philosophy: in particular, on the fate of the Modern Synthesis in light of later developments in biology, the purpose and nature of evolutionary developmental biology, and disputes between those who emphasize a mechanistic conception of biology and ‘processualists’. Thus, my current research has two objectives: the first is to clarify the relationship between cybernetics and reductionism, and the second is to demonstrate the relevance of cybernetics to evo-devo. To accomplish the first objective, I will provide positive arguments for the thesis that, in contrast to the predominant view, cybernetic explanations within biology, when properly understood, are non-reductionistic, and do not have, at their core, any heavyweight metaphysical commitment to the mechanistic nature of life. To accomplish the second objective, I will disentangle the nature of cybernetics and reappraise its history in order to show how it offers new tools for approaching well-known neo-Darwinian controversies that have emerged in recent years.

52 The Extended Evolutionary Synthesis debate: some epistemological and sociological dimensions Alejandro Fábregas-Tejeda, Francisco Vergara-Silva Universidad Nacional Autónoma de México, Mexico

For the proponents of the Extended Evolutionary Synthesis (EES), the narrow and ‘gene- centric’ stance of the Standard Evolutionary Theory (SET) fails to capture the full gamut of causal processes entangled in the evolution of biological diversity (in particular, developmental bias, niche construction, developmental plasticity, and extra-genetic inheritance). In that sense, the EES denounces an incomplete ontology of evolutionary processes in the SET. The EES, as a conceptual framework that emphasizes organismal causes of development, inheritance and differential fitness, the role of constructive processes in development and evolution, and reciprocal representations of causation, delivers an extended ontology of evolutionary processes, including those that generate novel variants, bias selection, modify the frequency of heritable variation, and contribute to inclusive transgenerational inheritance. Here, we discuss two far-reaching dimensions of the EES debate: i) epistemological: by focusing on the explanatory goals of this framework, (purported) interdisciplinary integration, the conceptualization of the ‘organism-environment’ relationship, and tensions subtending the notions of ‘developmental system’, ‘organismal agency’ and ‘developmental niche(s)’; ii) sociological: by analyzing discursive strategies involved in the disagreement between the defenders of both the SET and the EES, the dispute for ‘symbolic and scientific capital’ (sensu Bourdieu), ethical-political implications of this new view of evolution, and other issues that can be unmasked by STS-studies and ideas coming from recent sociology of science. Finally, we argue that a practice-oriented analysis of the EES debate may bridge some of the epistemological and sociological dimensions previously discussed.

53 Beyond standard views of reproduction: pregnant females as individuals and their role in evolution Arantza Etxeberria, Laura Nuño de la Rosa, Mihaela Pavlicev Complutense University of Madrid, Spain

While reproduction is generally agreed to be a crucial feature of life on Earth, its roles in evolution have been neglected, either by considering it secondary to self- maintenance, or by reducing it to replication and copy making. In the latter case, the highly diverse modes of reproduction explored in evolution are examined as reproductive strategies with a single purpose: to maximize fitness, without considering the effects of diverse reproductive modes, for example, on relations between biological entities. Recently, there has been an increasing call to consider the material processes involved in reproduction, and evo-devo seems to be the natural disciplinary candidate for integration of reproduction and development into the structure of evolutionary theory. The theoretical implications of an “evo-devo of reproduction” have remained largely unexplored so far, also due to its focus on the evolution of body parts and their interactions. In this presentation, we will reflect on the crucial implications of recent research in evolution of therian reproduction, particularly of the origins of eutherian pregnancy, for the status of the pregnant female. This problem, namely if the pregnant female is a carrier of fully-fledged developing individuals or a single individual by itself, is becoming a focus of debate in the philosophical literature. Our goals will be to 1) question some common assumptions about biological reproduction and individuality for evolution; 2) explore, in the light of recent work in evo-devo, how mammalian reproduction has evolved as a merging of female and egg that involves major innovations at different (morphological, physiological, developmental, genetic) scales of organization; 3) consider what kind of individuals are involved in pregnancy from the physiological, evolutionary, and interactive perspective; 4) propose that, from a reproductive perspective, the pregnant female as a developmental stage can be considered tantamount to a evolutionarily novel kind of temporary individual.

54 Evolution of multicellularity: cheating done right Walter Veit University of Bristol, UK

For decades Darwinian processes were framed in the form of the Lewontin conditions: reproduction, variation and differential reproductive success were taken to be sufficient and necessary. Since Buss (1987) and the work of Maynard Smith and Szathmáry (1995) biologists were eager to explain the major transitions from individuals to groups forming new individuals subject to Darwinian mechanisms themselves. Explanations that seek to explain the emergence of a new level of selection, however, cannot employ properties that would already have to exist on that level for selection to take place. Recently, Hammerschmidt et al. (2014) provided a ‘bottom-up’ experiment corroborating much of the theoretical work Paul Rainey has done since 2003 on how cheats can play an important role in the emergence of new Darwinian individuals on a multicellular level. The aims of this paper are twofold. First, I argue for a conceptual shift in perspective from seeing cheats as (i) a ‘problem’ that needs to be solved for multi-cellularity to evolve to (ii) the very ‘key’ for the evolution of multicellularity. Secondly, I illustrate the consequence of this shift for both theoretical and experimental work, arguing for a more prominent role of ecology and the multi-level selection framework within the debate then they currently occupy.

55 The explanatory robustness of developmental types: a propensity view Cristina Villegas Complutense University of Madrid, Spain

In this talk I want to provide a philosophical frame for understanding the explanatory role of the properties of developmental systems explored in the field of evo-devo. A propensity interpretation of variability and evolvability is first introduced. These properties will be claimed to be probabilistic dispositions responsible for the variational patterns represented in evo-devo models. Following Roberta Millstein’s suggestion, in the context of the nature of fitness discussion, that a long-run propensity interpretation of biological probabilities is best understood as a type propensity interpretation, I will defend that the explanatory power of such evo-devo properties partially relies on their reestablishment of a certain kind of typological thinking without necessitating the – always controversial– reification of types. I will then argue that this feature provides evo-devo explanatory models with greater counterfactual robustness, enabling them to address how-possibly questions about the generation of variation and innovations more satisfactorily than populational explanations.

56 The cultural evolution of cultural evolution Jonathan Birch, Cecilia Heyes London School of Economics and Political Science, UK

Human cultures are not the only cultures. Other primates, as well as cetaceans and birds, pass learned skills down the generations. But human cultural evolution is different. It is cumulative: small improvements to skills build up. Moreover, the accumulation is fast: improvements spread rapidly, no longer tied to the timescale of biological generations. How does fast cumulative cultural evolution work? Where did it come from? And why is it the sole preserve of humans? One approach to these puzzles appeals to genetic adaptations that only humans possess. Heyes (Cognitive Gadgets, HUP, 2018) develops a novel approach on which the mechanisms of cultural learning are products of cultural, not genetic, selection. Here we turn this explanatory strategy towards the process of “cultural selection"" itself. In short, we propose that cultural selection itself evolves culturally. To motivate this idea, we highlight a crucial distinction between two types of cultural selection: CS1 and CS2 (Birch, The Philosophy of Social Evolution, OUP, 2017). CS1 is variation in biological fitness due to culturally transmitted variation, whereas CS2 is variation in cultural fitness (roughly, the number of learners a model recruits) due to culturally transmitted variation. We explain how ""cultural fitness"" should be defined, and we distinguish two cognitive “grades” of CS2, depending on whether model selection by learners is based on attentional or metacognitive processes. This allows us to pose the question: which type of cultural selection enables fast cumulative culture, and how did this type of process originate? We hypothesise that metacognitive CS2 (and not merely attentional CS2) is essential for cumulative culture. We further hypothesise that metacognitive CS2 was assembled by simpler, earlier processes of CS1. We draw on evidence from cognitive science, and especially developmental psychology, to support our hypotheses."

57 Evolution by natural induction Richard A Watson University of Southampton, UK

It is widely assumed that natural selection is the only possible source of adaptation occurring spontaneously in natural systems. This assumption short-circuits any debate over the significance of other processes in evolutionary change, e.g. the ‘active roles of phenotypes’. Induction, characteristic of learning systems, can also occur spontaneously in suitable dynamical systems. For example, Hebbian learning, sufficient for many non-trivial adaptive behaviours, occurs spontaneously in a physical system of particles connected by springs if the springs are slightly plastic (imperfectly elastic). When the system is repeatedly exposed to external forcing, its ability to find low-energy state configurations under future forcing is increased - not because it was designed or evolved to do so, but because of the direct action of differential forcing on its internal organisation. Biological networks with evolving components are capable of such inductive learning. This involves the action of natural selection only on individual links in the network using only local selective feedback (equivalent to imperfect elasticity). The result is that evolving systems, e.g. gene-regulation networks, can increase their evolvability without being selected for this function. Interestingly, evolutionary models cannot exhibit induction unless they include active phenotypes (e.g. dynamical developmental processes). We thus conclude that evolution in natural systems proceeds by a combination of natural selection and natural induction, that natural induction is an adaptive mechanism in its own right (not a product of natural selection), that the active roles of phenotypes are necessary for evolving systems to exhibit natural induction, and that the adaptation provided by natural selection alone is significantly limited in comparison. These conclusions support the view that the active roles of phenotypes are important to natural evolution, not because they are proximal details, but because they enable access to a different adaptive mechanism that is absent in the standard model.

58 Generalized pathways to specific outcomes: stress links maternal environment and adaptive offspring trait variation Ahva Potticary, Renee Duckworth The University of Arizona, USA

The evolution of an adaptive maternal effect requires complex coordination of a maternal response to environmental cues, coevolution of maternal physiology and offspring embryonic development, and subsequent matching of adult offspring phenotype to a future environmental context. How can such complex multi- generational and multi-environmental coordination evolve? One possibility is that evolution of maternal effects does not require de novo evolution of maternal physiology, but instead coopts pre-existing pathways to link ecological cues to aspects of maternal physiology that are primed to influence offspring development. An obvious candidate for such cooption are maternal stress pathways, as they are highly conserved across taxa and reactive to environmental context, providing a generalized cue of environmental quality. We test this idea using a known adaptive maternal effect in western bluebirds, where competition over a limited breeding resource (cavities in trees) results in mothers influencing offspring dispersal strategies. We combine 17 years of data on natural variation in female corticosterone levels and environmental conditions with experimental manipulation of breeding resources to determine whether multiple cues influence maternal stress, and whether maternal stress, in turn, induces the maternal effect in offspring. We found that females had higher levels of baseline corticosterone when breeding on territories that had either heightened competition for nest cavities or more difficult foraging conditions during inclement weather. Moreover, we found that irrespective of the environmental cue, the maternal effect was associated with stress females experienced during oogenesis: females with lower corticosterone levels produced more philopatric sons compared to females with higher corticosterone levels. These results suggest that stress pathways are a general link between ecological cues and offspring development and may foster the evolution of maternal effects by enabling species-specific stressors to produce adaptive variation in the offspring generation.

59 Evolution of migratory behavior by barnacle geese: the roles of experience, memory and social learning Thomas Oudman, Ingunn Tombre, Paul Shimmings, Graeme Ruxton, Jouke Prop NIOZ Netherlands Institute for Sea Research, The Netherlands; University of St Andrews, UK

Many migratory animal populations adjust their migratory decisions to changes in spring phenology – some even stop migrating altogether. In many species, individuals may learn their migratory strategy by following others. This opens up the possibility for the cultural inheritance of strategies that are adjusted in response to individual experiences. Forty years of observations at multiple locations show strong changes in the use of staging sites by barnacle geese along the Norwegian coast, on their way from the UK to the breeding grounds on Svalbard. Barnacle geese are highly social animals, and the choice of staging site strongly affects their reproductive success. This choice is not straightforward. A reconstruction of foraging conditions at the different staging sites from daily local temperature and radiation data showed that annual variation was high, and that each staging site showed a different trend over the last forty years. To understand how barnacle geese navigate this dynamic environment, we designed a set of individual-based simulation models, each of them assuming a different set of processes to underlie the choice of staging site. These include group size, reliance on individual memory and exploration, and different social copying rules (e.g. based on chance, relatedness, previous success or experience). This allowed us to explore how each of these processes and their interactions would affect the population response to the observed environmental variation. By comparing these simulated responses to the actual population trends, we infer which of these processes are most likely to underlie the migratory behavior of barnacle geese. We find that both social and individual learning required assumptions for the simulations to resemble the actual population trends. The simulations also point out additional types of field observations that will help to discern the processes by which migratory behavior adjusts to changing environments in other species.

60 Theory-based ecology and niche construction Liz Pásztor Biological Institute, Eötvös Loránd University, Hungary

Niche construction has been discussed in several ecological contexts. The theory of niche construction differs from the standard theory of evolution in focusing on the environmental effects of the organisms and on the selection processes induced by these modifications. It can be linked easily with the theory of ecosystem engineering by considering the selective effects of the nutrient cycles modified by the organisms. This was modeled directly in a framework called „ecological niche construction”. The selective effects of niche construction can also be integrated to community ecology by considering selection in the context of species interactions. However, these achievements have not revealed the common roots of the theory of niche construction and the Darwinian theory of evolution and ecology. A theory-based ecology (TBE) has been developed, which is founded on Darwinian principles and defines ecological niches deductively. Ecological niches are described in terms of population regulation including both positive and negative interactions among populations. As TBE considers the environmental factors regulating population growth explicitly the impact of population growth on these factors and reciprocal causation are inescapable elements of this theory. Thus, evolution by niche construction follows from first principles. This will be explained and illustrated in this talk.

61 The Hot Spring Hypothesis for the origin of life: implications for niche construction theory in the first major transition in evolution from non-life to life Bruce Damer, David Deamer, John Odling-Smee The Biota Institute, USA; University of California, Santa Cruz, USA

Until recently the origin of life was thought to have occurred deep in the oceans at hydrothermal vents as some energy sources and building blocks are available there. However, chemical, thermodynamic and combinatorial barriers have led to an alternative hypothesis situated in an updated version of Charles Darwin’s “warm little pond”: fresh water volcanic hot spring pools that undergo cycles of hydration and dehydration. The hypothesis involves construction of proto-niches that self-assemble from membranous structures and guide chemical reactions related to synthesis and evolution of polymer systems. Combinatorial selection amplifies functional polymers within protocells ultimately incorporating heritable genetic memory. This hypothesis has been tested experimentally in laboratory and volcanic hot spring conditions suggesting that the first major transition from non-living compounds to living molecular systems will be characterized by: 1. A cycling natural “engine” utilizing crowding to shape probabilities analogous to loading the dice, a network effect of interacting polymers in protocell aggregates, and Darwinian selection arising through an emergent memory system; 2. Communities of dividing living cells emerging from earlier self-assembling transitory protocells; 3. The first genotypes arising through combinatorial selection of proto-phenotypes; 4. Vertical descent in genetic evolution originating from horizontal network sharing of the products of chemical evolution. The hot spring hypothesis not only provides an experimentally-accessible predictive chemical framework for the transition to life but also a simplified exploratory laboratory which could provide insights into the fundamental operating principles of evolution itself. An international community has assembled around the testing of this hypothesis and we will present compelling empirical results by the date of the meeting.

62 Developmental bias and morphological convergence in Anolis lizards Nathalie Feiner, Illiam Jackson, Tobias Uller Lund University, Sweden

Instances of convergent evolution provide insights into the processes that give evolution directionality. While convergent phenotypes are usually attributed to similarity in selective pressure, it remains unexplored to what extent convergence is shaped by developmental biases. Yet, the fact that different morphologies can be functionally equivalent (many-to-one mapping of phenotype-to-fitness) suggests that morphological convergence is partly caused by biases in how development responds to genetic or environmental perturbation. After colonizing the four main islands of the Greater Antilles, Anolis lizards have independently and repeatedly evolved into six ecomorphs that are well adapted to running and climbing in certain microhabitats (e.g. grass, twigs, tree trunks or tree crowns). In contrast, Lesser Antilles and mainland Anolis species do not obviously fall into ecomorph categories, despite that they are typically found using similar habitat. Using a morphological dataset (µCT-scans) of 350 Anolis species, we quantified the extent of divergence and convergence in the relative size and shape of limbs, pelvic and pectoral girdles between independent radiations of Anolis species. We quantified if morphological divergence between species tends to fall along the same axis that describes maximum variation within species. Furthermore, we tested the prediction that adaptive divergence and convergence can be directed by developmental plasticity. In an experimental setting with twelve species raised either in a ‘running’ or ‘climbing’ habitat, we established to what extent the relative size and shape of limbs and girdles respond to mechanical stress induced by locomotion. These data allow us to assess to what extent the patterns of adaptive divergence and convergence in morphology are channelled by the process of development.

63 Measuring developmental bias: quantitative genetics meets developmental biology Lisandro Milocco, Isaac Salazar-Ciudad University of Helsinki, Finland

The patterns of inheritance of complex traits such as shape can be approximated by quantitative genetics models. The models from this field have the core implicit assumption that all phenotypic variation is symmetric, arising with equal likelihood. There is however evidence that this assumption does not hold in many cases, a phenomenon which has been called developmental bias. The term bias implies that some phenotypic variation is more or less likely to arise, compared to a base expectation, due to the dynamics of the developmental system involved in producing that variation. The importance of this developmental bias in determining the direction of evolution in the short term, however, remains heatedly debated. In this work, we use the canonical equation from quantitative genetics, the multivariate breeder's equation, as our base expectation for evolutionary change. We compare that base expectation with the observed evolutionary change in an evolutionary simulation. The latter combines a realistic genotype-phenotype map, from a tooth development model, with a population-level model with sex, mutation and selection on the phenotype. The deviations between the expected and the observed change are a measure of the role that developmental dynamics play in constraining the directionality of morphological evolutionary change. We found that the breeder’s equation often fails to accurately predict evolutionary change because the assumptions of quantitative genetics do not always hold for the morphological variation arising from development. We quantify both the frequency and magnitude of these inaccurate predictions, which can be as common as 31% for some traits and of large effect size. We found that these prediction errors most commonly occurred for traits that are determined by the highly nonlinear reaction-diffusion mechanism involved in tooth morphogenesis. These results give a tangible measurement of the effect that developmental interactions have in determining the direction of evolution in the short term.

64 The evolutionary potential of environmentally induced phenotypes Reinder Radersma, Daniel WA Noble, Tobias Uller Lund University, Sweden

Environmentally induced phenotypes have been proposed to initiate and direct adaptive divergence. The evolutionary potential of environmentally induced phenotypes depend in part on how well plastic responses are aligned with the additive genetic variance and covariance among traits, which is captured by the genetic covariance matrix (i.e., G matrix). Using meta-analyses, we show that the direction of plastic responses to novel environments are generally aligned with the major axes of standing genetic variation. Selection for or against environmentally induced phenotypes is therefore likely to be effective. Furthermore, reciprocal transplant studies reveal that evolutionary divergence tend to follow the direction of plasticity. These results suggests that environmentally induced phenotypes may commonly initiate adaptation to novel environments. However, the alignment between the direction of plasticity and the main axis of genetic variation suggests that developmental bias in response to environmental and genetic perturbation often may be similar, making it challenging to distinguish between ‘plasticity-first’ vs ‘genetic constraints’ scenarios of evolutionary divergence.

65 Developmental interactions established during speciation leave a lasting trace in descendent populations Laura Mears, Tom Ezard, Richard A Watson University of Southampton, UK

Relationships among traits change the way that populations respond to selection. Traits untethered by developmental interactions are free to evolve independently to match the adaptive landscape. But, where developmental interactions exist, selection on one trait elicits a response in others. To extract clues about how developmental programming affects population divergence, we combine an adaptive dynamics framework for modelling speciation with an individual-based model of evolvable pleiotropic constraints. Using this system, we study how changes in development alter the way that populations move over adaptive landscapes. Evolving independent traits under strong selection results in among-individual variation orthogonal to the direction of divergence. This orthogonal variation persists until populations reach static adaptive peaks, at which time trait independence allows for rapid removal of non-adaptive variation. But when evolvable interactions connect traits, among-individual variation turns to align with the direction of divergence. We show that this directionality, and the underlying developmental architecture, persist even after populations reach stable adaptive peaks, preventing the loss of non-adaptive variation. Thus, the descendent populations remain in alignment with one another, revealing the trajectory they followed as the ancestral population diverged.

66 What kind of evolutionary novelty (if any) could be language? Salvatore Ivan Amato, Giuliana Pulvirenti, Alessandra Falzone University of Messina, Italy

The concepts of Novelty and Homology are as central in Evo-Devo approaches and in the emergent Evolutionary Extended Synthesis as problematic and intertwined. These could indeed be applied to different level of biological organization analysis; from genetic and cellular to morphological as well as behavioral ones (Müller & Newman 2005; Minelli 2003). Studies at the phenotypic level have been able to identify several generative mechanisms responsible of the emergence of evolutionary novelties (genetic, epigenetic, behavioral, environmental). This theme has rightfully assumed a great relevance within the biolinguistic area of research opening a theoretical debate among Hauser-Chomsky-Fitch Vs Pinker-Jackendoff regarding the problem of the emergence of the language faculty in our lineage: is it a unique feature of our human nature or a mixture of several traits both shared and unique? Does it represent the result of a gradual process of adaptation involving several components of the human brain, or is it the result of a genetic mutation with a discontinuous outcome at the phenotypic level? We think that the biolinguistic field is lacking of an adequate problematization of the concept of novelty. This may be achieved widening the dialogue with Evo-Devo/EES area of research, especially considering developmental plasticity and niche construction processes (Pigliucci & Müller 2010; West-Eberhard 2003). An analysis of the role that genetic, epigenetic and ecological factors may exert on the formation of a phenotype, as well as the causal mechanism responsible of the emergence of evolutionary novelties may foster a better characterization of the linguistic phenotype and help elucidating the ways in which such phenotype comes to be during the course of ontogenesis. This sort of investigation could also offer novel insights about the evolutionary events and the possible mechanisms that have driven its emergence during the phylogenesis.

67 It’s a horn, a wing, a helmet: the role of wing serial homologs in insect innovations Yonggang Hu, David M Linz, Armin P Moczek Indiana University, USA

Understanding the origin of novel complex traits is a foundational challenge in evolutionary biology. The most commonly used definition of novelty entails the absence of homology to ancestral traits. This definition, however, is increasingly difficult to reconcile with empirical findings across diverse taxa, which instead emphasize the role of differential re-purposing of conserved developmental modules outside their traditional developmental context as a dominant route to innovation. Yet how descent with modification in developmental evolution may lead to morphological innovation remains poorly understood, including the role of pre-existing gene networks in enabling, biasing, or hindering such innovation. Here we employed RNA interference mediated analysis of wing gene function with Hox-gene mediated transformations of body segments to investigate the origin of the thoracic horns of scarabaeine beetles, one of the most dramatic classes of secondary sexual traits in the animal kingdom. We find (1) that thoracic horns derive from bilateral source tissues that fuse to form a single medial outgrowth during development, (2) that diverse wing genes are functionally required for instructing this process, (3) and that in the absence of Hox-input thoracic horns transform into ectopic wings. Combined our results provide strong evidence for the serial homology between thoracic horns and insects wings, and raise the possibility that other insect innovations may similarly derive from wing serial homologs.

68 Beyond Darwin: how every individual adapts to its internal perturbations? Yoav Soen Weizmann Institute of Science, Israel

A critical gap in the theory of adaptation is the inability to explain how every individual is able to cope with an unimaginably large number of epigenetic and symbiotic variations, including many that are encountered in novel combinations and are unique to that individual. Overcoming many of these novel, individual-specific variations cannot be explained solely by natural selection, because the latter only accounts for adaptation of the population as a whole. In other words, we understand how populations adapt to changing environments, but we have no idea how an individual adapts to stochastic internal changes of its own. To address this gap, we proposed a non-traditional principle of individual-specific adaptation, capable of extending the population-based adaptation by natural selection to dynamic adaptations that are newly forming within every individual cell, animal and plant. In some specific cases, acquired adaptations can even be inherited, thus enabling progressive improvements in a few generations. To investigate the validity of this Lamarckian theory, we established unique experimental systems for testing the predictions of the theory in flies (D. melanogaster) and in cultured cells. Both systems provide multiple lines of evidence supporting a capacity to adapt by exploratory dynamics over time in an individual. I will explain this concept of individual-specific adaptation and present substantial experimental and theoretical evidence in support of this hypothesis.

69 How the beetle head got its horns: integrating a novel trait into a conserved trait complex David M Linz, Armin P Moczek Indiana University, USA

The origin of novel traits is among the most intriguing and enduring problems in evolutionary biology. In particular, how novel complex traits are initiated and incorporated within existing trait complexes without disrupting ancestral functions remains poorly understood. The highly diversified horns of Onthophagus beetles represent a recent evolutionary invention seamlessly integrated within the adult beetle head, an ancient trait complex deeply conserved among insects. We sought to investigate the developmental genetic network that instructs postembryonic head formation in beetles, as well as identify the component parts that allowed horn induction and positioning to become integrated into the dorsal head while maintaining the integrity of head formation as a whole. To do so we executed a unique, fine-scale RNAseq approach in both horned males and hornless females. This approach allowed us to independently assess the transcriptional repertoires of six distinct head compartments in both sexes. Contrasting these transcriptomes within and among sexes then allowed us to (i) identify the transcriptional underpinnings of horn induction and positioning in horn forming head regions as well as (ii) transcriptional changes occurring in neighboring head regions putatively underlying compensatory, accommodating development in the remainder of the head. Our results identify suites of genes critical for patterning concise regions of the dorsal head and show that diverse embryonic patterning genes have been repurposed to execute central roles in instructing the formation of the postembryonic head, with some functions paralleling those executed in the embryo, while others appear largely decoupled. Further, we identify points of integration where re-use of pre-existing head patterning mechanisms enabled the positioning of a novel trait – cephalic horns – within an otherwise conserved trait complex – the dorsal head. We discuss our results by examining the roles of - and interplay between - conservation and innovation in the origins of novelty and diversity.

70 Niche construction affects the consistency and strength of natural selection Andrew Clark, Dominik Deffner, Kevin N Laland, John Odling-Smee, John Endler University of St Andrews, UK

Consideration of the properties of the sources of selection potentially helps biologists to account for variation in selection. Yet, while investigations conducted over the last 40 years have generated many thousands of individual quantitative estimates of the direction and strength of natural selection in the wild, associated analyses of such environmental sources of selection and variation in selection are rare. Here we explore how the variability of natural selection is affected by organisms that regulate environmental variables through their activities (whether by constructing components of their local environments such as nests, burrows, or pupal cases, or by choosing suitable resources). Specifically, we test the prediction that organism-constructed sources of selection that buffer environmental variation will result in reduced variation in selection gradients, including reduced variation between (i) years (temporal variation), and (ii) locations (spatial variation), and (iii) weaker directional selection, relative to non-constructed sources. Using compiled datasets of 1045 temporally replicated, 257 spatially replicated, and a pooled dataset of 1230 selection gradients, we find compelling evidence for reduced temporal variation and weaker selection, in response to constructed compared to non-constructed sources, and some evidence for reduced spatial variation in selection. These findings, which remained robust to alternative datasets, taxa, analytical methods, definitions of constructed/non- constructed, and other tests of reliability, suggest that organism-manufactured or chosen components of environments may have qualitatively different properties from other environmental features.

71 Linking coral morphological traits and reef fish community structure Luisa Fontoura, Stephanie D’agata, Kyle JA Zawada, Andrew H Baird, Mariana Álvarez- Noriega, Rachael Woods, Michel Kulbicki, Maria Dornelas, Joshua S Madin, Joseph M Maina, Elizabeth Madin Macquarie University, Australia

The synchronized evolutionary expansion of small body size reef fishes with coral complex forms in shallow waters suggests that branching corals might have had an essential ecological role of providing shelter for fish to utilize. Losing structurally complex coral forms can reduce shelter availability significantly for reef fishes, limiting their ability to avoid predators and increasing competition for settlement area during early-life stages. In this study, we aim to understand how changes in coral morphological complexity, captured by coral convexity (a novel morphological trait), affect the structure of reef fish assemblages. We use a multi-year dataset to track coral community and reef fish assemblage structure, capturing a severe coral mortality event that shifted the complexity of the coral community. We show that juvenile fishes, rather than adults, are more responsive to shifts in morphological traits at the coral community scale, a likely consequence of the higher vulnerability of juvenile reef fishes due to competition for refuge in their early versus later life-stages. Changes in coral morphological complexity explained 23% of juvenile reef fish decline, however the magnitude of this effect varied by fish functional group. Although coral complexity explained the decline of small body size juvenile fishes with restricted home ranges, we found no relationship between changes in coral complexity and functional groups of juvenile fish with large body size and wide home ranges. Differences in the responses among fish functional groups due to loss of coral complexity suggest that structurally complex coral species construct ecological niches that are occupied by vulnerable reef fishes with small body size and restricted mobility. Importantly, our results shed light on an evolutionary link between fish vulnerability and coral morphological traits.

72 Microbial community structure and function driven by infaunal niche construction in intertidal sediments Adam Wyness, Andrew Blight, Patricia Browne, Matthew Holden, David M Paterson University of St Andrews, UK

The Extended Evolutionary Synthesis theory predicts that, through their niche construction, organisms greatly influence community structure, stability and productivity, and co-direct evolutionary responses to selection in ecosystems (Odling- Smee et al., 2003; Laland et al., 2015). The degree to which niche constructing activities are complementary or conflicting impacts the magnitude and stability of biochemical and physical gradients formed within sediment profiles. These gradients generated by the ecosystem engineers are selective pressures to the microbial communities in sediment. The aim of this work was to test how complementary and conflicting ecosystem engineering impacts microbial community diversity and function and the consequences on selected sediment functions such as surface stability, nutrient cycling and primary production. A well-established mesocosm system of intertidal sediments with treatments included the presence of infauna (natural bioturbation with either one or both of Corophium volutator, Nereis diversicolor), or manual turbation (proxy for bioturbation). Dissolved oxygen and redox potential gradients were measured using microelectrodes. Microbial community composition at the sediment surface and various depths was analysed using high throughput 16S rRNA sequencing, and functional metagenomes predicted. Evidence for the cooperative ecosystem engineering between the infauna of dissolved oxygen and redox gradients compared to individual infauna treatments was observed, and coincided with in shifts in diatom and prokaryote community composition. Surface communities were compositionally and functionally different between infauna treatments, with dominating influences observed in mixed treatments rather than an intermediary response. These results illustrate that niche construction is a powerful evolutionary force where organisms modify their own and others’ selective environments. Furthermore, observed differences in microbial community response to physical and bioturbation hold significant inference for the impact of anthropogenic disturbance on biogeochemical cycling by microbial communities. Now that community level effects have been shown, this work will be taken further to examine ecological trajectories among target bacterial populations.

73 Keeping close neighbours, corals are spatially clustered within and across species Maria Dornelas, Shane Blowes, Tory Chase, Grace Frank, Mia Hoogenboom, Janine Illian, Oscar Pizarro, Andrea Walker, Stefan Williams, Kyle Zawada, Joshua Madin University of St Andrews, UK

Spatial distributions of sessile niche constructing species determine coexistence patterns. Major biodiversity models make two assumptions about spatial distributions: that individual species are spatially clustered, and that the distributions of different species are independent. These assumptions are common across all models that aim to predict the global distribution of biodiversity. However, there is remarkably poor empirical knowledge to support these assumptions. Moreover, to our knowledge these patterns have never been quantified on reef corals, the niche constructors of the most diverse marine ecosystem on the planet. We used AUV collected imagery to build 3D topographic maps of a 7500 m2 of a reef at Lizard Island, Great Barrier Reef. These models were annotated with the species identities and locations of over 11,000 coral colonies, identified in situ by divers, to resolve the spatial distribution of the 120 species in the surveyed area. We analysed these spatial distributions and quantified intra and inter specific clustering. We show that across a range of scales, all but the rarest species are spatially clustered with conspecifics, a finding that is consistent with model assumptions. However, neighbouring species are remarkably diverse, and we find evidence for interspecific clustering rather than independent distribution of different species. We suggest that these patterns are likely to arise from niche constructing mechanisms, such as clustering at the recruitment stage, when the physical structures built by corals alter flow regimes in ways that facilitate settling of coral larvae.

74 Holed-up: nesting in a cavity as an evolutionary driver of parental care in fishes and birds Karina Vanadzina, Michael Webster, Kevin N Laland University of St Andrews, UK

Parental care is any form of behaviour that increases the survival rate or growth of a parent’s offspring. Despite the wide taxonomic spread of parental care behaviours, we know little about the evolutionary drivers of stable bonds between parents and their young. The use of spatially restrictive breeding habitats does promote parent–offspring association and could therefore contribute to the origin of care in clades such as ray- finned fishes where more than 75% of families abandon their eggs shortly after fertilisation. My analysis of reproductive behaviours in North American freshwater fishes shows that cavity-spawners are 34 times faster at evolving parental care from the ancestral state of no care than species that spawn in open waters. The evolutionary impact exerted by the breeding habitat can also extend to clades where the majority of species provide advanced care and raise their young in stable microhabitats such as nests. For example, it has been hypothesised that hole-nesting passerines take care of their young for longer periods and give rise to larger clutches when compared to birds that nest in the open. This study is the first to explore the global distribution of parent– offspring bonds in relation to breeding grounds in ray-finned fishes and passerine birds. I use species-specific accounts of reproduction and the latest phylogenetic comparative methods to analyse the co-evolution of parental care traits and breeding habitats. My work fits within a framework of a larger project on niche construction that tests whether artefact building and habitat choice generate consistent and predictable selection pressures on a macro-evolutionary scale across a broad taxonomic spectrum of animals.

75 Trait specialisation, innovation and the evolution of culture in changing environments Dominik Deffner, Anne Kandler Max Planck Institute for Evolutionary Anthropology, Germany

Organisms often respond phenotypically to environmental challenges by innovating and adopting novel behavioural variants. Behavioural (or 'cultural') variants are alternative ways to solve adaptive problems, such as finding food or building shelter. In unpredictable environments, individuals must both be able to adapt to current conditions but also to cope with potential changes in these conditions, they must ‘hedge their evolutionary bets’ against the variability of the environment. Here, we loosely apply this idea to the context of behavioural adaptation and develop an evolutionary model where cultural variants differ in their level of specialization, i.e. the range of environmental conditions in which they provide fitness benefits: generalist refers to large ranges whereas specialist refers to small ranges. We use a Moran model (with additional learning opportunities) and assume that each individual’s propensity for asocial learning is genetically determined while the characteristics of cultural variants are changed through asocial learning. This model demonstrates that adjusting the level of specialisation allows organisms to navigate the trade-off between fast initial adaptation and the potential for long-term improvements. In situations with many learning opportunities, no adjustment of the innovation rate (propensity to learn asocially) is required. Fast adaptation is guaranteed by solely adjusting the level of specialisation of cultural variants. Few learning opportunities, however, require both processes —innovation and trait specialisation — to work hand in hand. To explore the effects of different modes of innovation, we contrast independent invention and modification and show that a large proportion of modifications improves both short- and long-term adaptation. Similarly, inaccuracies in social learning provide another source of variant variation that facilitates adaptation after an environmental change. However, unfaithful learning is detrimental to long- term cumulative culture. Our results demonstrate that trait characteristics themselves can play a major role in cultural dynamics and influence the evolution of learning strategies.

76 A New Caledonian crow's-eye-view on how to develop probing intelligence: the evolution of tool-use and tool-making as embodied, distributed and socially situated perception action processes Joanna E Fairlie1, Russell Gray2, Mei-Shin Wu2, Natalie Uomini2 1associated with the Max Planck Institute for the Science of Human History, Germany as a guest researcher, 2Max Planck Institute for the Science of Human History, Germany

The behaviours of the tool-using Corvus moneduloides (New Caledonian crows or NCCs) are often mentioned in support of existing theories about the evolution of Homo sapiens cognition. Theories about their behaviours have been interpreted within anthropological frameworks that have rarely considered modern evolutionary biology, neuronal plasticity, ecological or developmental issues, or niche construction. Here we discuss fresh cognitive models and behavioural approaches which allow us to consider the NCCs in their own right as modern birds, but also as members of social groups that have developed their own cultural evolutionary pathway. We use a new methodology for coding gestural sequences developed by the first author from footage collected by the second author during two field trips in Grand Terre, New Caledonia. We set out to establish whether repeated interactions between fledged juvenile birds and accompanying adults in the context of using tools to forage, can be considered as a recurring form of learning or even a teaching process. The theory models include ecological psychology and dynamic cognition. They allow us to consider learning processes as dynamics of directed, selective, variable and adaptive physiological and neuronal plastic change which can only occur when the learner is embedded in an appropriate social, cultural and ecological mileu. Our qualitative results show consistent gestural patterns embedded in interactions between NCC juveniles and adults. The patterns suggest that juveniles mature into expert tool-users because they are cognitively capable of learning. They benefit from having activities graded for them through time by their adult mentors, and from their own initially unrewarded perseverance. We end with some brief speculation about the implications of our NCC- centred findings for evolutionary anthropology.

77 Generalized Darwinism revisited: how a new synthesis changes our view on cultural evolution Karim Baraghith Düsseldorf Center for Logic and Philosophy of Science, Heinrich-Heine University, Germany

Theories of cultural evolution, dual inheritance theory or generalized Darwinism have been around for some decades now. The idea is that cultural systems obey the same abstract principles of variation, selection and reproduction in their development as biological systems - organisms, genes and populations - do. As well as heavy enthusiasm, like the ability to provide a long expected scientific synthesis for the social sciences (resembling the “modern synthesis” for the life sciences, cf. Mesoudi 2011), such approaches faced severe skepticism and critiques from various directions. Exceptional aspects of cultural transmission - like conformist bias, blending inheritance or guided variation - led some researchers doubt that cultural evolution is truly “evolution”. Furthermore, the identification problem of cultural units or variants (memes) and of cultural “generations” seemed to undermine the application of population dynamical or phylogenetic methods for the study of cultural phenomena. But meanwhile, our biological understanding of "evolution" itself has evolved in various directions as well. Aspects like plasticity, epigenetic regulation, inclusive inheritance and niche construction seemingly change our view on evolution in direction of an “Extended Synthesis” (cf. Pigliucci/Müller 2005). In this paper, I am going to show that most approaches in the field of cultural evolution today still adopt and assume a very classical notion of evolution as e.g. given by population genetics. I want to argue that many critiques are loosing grip, when we involve a more fine grained notion of cultural “evolution”, which takes into account these new findings and makes evolutionary models much more applicable to the study of sociocultural systems. I am going to demonstrate that in three case-studies.

78 Stress-induced mutagenesis systems and the adaptive immunity Bartlomiej Swiatczak University of Science and Technology of China, China

Stress response is a basic survival reaction in all living things from bacteria to humans. In the case of evolving cells, it often takes the form of genome restructuring, which amplifies variation and accelerates the process of collective adaptation of these cells to altered environmental conditions. While stress-enhanced evolution is suppressed in most bodily cells due to their subordination to somatic duties, certain cells of multicellular organisms like cancer cells and adaptive immune cells do evolve and along with reclaiming their evolvability, they reemploy their genome restructuring capacities to diversify their protein products in response to stress or danger. The aim of the presentation is to show that due to their potential to provide population-level protection from environmental threats the conserved mutagenesis systems act as primordial systems of defense for these populations. In fact, stress and danger responses have been considered key to immunity and unicellular communities are already able to generate ones. Harnessed by vertebrate organisms, these ancient danger response systems extend their protective effects from the cell population to the organism as a whole. Overall, an attempt is made to demonstrate that instead of being a new invention in the history of immunity, stress-induced hypermutation is its primordial form, which resuscitated in vertebrate immune systems helps to direct somatic evolution of adaptive cells towards an adaptive success.

79 Phenotypic integration as both developmental bias and developmental facilitation of evolutionary diversification: empirical examples from teeth, toes, and torsos Kathryn Kavanagh University of Massachusetts Dartmouth, USA

Most theory considering developmental bias on evolution presumes that developmental integration during morphogenesis acts to reduce variations in a population, which then biases evolution since selection can only act on variation that exists. Here I show that it can act as both bias and facilitator of morphological diversification. An empirical example will be presented of developmental integration among segments via an activator-inhibitor based mechanism (the Inhibitory Cascade), which leads to biases in evolution of segment proportions in diverse tissues such as molars, phalanges, and somites. However, we will also present a study which shows a counter example, where developmental integration among adjacent tissues (jaw and tooth) facilitates novel variations and thereby increases evolutionary diversification. Other examples will show that, in some cases, a large number of rib cartilage variants are maintained in a population at an extraordinarily high frequency, while, in other cases, cartilage initiation pattern is extraordinarily stabilized, even when there is apparent selection against the stable form. This two-sided role of developmental integration in evolution, and how it affects evolutionary theory, will be explored.

80 Can plasticity create irreversible constraints? Alfredo Rago, Miguel Brun-Usan, Kostas Kouvaris, Tobias Uller, Richard A Watson University of Southampton, UK

Many traits in nature show no or limited viable variability. This lack of variability suggests the existence of fundamental boundaries to evolution, known as developmental constraints. While constraints can result from non-evolvable physical and chemical properties of life, many biological features that are the result of past evolution are also highly constrained. This can be explained if evolving in one direction creates constraints against moving backwards. For example, evolving a new trait may eliminate the ability to produce viable variants that remove the trait. While empirical observations overwhelmingly support that evolution can be irreversible, few theoretical models try to explain how new constraints evolve, and why they prevent reversible evolution. Here we focus on the hypothesis that developmental interactions cause constraints. To do so, we simulate the evolution of developmental plasticity using a developmental model based on gene regulatory networks (GRNs). We demonstrate that our developmental model can evolve traits that respond to environmental cues (developmental plasticity) according to simple rules, such as proportional increase and non-linear dynamics. Evolution of developmental plasticity changes developmental organization in a way that causes genetic mutations to have similar phenotypic effects as changes to environmental inputs. Conversely, mutations with phenotypic effects that are different from plastic responses become rare (bias). Finally, we show how these biases constrain evolution to proceed in the direction of plasticity even when alternative directions would provide greater fitness, and prevent the selection of less biased mutants. Our work thus highlights how models that integrate general assumptions about development can explain the evolution of constraints in present-day organisms. This reduces the need to invoke ad-hoc constraints and helps us understand when evolutionary changes are expected to be irreversible.

81 Cultural evolution, revolution and transmission of humpback whale song Ellen C Garland University of St Andrews, UK

Culture, the sharing of behaviours or information within a community acquired through some form of social learning from conspecifics, represents a ‘second inheritance system’. Male humpback whales (Megaptera novaeangliae) sing a long, stereotyped, and culturally transmitted song display. At any point in time most males within a population will sing the same version (arrangement and content) of this complex sexual display. However, the song is continually evolving and males must constantly learn and incorporate these changes into their own song to maintain cultural conformity. In addition to evolutionary change, song also undergoes radical ‘revolutions’ where a novel song introduced from a neighbouring population rapidly and completely replaces the existing song. Multiple humpback whale song revolutions have spread across the South Pacific region from the east coast of Australia across to French Polynesia, with a one to two year delay. This has occurred regularly, rapidly and repeatedly across the region; however, we still have a limited understanding of the underlying mechanisms driving this cultural phenomenon and its implications for extra-genetic inheritance. Using empirical data, I will present our current understanding of the mechanisms involved in the song learning process, how these processes may be disrupted, and finally the evolutionary implications for this cultural phenomenon.

82 Niche construction: resolving the process debate Tim Lewens University of Cambridge, UK

The lively debate going on among evolutionists regarding the nature and significance of niche-construction is seamlessly scientific, historical and philosophical. Many scientists also consider the debate’s outcome to have significant implications for biological practice. In this talk I assess three claims about niche-construction. The first (from some niche-construction enthusiasts), is that if we are to explain adaptation, we must supplement natural selection with an additional force that runs from organism to environment. The second (from some niche-construction sceptics) is that while selection (and other basic evolutionary processes) have direct impacts on evolution, the influence of niche-construction is only indirect. The third (from the most prominent enthusiasts) is that niche-construction is a fundamental process of evolution that has not been recognised as such, but which has importance for our understanding of adaptation. I cast doubt on the first two claims. Regarding the third, I draw on Fisher’s case for runaway sexual selection to point to a pragmatic strategy that can secure niche construction as an important evolutionary 'process', regardless of whether it is also judged to be ‘fundamental’.

83 Comprehensive transcriptomics tracks changes in gene expression and coding sequences during C4 evolution Lamiaa A Munshi, Pascal-Antoine Christin The University of Sheffield, UK

C4 photosynthesis is a novel adaptation that increases plant productivity in tropical conditions. It is a complex trait requiring the coordination of many biochemical, anatomical and genetics components. Despite its complexity, the C4 trait evolved more than 60 times independently in flowering plants. However, the history of genetic modifications leading to the C4 trait remain poorly understood. The Molluginaceae family includes closely related species that span C4 photosynthesis as well as a variety of non-C4 types. Capitalizing on this system, we used comparative transcriptomics to track the changes gene expression levels and coding sequences along the phylogenetic tree. Genes encoding core C4 enzymes are upregulated in C4 accessions, as expected. However, we show that these genes were already upregulated in some non-C4 ancestors, which likely facilitated transitions to a C4 biochemistry. By sampling multiple populations per species, we were further able to demonstrate that amino acid replacements adapting the proteins for the C4 function followed the upregulation of genes, and continued when the plants were already C4. Our comparative work therefore shows how a complex biochemical pathway can be gradually assembled through multiple rounds of changes in gene expression and coding sequences.

84 Computational complexity is an ultimate constraint on evolution Artem Kaznatcheev University of Oxford, UK

Experiments show that evolutionary fitness landscapes can have a rich combinatorial structure due to epistasis. For some landscapes, this structure can produce a computational constraint that prevents evolution from finding local fitness optima -- thus overturning the traditional assumption that local fitness peaks can always be reached quickly. I introduce a distinction between easy landscapes of traditional theory where local fitness peaks can be found in a moderate number of steps and hard landscapes where finding local optima requires an infeasible amount of time. Hard examples exist even among landscapes with no reciprocal sign epistasis; on these semi-smooth fitness landscapes, strong selection weak mutation dynamics cannot find the unique peak in polynomial time. A candidate example of such a landscape is yeast’s tRNA gene for recognizing the arginine codon AGG. To fully model myriad proximal constraints on adaptation (methods of generating variation, population structure, stochasticity, etc), I consider all polynomial time algorithms implementing a step of evolutionary dynamics. On hard rugged fitness landscapes that include reciprocal sign epistasis, even these arbitrary evolutionary dynamics cannot find a local peak quickly. This negative result allows reasoning rigorously from disequilibrium to establish the positive results that – even on static landscapes – costly learning can remain adaptive and hitchhiking can maintain cooperation for-effectively-ever. Moreover, on hard landscapes, the fitness advantage of nearby mutants cannot drop off exponentially fast but has to follow a power-law that long term evolution experiments have associated with unbounded growth in fitness. Thus, the constraint of computational complexity enables open-ended evolution on finite landscapes. Knowing this constraint allows us to use the tools of theoretical computer science to characterize the fitness landscapes that we expect to see in nature. Just how ubiquitous hard landscapes are in nature remains on open empirical question.

85 Population-level variation in sperm-mediated paternal effects in threespined sticklebacks Jennifer Hellmann, Alison Bell University of Illinois Urbana-Champaign, USA

In addition to genetic adaptation and developmental plasticity, parental experiences can have lasting effects on offspring phenotypes (parental effects). This has long been appreciated with respect to maternal experiences; however, only recently have we realized that paternal experiences can be transmitted to offspring through multiple mechanisms, including via modifications to sperm. There are many outstanding questions regarding the function and ubiquity of these sperm-mediated paternal effects. Theory predicts that offspring should attend to parental cues when they are reliable and informative of future environmental stressors, but should ignore parental cues when parental environments are not highly correlated with offspring environments. To understand if there is variation in the extent to which parental cues are integrated into offspring phenotypes, we assessed the direction and magnitude of sperm-mediated paternal effects across populations of Alaskan threespined stickleback (Gasterosteus aculeatus). We found no significant effect of paternal exposure to predators in a newly established freshwater population of sticklebacks. However, in a marine and well-established freshwater population, we found that paternal exposure to predation risk impacted both the variance and mean of offspring traits. For both populations, variance in behavioral and morphological traits was greater in offspring of predator-exposed fathers compared to offspring of control fathers. However, the shift in the mean values of offspring traits differed across the two populations; for example, offspring of predator-exposed fathers were less active than offspring of control fathers in a marine population, but in the well-established freshwater population, offspring of predator-exposed fathers were actually more active than offspring of control fathers. These data demonstrate that there is population-level variation in both the direction and magnitude of paternal effects, suggesting that environmental conditions and genetic cues influence the extent to which parental cues alter offspring phenotypes and potentially facilitate adaptation to environmental change.

86 The effect of parental care on population evolvability Sonia Pascoal, Hideyasu Shimadzu, Rebecca M Kilner University of Cambridge, UK

A population’s evolutionary history potentially affects its capacity to evolve because past selection events determine the extent of current genetic variation. Many animals are exposed to selection from a social environment - the conspecifics with whom they routinely interact. However, the effect of past selection by the social environment on current genetic variation is poorly understood. Here we use experiments with the burying beetle Nicrophorus vespilloides to investigate whether selection by parents changes the extent of genetic variation in a population. We measured mutation load by inbreeding individuals from replicate experimental populations that had experienced different levels of care for 25 generations. We tested two predictions: 1) if parental care conceals the negative effects of new deleterious mutations, then inbred individuals with a history of parental care should be more likely to go extinct than those with a history of no care; 2) if parental care conceals the positive effects of new beneficial mutations, then inbred individuals with a history of parental care should be more resilient to inbreeding than those with a history of no care. Our results support the former prediction. We conclude that parental care influences the quality of a population’s standing genetic variation, so changing the raw material available for future evolutionary adaptation.

87 The thrifty watchmaker Michael Lachmann, Wim Hordijk Santa Fe Institute, USA

Why do we need to exercise to keep our muscles in shape? Why not develop the right muscles without such input? Wolff’s law states that bones adapt to the stresses placed on them. A similar law, Davis’s law applies to soft tissue. Plants use light to direct their growth, and blood vessels oxygen concentration. In these cases the underlying developmental mechanism seems to “understand” its high level function. How did evolution reach that fitness peak, when it can only see the local benefits to the lineage? In my talk I will claim that a key to understanding all these and other related phenomena is to switch our view of what a genotype is. Instead of thinking of a genotype as coding for a phenotype, or maybe a G by E matrix coding for a phenotype, which should think of the genotype as an interpreter of environmental information into phenotypic information. The genotype is interpreting the micro-environments that the developmental process encounters to form the phenotype. This view allows us to understand why developmental processes seem so general.

88 Variable phenotypic plasticity and its consequences for evolutionary potential Franziska S Brunner, Alan Reynolds, Ian Wilson, Stephen Price, Steve Paterson, David Atkinson, Stewart J Plaistow University of Liverpool, UK

It is often hypothesized that the pattern of phenotypic plasticity in a population may direct the course of subsequent evolution. As a result, plasticity in populations is often summarized as an average response and labelled as being either adaptive or maladaptive. However, this approach ignores the genetic variation in phenotypic plasticity present within populations and assumes that the mean plastic response is representative of the whole population. Whether this is acceptable or not depends on both the magnitude and the type of plasticity variation maintained in populations; something that is rarely measured. In this study, we quantified the multivariate life history plasticity of 43 clones collected from the same population in response to both temperature and resource level differences. We calculated population-level variation in both the phenotypic direction and magnitude of plastic responses and compared this to two reference populations from very different latitudes. In all cases, we found that within population variation in plastic responses is high and context-dependent. Based on these findings, we discuss the reasons for the maintenance of plasticity variation within a population and its consequences for evolutionary potential. We conclude that there may be many instances where the effect of phenotypic plasticity on the rate and direction of evolution is not accurately represented by an average reaction norm.

89 Comparative genomics of convergent evolution in the emergence of complex traits Claudio L Flores Martinez University of Hamburg, Germany

With the ever-increasing number of sequenced metazoan genomes, it becomes apparent that several complex traits, which had long been held to have evolved in a homologous manner, emerged convergently in distantly related animal lineages. Complex body plans, nervous systems and eusociality certainly are among the most interesting traits that – under the scrutiny of comparative genomics and functional analysis – are breaking away from the hegemony of homology. However, the genomic basis of convergent evolution is not well characterized. Nevertheless, it constitutes a decisive element in understanding the prevalence and adaptive value of convergence throughout metazoan bio-complexity. This first systematic and theoretical study on the genomic underpinnings of convergent evolution focuses on the current rewriting of nervous system evolution in Metazoa. More generally, at least four major mechanisms driving convergent evolution of metazoan genomes can be distinguished: 1) protein family expansion, 2) convergent amino acid substitution, 3) de novo recruitment and 4) transposable elements. Information and (biological) systems theory are reframing the phenomenon of convergent evolution in terms of top-down causation (the process whereby higher levels of organization in structural hierarchies constrain the dynamics of lower levels) and inferred functional equivalence classes. This project explores the convergent emergence of complex traits across multiple scales and at different levels of organizational bio-complexity (from molecular instantiation to cell type, tissue and beyond) within a phylogenetically-inspired context. It aims at establishing a coherent approach allowing, first, to identify problems in evolutionary biology that can be solved by using comparative genomics; second, outline the kind of bioinformatic pipelines needed for solving the problem (e.g. protein family expansion vs. amino acid substitution) and, third, interpret these results within newly forged theoretical and quantitative frameworks that deal with the highly dynamic genomic processes and mechanisms underlying the convergent emergence of complex traits.

90 Epigenetic potential in native and introduced populations of house sparrows (Passer domesticus) Haley E Hanson, Holly J Kilvitis, Aaron W Schrey, J Dylan Maddox, Lynn B Martin University of South Florida, USA

Epigenetic potential, defined as the capacity for epigenetically-mediated phenotypic plasticity, may play an important role during range expansions. During range expansions, populations may encounter relatively novel challenges, including novel pathogens, while experiencing lower genetic diversity due to bottlenecks and/or founder effects. Phenotypic plasticity may allow individuals to rapidly cope with these challenges. Here, we asked whether one form of epigenetic potential (i.e. the abundance and/or position of CpG sites within gene promoters) varied among native and introduced populations of house sparrows Passer domesticus in three microbial surveillance genes: Toll-like Receptors 1B (TLR1B), 2A (TLR2A), and 4 (TLR4). Evidence suggests that increased expression of TLR2 and TLR4 may play a role in house sparrow range expansions, and that methylation at one CpG site within the putative promoter region of TLR4 was a strong predictor of TLR4 expression in house sparrows. We characterized i) total CpG sites, ii) total gains of CpG sites, and iii) losses of CpG sites in TLRs 1B, 2A, and 4. We hypothesized that introduced populations would have more total CpG sites, more gains of CpG sites, and fewer losses of CpG sites (or overall higher epigenetic potential). We found that introduced populations had a higher total number of CpG sites in TLR2A and TLR4, but not in TLR1B. Introduced populations had more losses of CpG sites in TLR1B, more gains of CpG sites in TLR2A, and fewer losses of CpG sites in TLR4 compared to native populations. These trends were not driven by genetic differences between populations, nor by genetic diversity. This suggests that selection is acting on CpG sites in introduced populations, but possibly through different mechanisms, and that epigenetic potential may be playing a role in house sparrow range expansions.

91 Why does natural selection favour the transgenerational inheritance of social adversity? Bram Kuijper, Rufus A Johnstone University of Exeter, UK

Numerous studies have shown that social adversity in early life can have long-lasting negative consequences for social behaviour in adulthood, consequences that are often propagated to future generations. Given these transgenerational effects, it is puzzling why natural selection might favour such sensitivity to an individual's early social environment. To address this question, we model the evolution of social sensitivity in the development of helping behaviours, showing that natural selection indeed favours individuals whose tendency to help others is dependent on early-life social experience. We find that natural selection typically favours positive social feedbacks, in which individuals who received more help in early life are also more likely to help others in adulthood, while individuals who received no early-life help develop low tendencies to helping others later in life. This positive social sensitivity is favoured because of a transgenerational relatedness feedback: patches with many helpers tend to be more productive, leading to higher relatedness within the local group, which in turn favours higher levels of help in the next generation. By contrast, patches with few helpers tend to be less productive, leading to lower relatedness, which in turn favors lower levels of help in the next generation.

92 Trait mean-variance relationships impact speed of adaptation Rebecca B Hoyle, Thomas HG Ezard University of Southampton, UK

Quantitative genetics models of trait evolution typically assume that trait (co)variances remain constant owing to mutation-selection balance. Mean-variance relationships are, however, common in biological systems. Morphological trait data from fossil planktonic forminifera suggest that the trait variance may vary linearly with its mean. We present a quantitative genetics model of trait evolution in which the trait variance changes adiabatically with the mean and show that this leads to faster trait evolution and long- term maladaptation following an environmental shift. This raises interesting questions about the extent to which evolutionary trajectories favour speed of adaptation to changing environmental conditions over fitness outcomes and the limitations of the ubiquitous assumption of constant trait variances through mutation-selection balance.

The influence of multi-level demographic and ecological feedbacks in the evolution of cooperation Antonio MM Rodrigues University of Cambridge, UK

Theoretical models of social evolution typically seek to uncover which genetic and ecological factors mediate the evolution of cooperative behaviours. In addition, most models assume that these behaviours have small phenotypic effects and therefore the interdependency between the behaviour and the environment is considered to be minimal. More generally, behaviours will lead to significant changes in phenotypes, which have the potential to impact both the local and the global environment, which, in turn, will feedback to change the evolutionary trajectories of cooperation. These processes, which take place at different levels of biological organisation, are still poorly understood. Here, I will outline a model that captures the effect of cooperative behaviours at different levels, from the individual to the population, and therefore includes the impact of ecological feedbacks on social phenotypes. I will show how the evolution of cooperative behaviours modifies local demography as well as population density across multiple-generations and I will describe how these multi-level processes can influence the evolutionary trajectories of behaviours. Moreover, I will analyse cases in which individual conditions can be transmitted over multiple generations, and I will show how this can either prevent or promote the evolution of cooperation. 93 The evolution of teaching and anti-teaching James Allen, Rufus A Johnstone University of Cambridge, UK

Alongside genes, culture is also transmitted between generations. Whilst both empirical and theoretical work has established social learning in animals as crucial for the transmission of skill, definitive examples of teaching are rare, and what examples there are exist in very different species. Defining teaching as an enhancement of social learning, we develop a theoretical framework of skilled individuals in a structured population, and investigate two outstanding issues within the evolution of teaching: the importance of demographic competition, and the prevalence of cooperative breeders within empirical examples. By including the cost of demographic competition, we pinpoint the existence of a novel adaptive behaviour: the costly reduction of other’s social learning. In parameter regimes where competition dominates (high vertical transmission and low philopatry) a reduction of social learning is selected for, a behaviour we label anti-teaching. Just as is observed for teaching, we find that anti-teaching is most adaptive when the skill is highly adaptive and difficult to learn. Introducing a trade-off between teaching effort and mortality, anti-teaching is adaptive even when it is costly to reduce the learning of others. Notably, when competitive considerations dominate, individuals are willing to reduce the social learning of others even at a cost to their own mortality. Furthermore, we extend our framework through the inclusion of age-dependent teaching efforts, enabling us to study the age at which teaching is adaptive, and the effect of reproductive skew on the evolution of teaching. By introducing the novel behaviour described above, and age-specific teaching efforts, this work contributes to our understanding of the evolution of teaching, and more generally the spread of culture in animals.

94 Evolution of small RNA production under fluctuating environmental conditions Willian T.A.F. Silva, Sarah P Otto, Simone Immler Uppsala University, Sweden

Recent empirical studies have addressed the importance of small RNAs (sRNA) not only in their primary function of post-transcriptional regulation of gene expression but also in their ability to respond to fluctuations in environmental conditions and be transmitted across individual generations, creating a mechanism of non-genetic inheritance. Small RNA function has been particularly associated with an organismal response to suboptimal environmental conditions caused by biotic (e.g. viral infection) or abiotic factors (e.g temperature fluctuations) and its transgenerational, potentially adaptive effect. We created a model of sRNA dynamics including the major biological components involved in sRNA biosynthesis via the ping-pong amplification process that is characteristic of Piwi-interacting sRNAs. Additionally, we include a transgenerational effect and responsiveness in amplification rates, and explore the sRNA dynamics across individual generations under different fluctuating environmental conditions. Our results show that mechanisms of sRNA amplification become highly invasive when coupled with responsiveness to environmental conditions, and responsiveness in the soma or in the germline can result in different evolutionary outcomes depending on levels of environmental correlation. We consider our findings in the context of current empirical studies and discuss the possible evolutionary consequences of non-genetic inheritance.

95 Inheritance of acquired traits - implications for evolution? Sophie Juliane Veigl University of Vienna, Austria

The inheritance of acquired traits (IAT) has regained prominence in the current heyday of epigenetics. The transmission of acquired information from parents to offspring, even for several generations has been reported in a wide range of species. Proponents of the Extended Evolutionary Synthesis (EES) argue that modes of inheritance and evolution, which are not Neo-Darwinian, are certainly possible. Defenders of the Modern Synthesis (MS) mobilize several arguments to counter such claims. For instance, they argue that IAT has no bearing on evolutionary timescales, as it is too short-lived! Or, they argue that no matter what the specific mechanism of IAT is, all effector molecules are encoded in DNA, and thus subjected to random mutation and natural selection. Is IAT as a mode of evolution thus even plausible? What can be said in its defence? In this paper I discuss two scenarios in which IAT might have an impact on evolution. I use the example of the inheritance of RNAi in the nematode C elegans. Specifically, I present two arguments that address criticisms of defenders of the MS. First, I point to amplification systems in C. elegans, that prolong IAT for up to 80 generations, and counting. Thus, evolutionary timescales of IAT are possible, at least in principle. Second, I consider the active role of small RNAs on restructuring the genome: small RNAs alter the susceptibility of individual stretches of DNA for random mutations, by making them more or less accessible. Thus, they modulate the impact of classical modes of evolution. Based on these two examples I conclude that by looking at the mechanisms underlying IAT, arguments of defenders of the MS can be rejected. Focusing on the specific mechanisms also provides incentives for testing the role of IAT in evolution.

96 Egg development plasticity and the transmission of maternal effects in Daphnia Stewart Plaistow, Aishah Aslam, Megan Hasoon, Marco Marcello University of Liverpool, UK

Environment-induced non-genetic inheritance (NGI) may play a key role in facilitating rapid adaptation to changing environments. However, the mechanisms underpinning NGI remain obscure and we currently know little about how genetic, non-genetic and environmental cues are integrated. The egg phenotype is critical in this respect because it provides many routes for information transfer from one generation to the next and defines the initial environment that inherited genes are exposed to. Parthenogenetic organisms such as Daphnia are ideal models for investigating the integration of developmental cues because large numbers of genetically identical individuals can be reared in different environments in parental and offspring generations, making it easy to separate genetic and non-genetic influences. However, variation in egg phenotypes is typically only ever studied at a simplistic level. Here we use metabolomics to compare the egg phenotypes produced by mothers exposed to different environments, or mothers occupying different states (sexual vs asexual, old vs young). We then use confocal microscopy to compare the embryology of offspring from mothers from different environments, or different states. Our results suggest that (1) changes in the maternal environment or state result in changes in the egg phenotype, and (2) that changes in the egg phenotype results in egg development plasticity that can explain life- history variation in adults. We conclude that offspring provisioning is likely to be a key factor explaining how genetic, non-genetic and environmental cues are integrated.

97 Environmental heterogeneity mediates the evolution of trans-generational effects on fitness Martin I Lind, Martyna K Zwoinska, Johan Andersson, Hanne Carlsson, Therese Krieg, Tuuli Larva, Alexei A Maklakov Uppsala University, Sweden

Theory maintains that environmental heterogeneity can shape the evolution of trans- generational parental effects on offspring fitness but the experimental evidence is limited. Constant or predictably fluctuating environments should favour the evolution of adaptive trans-generational parental effects. In contrast, the absence of correlation between parental and offspring environments should select against trans-generational effects. Here we investigated the role of environmental heterogeneity in the evolution of trans-generational parental effects using experimental evolution in response to temperature. Genetically diverse populations of the outcrossing nematode worm Caenorhabditis remanei, adapted to 20°C, were exposed to a novel, stressful high temperature (25°C) for 30 generations, in constant, increasing or fluctuating temperature regimes, with either strong or zero correlation between parent and offspring temperature environment. We found evolution of trans-generational effects, since fitness in the novel, high temperature depended upon parental environment. While lines evolving in correlated environments required parental exposure to high temperature to achieve high reproduction in this environment, the total reproduction of lines evolving in unpredictable environment was independent of parental environment. This was caused by the adaptive maintenance of an anticipatory trans-generational parental effect in lines evolving in correlated environments, while this trans-generational inheritance had been adaptively lost in lines evolving under non-correlated environmental heterogeneity. Moreover, the individual fitness (lambda) of the lines evolving in uncorrelated environments was actually highest when parents and offspring where grown in opposite environments (simulating environmental fluctuations). Therefore, environmental heterogeneity influences the evolution of trans-generational inheritance.

98 Abstracts: Poster Presentations

P01 Expanding the conceptual scope of niche construction theory Bendik Hellem Aaby, Grant Ramsey KU Leuven, Belgium

Niche construction, commonly defined as organisms actively changing selection pressures by altering the feature-factor relationship between themselves and their environment, has become increasingly incorporated into evolutionary dynamics in contemporary evolutionary biology. The interest in niche construction was largely sparked by Lewontin’s work on biological constructivism, and more recently builds on niche construction theory (NCT) Odling-Smee et al. (2003). NCT argues that we can distinguish two fundamental kinds of niche construction—perturbative and relocational. These concepts refer to the way in which niche construction is brought about by the activities of the organisms. ‘Perturbative’ captures the cases in which an organism’s activities alter its physical (external) environment and thereby changes the selection pressures, e.g., ecological engineering. ‘Relocational’ captures the cases where the organisms relocate in space (without modifying the environment), and thereby changes the selection pressures, e.g., migration. The NCT conception of niche construction is considerably narrower than that of Lewontin. In this talk we defend a broader framework more in line with Lewontin’s original account. We hold that there are three fundamental ways in which organisms can construct their niche: ‘constitutive’, ‘relational’, and ‘external’ niche construction. We aim to show that NCT’s ‘perturbative’ and ‘relocational’ excludes important niche construction cases, and we suggest that our framework is able to subsume all the phenomena described by NCT while also allowing us to account for far more cases of niche construction, especially social niche construction. We end our talk by showing how this broader Lewontian framework can help us elucidate how niche construction can be treated as an evolutionary process as understood by the extended synthesis.

99 P02 Could a major evolutionary transition have happened in human evolution? Paulo C Abrantes University of Brasília, Brazil

Paulo C. Abrantes attempted to make explicit, in a 2013 paper, the conditions under which a major evolutionary transition could have taken place in human evolution by using Godfrey-Smith's criteria for telling marginal from paradigmatic Darwinian populations in the cultural domain. In that paper, I argue that the emergence of cooperation in hunter-gatherer bands during the Pleistocene could have been a major transition in individuality, associated with the evolution of a species-specific social psychology. Difficulties arise, however, when one attempts, by adopting this approach, to come up with a notion of fitness pertaining to cultural groups, among other problems in applying to those groups the requirements for biological individuality. I will examine in this talk the plausibility of another scenario, in which a major transition in individuality could have happened later, associated with ultrasociality in large-scale societies. Godfrey- Smith's conceptual scheme applies less problematically in this case. This scenario doesn't presuppose any further psychological capacities on top of those which emerged among hunther-gatherers, arguably by gene-culture coevolution. A major transition could have been the outcome of just cumulative cultural evolution. Furthermore, this scenario has the advantage to be compatible with other attempts to reconstruct the latest stages of the evolution in the Homo genus, including those proposed by Maynard Smith and Szathmáry, Turchin, Richerson, Boyd, and Tomasello, among others. However, this alternative scenario has to deal with criticism from those that reject adaptationism and push for an expanded version of the theory of evolution in able to integrate developmental and evolutionary processes. A more heterodox proposal along those lines would attempt to explain human ultrasociality by underlining the role of ontogenetic processes and niche construction, and playing down natural selection in accounting for that trait.

100 P03 Dialectics of evolution; epistemology versus practice Ozan A Altinok WWU Münster, Germany

My main aim is to distinguish the usages of evolution such as evolutionary models, evolutionary theory, evolutionary phenomena and “evolution proper” to understand the extent of change in the call for change in the Modern Synthesis (MS) in favor for the Extended Evolutionary Synthesis (EES). Gould (1981) distinguishes evolution as fact and theory. I would like to add evolution as instrument to have a broader perspective to understand change in evolution. Through elaborating that scientists often make the distinction between the theories which are “home truths” (Hacking, 1984) and theories which are under investigation, I will analyze the limits and the location of change in “evolution” that is happening and that is expected to happen by EES. Wray et. al, (2014) call for an understanding of evolution which permits the domains to be on its core, as long as they are empirically validated, while on the very same commentary Laland et. al (2014) call for an EES for biologists to grasp a better understanding of evolution. I would like to consider scientific practice more central rather than theories. Going away from “high theories”, Cartwright (1983), helps us to see the extent of flexibility in the usage of the concept of evolution, holding that evolutionary theory, as a theory, has limited relevance to the practices of biologists. I claim that scientists are generally very pragmatic and instrumentalist (Matthews, 2017), when considering their work in most cases, as guidelines of research. I claim that, Callebaut’s (2010) understanding of dialectics of unity and disunity should be extended to dialectics of practice of biology and biological theory with the scientists in the center to grasp the possibility of change in accurate aspects of evolution through EES and other possible scenarios that are in the form of a movement, not only a protest.

101 P04 Overprinting and the evolvability of novel protein-coding genes Zachary Ardern, Stefan Wichmann, Siegfried Scherer Technical University of Munich, Germany

As techniques to discover short proteins have improved, many overlapping genes encoding protein products have been discovered in viral, prokaryotic, and eukaryotic genomes. Methods such as ribosome profiling, large-scale analyses of sequence constraint made possible by modern databases, and improvements in proteomics have all contributed to this substantial increase in our ability to detect protein products. Studying the origin of these genes provides an important case study in evolutionary innovation. This class of genes is particularly useful for obtaining insights on gene origin as the genomic context prior to an overprinting event can be inferred with more confidence than is possible in the case of most intergenic sequences. Important factors in facilitating the origin through overprinting and subsequent conservation of novel protein coding genes include contributions of the pre-existing ‘mother’ gene’s sequence, the reading frame of the overlapping ORF, and the structure of the standard genetic code. Through a combination of phylostratigraphy, ancestral sequence reconstruction, and modelling of overlapping gene construction and alternative codes, we report on the contributions of each of these factors, placing them within a broader context of genomes’ inherent potential for evolutionary innovation.

102 P05 Morphological divergence in Drosophila ananassae due to nutritional variations through changes in allometric relationship and trait size Bhumika, Arvind Kumar Singh Banaras Hindu University, India

Developmental plasticity in response to nutritional changes plays a substantial role in adaptive evolution and diversification of morphological traits in organisms. We investigated the diversification in wing dimensions and its relative scaling with respect to body size, i.e., allometric scaling in response to dietary variations in carbohydrates and proteins quantity in Drosophila ananassae. For the experiments, ten isofemale lines of D. ananassae derived from Bilaspur, India and karyotypically characterized stocks of D. ananassae from Varanasi, India were utilized. We found that protein and carbohydrate variations in dietary media induced significant genetic variation in plasticity for allometric slopes of wing traits in males and females of Bilaspur population. However, Varanasi population refrained from showing such effect in response to dietary variations. The results suggest that changes in allometric coefficients due to dietary variation is population specific and this may lead to difference in morphological divergence in populations, thereby, causing evolutionary allometry. The analysis of reaction norm graphs suggested the involvement of alpha inversion present in 2L chromosome of Varanasi population, in conferring adaptation to different nutritional environmental conditions due to changes in static allometry of different wing dimensions. We detected that effect of dietary protein variations on plasticity of absolute body size and wing dimensions was highly genotype dependent when compared to variation in sucrose quantity, in both the sexes and in both the populations. The results signify that variations in protein in natural environment have greater tendency to induce selective pressure on absolute trait values of genetic variants as compared to carbohydrates and thereby had greater capability of inducing morphological diversifications through the process of genetic accommodation. Overall, this study reveals that the nutritional ecology governs the morphological divergence of traits size and shape in a genotype dependent manner in D. ananassae thereby resulting in morphological evolutions.

103 P06 Coral-built structural complexity enhances coral recruitment: niche construction in action? Viviana Brambilla, Andrew H Baird, Miguel Barbosa, Inga Dehnert, Joshua S Madin, Clare Peddie, Maria A Dornelas University of St Andrews, UK

Coral larval recruitment is a key process for coral reef persistence. Micro-habitat scale rugosity enhances settlement success; however, the effect of structural complexity of the habitat forming coral colonies remains undetermined. Here, we test the effect of habitat scale structural complexity on settlement and juvenile presence on artificial tiles. Structural complexity had a positive effect on settlement (+ 15.71 % of settler presence probability along the gradient), but no effect on juvenile abundance was detected. While complexity might be trivial for early stages survivorship, our results suggest that structural complexity as result of coral morphology creates conditions that facilitate coral settlement and serve as foundation for further studies of coral niche construction.

104 P07 When are more experiments necessary? Using evolutionary considerations in investigating animal minds Simon Brown Columbia University, USA

In the study of animal minds, available neuroscientific and behavioural data often underdetermine which of several radically different models of a particular species' mind is best. Morgan's Canon traditionally offered one way of responding to this challenge, but it has well-known problems. One alternative, considered by Sober (2005, 2015), is cladistic parsimony. But as Sober points out, this principle too will often only very weakly constrain our attribution of mental traits to animals. One possibility here (Sober 2005, Fitzpatrick 2008) is that we should abandon the search for principles of this kind and treat these issues entirely on a case by case basis. This paper, however, shows that we can usefully introduce more general principles for combining evidence about pairs of species, taking account their phylogenetic relations on the one hand and similarities and differences in their niches on the other. We can thereby systematically determine how confident we should be in attributing any given mental state to a given species. The key to doing this will be to understand a general notion of evolutionary dependence - roughly, considerations about the order in which various traits must appear in evolution. The generality and utility of this sort of notion has been underappreciated in the evolutionary sciences of animal minds. With a computational account of a cognitive trait, we can understand what it might evolutionarily depend on. This notion can include the numerous cases where a trait of interest might be an exaptation based on a computationally similar mechanism used for a rather different function. And this in turn means that we can use evidence from one species to generate predictions about related species with much more robustness than might be expected, and without the need for extensive, expensive and invasive experiments for each species of interest.

105 P08 The evolution of unicellularity in a multicellular world Charlie Cornwallis, Maria Svensson-Coelho, Karin Rengerfors, Lars-Anders Hansson Lund University, Sweden

Explaining how life evolved from simple replicators in a primordial soup to the array of biodiversity observed today remains extremely challenging for evolutionary biology. Increases in the complexity of life have occurred through a series of transitions where solitary replicators, such as genes, cells and animals (simple life forms), committed to living in groups, such as genomes, multicellular bodies and societies (complex life forms). However, not all organisms are complex so what explains when transitions occur? This question has been addressed by comparing complex and simple organisms. This allows inferences about the changes that take place during transitions, but reveals little about the adaptations in simple organisms that facilitate and hinder the evolution of complexity. Uncovering such ‘precursors’ to complex life is extremely challenging, as it requires identifying the mechanisms underlying complexity and tracing their origins back over time. Here I will present on-going work that integrates experimental evolution and comparative work to understand whether there are ‘precursors’ to both multicellular and unicellular life using the model system of green algae collected from non-model wild populations.

106 P09 Enactive evolution Amanda Corris University of Cincinnati, USA

According to the enactive approach to biological organization, an organism can be said to enact or ‘bring forth a world’ through structural coupling, where constructive interactions between organisms and their environments result in a relationship of co- evolution between those organisms and their environments as niches. While the enactive approach mainly aims to provide a theory of cognition, it also provides a theoretical framework for an alternative to received conceptualizations of adaptation. I argue that this framework is complementary to niche construction theory and can bolster a niche construction theory of evolution by supplying conceptual tools that help to explain the processes involved in the generation of selection. The enactive concept of evolution by natural drift (Varela, Thompson and Rosch 1991), whereby self- organizing, structurally coupled configurations of organisms and their niches generate selection, provides an account of feedback in evolution that can build on similar accounts offered by niche construction theory. The enactive approach has significant contributions to make to theories that aim to reconceptualize evolutionary processes and is worthy of consideration in contemporary debates regarding development and evolution.

107 P10 Behemoth: Henri Bergson's and Gilles Deleuze's vitalism, a proto evolutionary biology Jack Robert Edmunds-Coopey University of Durham, UK

Bergson's philosophy of life and its organicism inspired Deleuze's vitalistic tendencies in the 20th century which attempt to not only capture life as it is but to use scientific explanations of life integrated with philosophical ones. The study of life itself is what fascinated these thinkers which influenced not only their understanding of time in how the life experience of time provides more insight than a metaphysical account deriving from Kant. In light in how science despite the instrumental critiques of reason raised against it by the Frankfurt School in how 'science does not think' in Heidegger's terms continues to innovate and destroy life as we know it as and not as an object of science. Thus, it is proposed that the science of life as such is argued to study inanimate things but if we question like Bergson and Deleuze did this assumption of the binary between animate and inanimate objects through vitalistic tendencies, arguing that in fact life is not limited to biological entities but all forms of objects around us in a new understanding of what constitutes life as such outside a Darwinian and Newtonian definition which is perhaps restricted to the plant and animal kingdoms. Thus, life as we know it becomes dissolved if we re-consider the assumptions which delimit animate things from inanimate things and consider objects as examples of manifestations of life and how our desires, values, morality and meaning give 'life' to all sorts of objects and things around us consequently. In conclusion, this paper shall explore the vitalism of Henri Bergson and Gilles Deleuze and argue that their interactions with reconceiving of life not only revolutionizes understandings of biology, religion and the human meanings associated with societies, but that the boundaries and barriers between the humanities and sciences should not be existent.

108 P11 Effects of phenotypic robustness on adaption and evolvability Emanuele Rigato, Giuseppe Fusco University of Padova, Italy

Theoretical and experimental studies have provided evidence for a positive role of phenotype resistance to genetic mutation, or “phenotype mutational robustness”, in enhancing adaptation and fostering evolvability. With the aim of contributing to an understanding of the origin and evolution of phenotypic robustness in living systems, we adopted a theoretical approach, elaborating on a standard model of evolutionary dynamics, complemented by computer simulations. Results show that, under common selective regimes, a certain level of phenotypic robustness is a necessary condition for adaptation to occur, even in the absence of any drift effects. We also tested the predictions of the model by exploring some selected features of genome evolution, through a phylogenetic comparative approach on a sample of 210 eukaryote taxa. Phenotypic robustness qualifies as major quantitative determinant of biological system’s evolvability, a key feature of the genotype-phenotype map which would deserve to be formally included into a more inclusive explanatory framework of the evolutionary theory.

109 P12 Spadefoot toad tadpoles incur maintenance costs of developmental plasticity in the form of increased oxidative stress Pablo Burraco, Miguel A Rendón, Carmen Díaz-Paniagua, Ivan Gomez-Mestre Doñana Biological Station, CSIC, Spain

Adaptive plasticity enables organisms to improve the match between their phenotype and the environment under fluctuating environmental conditions. However, despite this advantage of plastic genotypes, we often observe substantial variation in the degree of plasticity across genotypes within populations. Costs of maintaining the sensory and response machinery of plastic traits have been proposed as a plausible explanation for the maintenance of genetic variation for plasticity, although only a few studies have detected them empirically. The majority of amphibians have an aquatic larval phase that typically shows a high degree of developmental plasticity, as they can alter their growth rate, morphology and behavior against various environmental inputs. We have tested for physiological maintenance costs of developmental plasticity in larval spadefoot toads in response to predation risk. We determined the plastic responses of 20 sibships to water-borne predator cues, and estimated parameters associated with metabolic rate, oxidative stress and immune status on tadpoles under control conditions. We found that plasticity in response to predator presence entailed maintenance costs in terms of antioxidant and immune imbalance. Our results indicate that being plastic can be physiologically costly, suggesting that the role of maintenance costs in the evolution of plasticity should not be dismissed.

110 P13 The evolution of barriers to exploitative behaviors Jonathan R Goodman, Paul Ewald University of Cambridge, UK

This presentation aims to show how barrier theory, which was generated to understand how cancer develops through the abrogation of mechanisms that block cellular reproduction and survival, can be more broadly applied to other systems, including animal populations and human societies. Barrier theory suggests how Red Queen–like evolutionary relationships may be ended, or at least paused. Barriers may be understood as naturally evolved or artificially implemented mechanisms for blocking the risk of exploitation. Selection will therefore favor those entities capable of abrogating these barriers: oncogenic viruses, for example, evolved enhanced survival by blocking the suicide mechanisms of their host cells. We propose a model of barrier theory suggesting that, for any system, barriers to exploitation and the degree of exploitation are inversely and logarithmically related. The mode of exploitation may, furthermore, be unpredicted, which cannot be accounted for in a standard signaling game model. Modes of exploitation differ depending on the system: invasive organisms may overcome the evolved barrier to cellular survival and reproduction; a plant may temporarily escape herbivory by evolving a chemical that none of its herbivores can detoxify. Together these premises highlight some important consequences for competitive co-evolutionary relationships.

111 P14 Three lessons of Bergson's creative evolution in the wake of transhuman evolution Daina Habdankaitė Vilnius University, Lithuania, and Turin University, Italy

The fast-paced advancement of AI, robotics, gene editing, and other contemporary technologies is considered by several theorists as the wake of transhuman evolution. The idea behind it is that human evolution is about to transcend the biological boundaries of the human organism in order to complete itself in a technically mediated form. The idea of transhuman evolution brings into question the process of evolution as well as the notions of species, intelligence, continuity and predictability of the process of development. By exploring a few examples of AI, robotics, and gene manipulation in creative practices, I will suggest that Bergson‘s concept of creative evolution could provide us several valuable lessons in conceptualizing and evaluating the idea of transhuman evolution. 1) Bergson’s strict division between intellect and intuition, grounded on their relation to the matter, helps to shine a light on the importance of materiality in the case of technological advancement of intelligence. To put in technological terms, once admitted that hardware decides what software is to be developed, is it possible for the software to influence hardware as well? And if so, what implications does this have on the conception of evolving intelligence and its transformations? 2) The line that Bergson draws between creativity and calculation helps to crystallize the problematic aspect of AI development: in order to be considered as AI, it must bear some resemblance to the human mind; but is creativity something that cannot be reproduced in and by a machine? What is it like for a machine to be creative? 3) Bergson’s notion of virtuality, when applied in the context of transhuman evolution, shines some light on the problem of uncertainty that part of AI and robotic technologies are aimed at solving. Is uncertainty a human fault or an evolutionary system’s default?

112 P15 Evolution of phenotypic complexity: a developmental perspective Pascal F Hagolani, Roland Zimm, Isaac Salazar Ciudad University of Helsinki, Finland

How does phenotypic complexity evolve? Indirect empirical evidence suggests that mutations which decrease complexity are more common than mutations that increase complexity. Additionally, it has been shown that complex phenotypes have a more complex relationship between genetic and phenotypic variation, or genotype- phenotype map, than simpler ones. The view from classical evolutionary biology is that complex genotype-phenotype maps hinder adaptation. Thus, these complex genotype- phenotype maps should be rare or, at least, become simpler over the course of evolution. In this paper, we show that, instead, such complex maps and the above mutational asymmetry are an inevitable consequence of how genes need to be wired into networks during development in order to lead to complex morphology. To explore this question, we randomly wired a huge number of genes and cell behaviours into networks in EmbryoMaker. EmbryoMaker is a mathematical model simulating collective changes in cell 3D spatial positions and gene expression due to such networks. In this way we obtained a “zoo” or an ensemble of in silico 3D embryos. Real gene networks are not random, but random networks allow an unbiased view on the requirements for the development of complex phenotypes. We found that the networks leading to complex phenotypes are: 1) rare; 2) need to be finely tuned to produce complex morphologies; 3) less stable to noise than networks leading to simple phenotypes; 4) more likely to lead to a decrease in complexity than to an increase when they are mutated and; 5) they exhibit a complex genotype-phenotype map. Our results suggest that, when complexity increases during evolution, it does so at a progressively slower rate as mutations which increase complexity become rarer, morphologies become less robust, and a more complex genotype-phenotype map arises.

113 P16 Timing of maternal exposure to toxic cyanobacteria and offspring fitness in Daphnia magna: implications for the evolution of anticipatory maternal effects Reinder Radersma, Alexander Hegg, Daniel WA Noble, Tobias Uller Lund University, Sweden

Organisms that regularly encounter stressful environments are expected to use cues to develop an appropriate phenotype. Water fleas (Daphnia spp.) are exposed to toxic cyanobacteria during seasonal algal blooms, which reduce growth and reproductive investment. Because generation time is typically shorter than the exposure to cyanobacteria, maternal effects provide information about the local conditions subsequent generations will experience. Here, we evaluate if maternal effects in response to microcystin, a toxin produced by cyanobacteria, represent an inheritance system evolved to transmit information in Daphnia magna. We exposed mothers as juveniles and/or as adults, and tested the offspring’s fitness in toxic and non-toxic environments. Maternal exposure until reproduction reduced offspring fitness, both in the presence and in the absence of toxic cyanobacteria. However, this effect was accompanied by a small positive fitness effect, relative to offspring from unexposed mothers, in the presence of toxic cyanobacteria. This effect was mainly elicited in response to maternal exposure to toxic cyanobacteria early in life and less so during reproduction. None of these effects were explained by changes in egg size. A meta‐ analysis using our and others’ experiments suggests that the adaptive value of maternal effects to cyanobacteria exposure is weak at best. We suggest that the beneficial maternal effect in our study is an example of phenotypic accommodation spanning generations, rather than a mechanism evolved to transmit information about cyanobacteria presence between generations.

114 P17 Studying the evolutionary process with genetic algorithms Edith Invernizzi University of St Andrews, UK

Genetic Algorithms (GAs) are a stochastic search algorithm that exploits the mechanisms of biological evolution -selection, reproduction, mutation and crossover- to reach optimal solutions in a large, complex fitness landscape where best-performing solutions cannot be found analytically. Despite having originally been devised as a tool for the study of the evolutionary process itself, they have had limited applications in evolutionary biology, mainly in the optimisation of analytical tools (e.g. in phylogenetics), or in the simulation of complex trait evolution (e.g. cognitive processes) – to the point that they are mainly used in engineering. Here, I use GAs to study how differences in the process of selection and reproduction may lead to different end points in the fitness landscape. I also analyse the dependence of this trajectory on other elements that are known to have an important role in the biological evolutionary process: namely, population size and mutation rate. I use the Sir Philip Sidney’s Game, a game theory problem applied to the study of the evolution of signalling, as a test bed for our query. I compare five ways of modelling selection and reproduction in the algorithm and I look at the relative frequencies of Evolutionary Stable Strategies, Evolutionary stable Sets and other, evolutionary unstable, solutions emerging under each variant, to analyse the reliance of each method on landscape exploration versus exploitation of the locally optimal solution. Because differences in the outcome results from how strictly reproduction depends on quality and on relative reproductive fitness of the higher to lower-quality solutions, these results have interesting implications for the effect of different population structures in biological evolution. This study highlights the usefulness of this simulation tool in improving our understanding of the evolutionary process itself.

115 P18 The evolutionary origin of our moral beliefs Man-Him Ip University of Birmingham, UK

In recent years, philosophers have discussed evolutionary debunking arguments of morality (EDA) extensively. These arguments basically start with an empirical claim that evolutionary forces have had a significant influence on what moral beliefs we have. Evolutionary debunkers then aim to show that the epistemic status of our moral beliefs is diminished due to that influence. However, we first need to know at least how evolution has influenced and shaped our moral beliefs, if it really has done so. I attempt to draw a very rough outline of the nature of moral beliefs. Basically, there are at least three essential features or key characteristics of moral beliefs: (1) There are some core contents of moral beliefs that are commonly shared in most communities (2) Moral beliefs are reliably connected to our motivation (3) Moral beliefs are connected to praise and blame I argue that a plausible view of the evolutionary origins of our moral beliefs must be able to explain how our moral beliefs came to have those three essential features. In this paper, I am going to introduce and evaluate two different theories of how evolution could have influenced and shaped our moral beliefs. I call these two views the adaptation account and the exaptation account. I will explain how these two accounts can provide a plausible explanation of how evolution could have influenced our moral beliefs. In other words, these accounts can explain why having moral beliefs with the three essential features is an adaptation and an exaptation respectively. However, I remain neutral on which of them could be the best account of the evolutionary influence on our moral beliefs. What I will demonstrate is that these two accounts have their own plausibility.

116 P19 Genetic assimilation in the fossil record: phenotypic plasticity and accommodation in Cambrian arthropods Illiam Jackson, Madeleine Bohlin, Richard Mann, Graham Budd Lund University, Sweden

Genetic assimilation describes a process in which an initially plastic response of an organism to the environment becomes constitutively expressed through quantitative genetic change. Here we suggest that the Cambrian Series 3 trilobite-like arthropod Agnostus pisiformis and subsequent members of its evolutionary lineage allow us to probe the fossil record for signals of genetic assimilation. The lineage is geographically broadly represented, as well as stratigraphically well-defined, stretching through a period of increasing anoxic/euxinic conditions interpreted as stressful to the organisms. Using elliptical Fourier analysis (EFA), we quantify the morphology of specimens recovered from geographically and stratigraphically varied assemblages, including several coeval but geographically disparate localities. For each of these assemblages we determine relative levels of anoxic/euxinic stress using redox-sensitive metals (Mo, U and V) as proxies, analysed via mass spectrometry of the sediment. We uncover an evolutionary pattern in which A. pisiformis and the subsequent member of the lineage, H. obesus, exhibit directional morphological change as well as increasing morphological variation in response to increasing anoxic/euxinic stress. The most recent member of the lineage, T. holmi, responds to the relaxation of this stress by reducing its variability around a new morphological mean within the range of the lineage’s earlier expansion. We interpret this pattern as (1) environmental stress inducing novel phenotypic variation, (2) selection acting on the induced phenotypic variation followed by (3) canalization of the initially plastic response; in other words, genetic assimilation.

117 P20 Extended cognitive systems, meaning, and evolutionary transitions Hilton F Japyassú Universidade Federal da Bahia, Brazil

Considering that non-biological entities are capable of information processing, and that cognition is by and large considered as adaptive information processing, the extension of biological cognition to worldly components is slowly turning into a sound scientific enquiry. This is relevant for an extended evolutionary synthesis, since worldly information processing that is tightly coupled to organism functioning can evolve as to be encapsulated by the organism itself: the organism extends towards specific components of the world, thus creating larger evolutionary units. Also, since extended cognition is tightly coupled to developmental learning processes, the new unit is not random, but rather a fully adjusted new phenotype, thus increasing evolvability. Extended cognition is thus a promising research program, but one should avoid important pitfalls that could halt its development. If information is taken as the only relevant aspect of cognition, with no room for meaning or representations, extended cognition risks excluding large fields of cognitive enquiry. We build on recent theoretical developments on biological autonomy to track the emergence of meaning within dynamical, information processing systems. Meaning requires the generation of subsystems that are both autonomous and integrative. A cognitive system emerge with new organisation levels, integrating informationally the performance of previously disparate subsystems; it requires regular responses to similar external triggers, contextual adjustment, and internal consistency. We depict successful instances of extended meaning systems, demonstrating that meaning expansion emerges from self- organisation within unpredictable environments, with the creation of hybrid brain- environment loops. It turns out that the extension of cognition to worldly components works by selecting out the expansion of meaningless information processing, paving the way to major evolutionary transitions.

118 P21 Mechanisms of hormone mediated parent-offspring conflict Neeraj Kumar1.3, Martijn van Faassen2, Bonnie de Vries1, Anja Lohrentz3, Ido Kema2, Manfred Gahr3, Ton GG Groothuis1 1Behavioural Biology, Groningen Institute for Evolutionary Life Sciences, University of Groningen, The Netherlands; 2Laboratory Medicine, University Medical Center Groningen, University of Groningen, The Netherlands; 3Behavioural Neurobiology, Max Planck Institute for Ornithology, Seewiesen, Germany

Vertebrate embryos are exposed to a varying degree of maternal hormones, having profound consequences for their later phenotype. Bird species have been used extensively to study these effects, with a strong focus on ecological and evolutionary factors affecting maternal hormone allocation to the egg, showing systematic variations among different eggs of the same nest, between-nests, and between-females. However, the embryo itself is often seen as a passive responder. Its actual role in adapting to and translating maternal hormonal signals, and its role in the evolution of parent-offspring conflict, has largely been ignored. By focussing on early hormone dynamics in the egg and embryo we found that (1) The active maternal hormones in the egg, such as progesterone and testosterone, are substantially and rapidly metabolized by the embryo, with a corresponding increase in much less potent metabolites; this suggests that the embryo itself can time its exposure to the maternal signal by converting less potent hormones back to potent ones when needed; (2) The dynamics of androgen metabolism differs systematically between eggs of different laying order; this opens the possibility that the embryo can deal with the maternal signal depending on the context it experiences, which may depend on yolk composition and incubation pattern; (3) The embryo expresses steroid receptors in its extra-embryonic membranes even before its own hormone production starts, indicating that selection has favoured the early perception of the maternal signal; and (4) The embryo downregulates these receptors in response to increased egg steroid levels, indicating that the embryo can regulate the effectiveness of the maternal signal. Altogether, the results imply that the embryo is not simply a ‘slave’ to the maternal signals but can potentially play its own role in parent-offspring conflict, opening new avenues to integrate mechanistic, developmental and evolutionary approaches to the study of maternal effects.

119 P22 The impact of individual immune experience on niche construction in red flour beetles, Tribolium castaneum. Lai Ka Lo, Caroline Müller, Martin Kaltenpoth, Joachim Kurtz University of Münster, Germany

Adult red flour beetles, Tribolium castaneum produce external secretions containing high levels of antimicrobial quinones (benzoquinone and hydroquinone), which have been shown to inhibit the growth of bacteria, yeast and fungi commonly found in flour and the beetle’s natural environments. As group living animals that share the same niches, potentially, the secretions from adult beetles could in turn influence the levels of pathogenic threat that their conspecifics or offspring are exposed to and thus modify the selective environment of both the niche constructors and recipients. In response to the sparse experimental tests of the ecology and evolutionary consequences of niche construction, we here start using the worldwide pest of stored grains, T. castaneum along with their natural parasite Bacillus thuringiensis tenebrionis (Btt) as a host- parasite model system for niche construction. Interestingly, T. castaneum demonstrated immune priming (i.e. increased immune response against previously encountered microbes) within and across generations. As the magnitude of animals’ niche construction likely varies with individual experience, i.e. exposure to environmental stressors, we hypothesise that individual immunological experiences in T. castaneum might lead to the construction of individualised niches. In our studies, individual beetles were either left naive, sham-exposed by injection of PBS or primed via injection of heat-killed vegetative Btt. Using spectrophotometry and bacterial 16S RT-qPCR, we compare both the quinone levels and microbial loads in the bodies and flour of adult beetles undergoing different treatments. We will present our preliminary work on the influence of individual immune experience on niche construction and our plan to test the evolutionary consequences of individual niche construction via experimental evolution.

120 P23 Trait-based theory leads to generality across successional systems Jessica Haghkerdar, Brian McGill, Maria Dornelas University of St Andrews, UK

Succession is the turnover in species and/or traits caused by a sudden availability of unexploited resources. Succession is one of the oldest fields in ecology, but we still do not have cross-system generality in our models of how succession progresses. If we want to achieve generality between systems as disparate as gut microbiomes, whale falls, and tropical forests, we need to be able to compare them empirically and with a broad taxonomic scope. Developing a general understanding of succession will entail standardising between different successional trajectories. The few cross-system succession studies that do exist are usually in a narrow range of habitat and disturbance types, but taking a trait- based approach will allow us to compare responses in any variable between successional systems. As a starting point for trait-based synthesis, we present a range of hypotheses on how key traits and life-history attributes will change during succession. These traits span all taxonomic groups and are relevant to any ecosystem type. While this framework should apply to any level of organisation, we particularly encourage its use in multitrophic communities, which have been under-studied, especially in succession. Traits used in succession studies should reflect reproduction (number and timing of offspring), performance (how an organism fares in one patch in space and time), and heterogeneity (how an organism copes with changing conditions). Building on traditional r-K life history divisions, this framework specifically highlights coping with changing conditions over time, as in succession. This deliberately cross-system, cross- taxon, multitrophic framework is suitable for both system-specific studies and multisystem comparisons. Its future application will help both applied and conceptual ecologists predict and manage systems undergoing succession and restoration.

121 P24 From anisogamy to intrasexual competition Ivain Martinossi, Mattias Siljestam Uppsala University, Sweden

The biological evolution of separate sexes in animals, females that produce eggs and males that produce sperm, is a fundamental question of evolutionary biology; furthermore, this initial difference between the sexes is thought to have resulted in a cascade of evolutionary steps that influenced many aspects of sexual dimorphism, including the intensity of competition for mating opportunities. Although it has been advocated that males compete more often and more intensely than females in animals, and that this pattern finds its roots in the state of anisogamy (gamete size dimorphism), we suggest that the evolution of anisogamy may not always spur the evolution of male- bias competition for mating. Using the method of adaptive dynamics, we develop a model of the coevolution of gamete size and competition for mating, starting in a population without sexual dimorphism. We show, in accordance with previous work, that the evolution of anisogamy is highly sensitive to population density and survival constraints. We also find that anisogamy is often accompanied by the evolution of a sex-bias in competition for mating. Finally and most importantly, competition for mating may evolve to be either female or male-biased, depending on the competition trait considered and which were the conditions that triggered the evolution of anisogamy in the first place.

122 P25 Diet-based developmental plasticity in a freshwater isopod Moritz Lürig, Kim Kaltenbach, Blake Matthews Eawag, Switzerland

Developmental plasticity is common in natural populations and can play an important role in adaptive evolution. When developmental plasticity is heritable and exhibits genetic variation in a population, it can evolve when selective environments change. Evidence for the evolution of plasticity can be obtained by (i) measuring the heritability of developmentally plastic traits, (ii) identifying the causes of plasticity in natural environments, and (iii) testing how putative agents of selection affect the distribution of developmental plasticity over time (within or across generations). The freshwater isopod, Asellus aquaticus, is a useful animal model to study the evolution of developmental plasticity in natural populations. Cryptic pigmentation of A. aquaticus is thought to evolve rapidly in response to predator-mediated selection. However, pigmentation is a developmentally plastic trait - individuals are born with low pigmentation, and irreversibly accumulate pigmentation over their lifetime. In a common garden experiment with 29 families (split-clutch, full-sib design), we show that the rate of pigmentation accumulation during development varies among families and increases with a high protein diet. Further, we found that fast growing individuals on low protein diets have a higher survival probability when they also have higher rates of pigmentation accumulation. To test whether pigmentation improves crypsis we performed predation trials with a visually foraging predator (i.e. stickleback fish), in which we manipulated both the matching background (i.e. dark and light sand) and frequency of background-matched individuals (i.e. low and high frequency of dark and light phenotypes). In these experiments, we find no evidence that fish impose either divergent or frequency-dependent selection on isopod pigmentation. Overall, our results suggest that natural isopod populations harbour genetic variation in diet-based developmental plasticity of pigmentation, but the adaptive value of this plasticity is still unknown.

123 P26 Global human niche construction: an evolutionary trap? Andra Meneganzin, Francesco Suman, Telmo Pievani Research Unit in Evolutionary Biology, Department of Biology, University of Padua, Italy

Niche construction theory posits that organisms can act as important agents of selection, by modifying biotic and abiotic environmental conditions. To test the presence and the evolutionary impact of niche construction, some criteria have been identified (Matthews et al 2014): 1- an organism must significantly modify environmental conditions, 2- organism-mediated environmental modifications must influence selection pressures on a recipient organism; 3- there must be an evolutionary response in at least one recipient population caused by the environmental modification. Homo sapiens has begun, by the late Pleistocene (Boivin et al 2016), in engaging in niche construction activities that meet those criteria, being however recognizable for some distinctive features: the rapidity of the processes, their global impact, the strong ecological inheritance of anthropic environmental modifications. Here we aim in particular at assessing the evidence for the third criterion, thus providing arguments for alterations in evolutionary trajectories due to anthropic activities. Moreover, human modifications have often brought to significant deleterious environmental consequences: it has been established that we’re losing ~11,000 to 58,000 species annually, changing ecosystem functions and services in a dramatic way (Dirzo et al 2014); by altering the climate, we have altered chemical composition of water, species distribution and their habitat. We discuss the extent to which global warming and biodiversity loss fall within the definition of niche construction activities. In fact, niche construction is often referred to as an adaptive process and global warming and biodiversity loss can indeed be intended as a by-product of adaptive human activity in the short term. However, this same activity might not result an adaptive one in the long term, as impoverished ecosystems might feedback negatively on human prosperity. Therefore, human niche construction so described might take the form of an evolutionary trap for Homo sapiens itself, resulting in a no longer adaptive process.

124 P27 A weapons-testes tradeoff in males extends to female traits Christine W Miller, Paul N Joseph, Rebecca M Kilner, Zachary Emberts University of Florida, USA

Trade‐offs among costly fitness traits are a central tenet of evolutionary theory. An organism cannot optimally invest in all traits associated with reproduction, development and survival, but must balance competing functions. Despite decades of research on such life history tradeoffs, we still know remarkably little about what governs resource allocation decisions. In this experimental study, we compared trait size in males and females and used the phenomenon of autotomy (dropping of limbs) as a form of phenotypic engineering to reveal allocation patterns. In Narnia femorata (Hemiptera: Coreidae) males use their hind limbs as weapons in male-male contests over access to females, while females use their hind limbs only for locomotion. We induced hind-limb autotomy in a sample of both males and females and measured resulting gonadal size to test the extent to which tradeoffs between sexually-selected weapons and testes in males are found in the homologous traits in females. As expected, insects that lost a hind limb during development grew larger gonads. Further, when we adjusted for physical differences in size, female ovaries increased more than male testes following autotomy. Thus, a documented weapon-testes tradeoff in males is also a limb-ovary tradeoff in females, a similarity that was not expected. These results suggest current theory on resource allocation may place an overemphasis on trait roles, while resource allocation patterns may more closely follow tissue type and expense of traits.

125 P28 Assessing adaptive and non-adaptive evolution of the glycolytic enzyme triosephosphate isomerase Ricardo Muñiz Trejo, Sergio Romero Romero, Daniel Alejando Fernández Velasco Universidad Nacional Autónoma de México, Mexico

Protein evolution studies have been dominated by accounts that explain physicochemical phenomena such as reversibility, folding, stability, oligomeric state, etc., as adaptive features, aligning with the tenets of Standard Evolutionary Theory. However, very few of them have inquired into the validity of these hypotheses from a broader evolutionary perspective. Understanding the underlying evolutionary processes in biochemistry might provide us with a better way to explain how evolution shapes the properties of macromolecules and how these properties set evolutionary pathways. In this work, we are interested in exploring which evolutionary processes, including those highlighted by the Extended Evolutionary Synthesis, are changing (or not) the thermostability of the glycolytic enzyme triosephosphate isomerase (TIM) in thermophilic archaea. For this enzyme, a change in the oligomeric state, from a dimer to a tetramer, is assumed to be the adaptation to life at high temperatures in Archaea. We are characterizing recombinant TIMs from archaea of the order Methanococcales with different growth temperatures (38 - 85 °C). Our experiments show that both tetrameric state and high thermostability (>90 °C) are preserved in TIMs from mesophilic representatives. Furthermore, hyperthermophilic representatives retain high catalytic activity even at 25 °C. These results suggest that there are some phylogenetic constraints involved in the evolution of TIMs from this order that are keeping these enzymes with high thermostability. Further studies regarding the thermostability of other proteins from these organisms need to be performed, but our results are coherent with other works that propose a readaptation of thermophilic Methanococcales, since thermostable proteins may contribute to the evolvability of this clade towards life at high temperatures.

126 P29 The evolution of hierarchical structure: separating the causes and consequences of evolvability Frederick J Nash, Loizos Kounios, Kostas Kouvaris, Richard Watson, Danesh Tarapore University of Southampton, UK

It has previously been suggested that the evolutionary causes of hierarchical organisation in biological systems are different from the evolutionary consequences of that hierarchical organisation. Specifically, it has been shown that evolved hierarchy can have the long-term consequence of facilitating evolvability, but that the short-term selective pressures that cause hierarchy to evolve can be a side effect of selection in systems where regulatory connections are costly. Here we investigate the conditions for the evolution of hierarchy and its relationship to evolvability in the context of development controlled with a gene regulation network. Such networks can exhibit non-modular structures, modular but not hierarchical structures, and modular structures with internal hierarchy. Using this model, we identify conditions under which hierarchical structures are favoured by short-term selection before any impact on evolvability may be observed. We rule-out the long-term advantages of evolvability as a necessary cause, and present cases where hierarchy evolves in static selective environments where no evolvability benefit can be observed. These results show that the distinction between the short-term selective causes of hierarchy and the long-term consequences of hierarchy can be much more distinct than previously documented, and can even be separated completely. We describe experimental modifications where hierarchy facilitates evolvability but does not evolve and cases where hierarchy evolves without facilitating evolvability. We discuss what would be required to determine whether the occurrence of hierarchy in nature evolved for evolvability or for other reasons, and whether there might be a deeper reason why short-term and long-term benefits might be coincident in natural systems.

127 P30 Spandrels and trait delimitation: no such thing as “architectural constraint” Mark E Olson Instituto de Biología, Universidad Nacional Autónoma de México, Mexico

40 years ago, Gould and Lewontin used the metaphor of a building’s “spandrels” to highlight that organismal traits could be the inevitable consequence of organismal construction, with no alternative configurations possible. Because adaptation by natural selection requires variation, regarding a trait incapable of variation as an adaptation could be a serious error. Gould and Lewontin’s exhortation spurred biologists’ efforts to investigate biases and limiations in development in their studies of adaptation, a major methodological advance. But in terms of the metaphor itself, over the past 40 years there are virtually no examples of “spandrels” in the primary literature. Moreover, multiple serious confusions in the metaphor have been identified and clarified, for example that the “spandrels” of San Marco are pendentives, and pendentives are perfect examples of adaptation. I look back over the sparse empirical fruits of the “spandrels” metaphor, and ask what the clarifications of the past 40 years mean for biological theory and practice. I conclude that if there is anything to be rescued from the clarified spandrels metaphor, it not “constraint” at all. Instead, it is the still-unresolved issue of trait delimitation, which is how to parse organisms into subsets that are tractable and biologically appropriate for study.

128 P31 With a little help from my friends: the role of the microbiota in dung beetle diversification Erik S Parker, Daniel B Schwab, Armin P Moczek Indiana University - Bloomington, USA

Understanding the processes that enable and shape the generation of biodiversity is a major objective of evolutionary biology. Traditionally, explanations have focused on the role of divergent selective pressures acting on generations of individuals. However, increasing appreciation for the vital role of host-associated microbial symbionts has added complexity to this notion of individuality, and raised the possibility that selection may occasionally, or perhaps frequently, act on teams of cooperating taxa rather than individuals. Yet few study systems exist where such perspectives can be assessed experimentally. My research aims to address this shortcoming by exploring the role gut symbionts play in the diversification of their dung beetle hosts. Despite feeding on nutritionally limited mammal dung throughout all stages of their life, Onthophagus dung beetles have been extraordinarily successful both ecologically and evolutionarily as measured by their expansion onto every continent except Antarctica, specialization onto an enormous array of dung types, and speciation into well over 2000 extant species. Using a combination of microbiota removal and transplant experiments we have shown that Onthophagus larvae inherit their gut microbiota vertically through a maternal fecal deposit known as a pedestal, that pedestal-derived microbiota enhance larval growth and survival especially under stress as well as protect against pathogens, and that different beetle host species have specialized onto different and functionally non-equivalent microbiota. Current efforts utilize recently established exotic dung beetle populations to assess host microbiota fidelity during the colonization of novel habitats as well as the role of microbiota in limiting, enabling, or biasing host range expansions. My talk will explore the most recent findings of this research area and their larger implications for understanding the role of host-microbiota interactions in the generation of biological diversity.

129 P32 Transposable elements: the unexpected junk that shaped genomes and challenged evolutionary biology Valentina Peona Uppsala University, Sweden

Barbara McClintock first described transposable elements in 1951 but her discovery remained ignored for decades, probably because her system of ""controlling elements"" did not fit any known genetic model although they shared some conceptual similarities with the (already at the time well-known) operons. In addition to be ignored, in the 1980 two important reviews published in the same Nature issue labelled and stigmatized repetitive DNA as junk DNA with no phenotypical relevance even though transposable elements themselves have been discovered because of their phenotypical impact. This label is still in use but thanks to scientific and technological advancement of the last decades, we have been able to appreciate them as a great substrate for evolution by molecular tinkering. Transposable elements changed our view of genes and genomes, from static entities to plastic and fluid. In fact, some transposable elements have been co-opted as telomerases, for the acquired immune system and placental evolution/development. They are inevitable genomic parasites from which genomes must be protected but also, in time, became essential components of eukaryotic genomes. Nowadays, we know that transposable elements regulate gene expression, play an active role during embryogenesis, provide neuronal plasticity and they also found use in the biotechnology industry as gene editing tools and certainly more roles and functionalities are waiting to be discovered. In this study I investigate the historic context in which Barbara McClintock discovery occurred, how the Modern Synthesis framework at that time influenced the reception and interpretation of her work, how transposable elements still challenge our comprehension of genome evolution (e.g. casting questions about what is actually neutral or under selection).

130 P33 Weismann, Crick, and others: flows of information in evolution Arnaud Pocheville CNRS, and Université Paul Sabatier, France

In previous work I have argued that one classical principled reason to separate development and evolution is that they are supposed to concern different levels and time-scales. I have also pointed to new theoretical approaches that aim at questioning this separation. In this talk I will discuss another kind of principled reason for this separation, one pertaining to biological information. Classically, development has been supposed to deal with the expression of existing information (mostly genetic information). This simplifies development to a one-way flow of information (from genes to the environment) at the individual level. Evolution, on the other hand, especially by natural selection, is supposed to deal with the inscription of new information. This simplifies evolution to another one-way flow of information (from the environment to the genes), but now at the population level. I will mention why the separation of flows of information can be attractive and briefly connect it to the question of the separation of time-scales.

131 P34 Delineating the boundaries of adaptation by tracing histories of environmental induction Ahva L Potticary, Erin S Morrison, Alexander V Badyaev The University of Arizona, USA

Adaptations change by reconfiguration of their components or by modification of homeostatic systems that maintained stability of preceding states. However, comparison of extant adaptations offers only indirect insight into the relative importance of these processes in evolution. Here, we overcome this limitation by directly tracing coevolution of adaptation components (diverse carotenoids taken up by growing ornamental feathers), with adaptation boundaries (sensitivity of developmental decision to forgo the feathers’ structural differentiation that limits accumulation of carotenoids). We found that along known historical trajectories linking native and 48 recently established populations, evolutionary changes in developmental boundaries of adaptation were not specific to individual carotenoids, but instead reflected the extent to which carotenoids were integrated into organismal processes and were environmentally predictable. Across all historical sequences, uptake of internalized carotenoids did not change the boundaries of feather developmental response, instead modifying the rate and precision of the response. In contrast, accommodation of uncommon dietary carotenoids strongly modified both the rate and the pattern of developmental response, especially in new populations. We found that largely inductive effects of carotenoid uptake on feather differentiation in younger populations were progressively converted into anticipatory effects as populations persisted in local environments, and show that this was due to increasing organismal coordination of feather differentiation decisions. These findings strongly implicate cooption and modification of generalized stress buffering mechanisms that shield feather growth and differentiation (i.e., adaptation boundary) as a starting point for the evolution of greater expression of external pigments within intricately structured feathers.

132 P35 The interaction of learning and evolution can enable adaptive behaviours that are not presupposed in the learning ability David Prosser, Alfredo Rago, Markus Brede, Richard A Watson University of Southampton, UK

Learning is a highly adaptive form of phenotypic plasticity where a behaviour can change over an individual’s lifetime as a result of experience. Innate behaviours can also be adaptive but do not change over an individual’s lifetime. In the absence of a mechanism for the inheritance of acquired characteristics, learned behaviours cannot influence innate behaviours directly. Nonetheless, the genetic assimilation of learned behaviours, often described as the Baldwin Effect, enables learned behaviours to guide the genetic evolution of innate behaviours indirectly but in a systematic manner. Previous models of this effect have, however, presupposed the ability of lifetime learning to produce appropriate adaptive behaviours. This makes a ‘plasticity-first’ concept of evolution based on such models open to the criticism that, although genetic evolution may follow behind a high-fitness acquired behaviour, the ability to produce the high-fitness behaviour in the first place was presumably the result of prior genetic evolution. Here we use a computational model to study the interaction of learning and genetic evolution in a multi-peaked fitness landscape where high-fitness behaviours cannot be quickly and reliably discovered by the learning process alone (nor by genetic evolution alone). We find conditions where the effect of genetic evolution is to incrementally reduce the dimensionality of the behavioural space that learning explores, not simply to assimilate the solutions that learning has already found, but in a way that causes the learning process to find higher fitness behaviours more reliably than was previously possible. This bidirectional interaction between evolution and learning is a form of plasticity first evolution capable of producing adaptations that are neither presupposed in the learning ability nor provided by prior genetic evolution.

133 P36 Naturalising the language faculty: the role of structures and developmental processes Giuliana Pulvirenti, Salvatore Ivan Amato, Alessandra Falzone University of Messina, Italy

The form/function dichotomy is one of the oldest and conceptually most pregnant distinction made in order to investigate the nature of organismal traits. Although the use of these two fundamental concepts is widespread in the field of contemporary evolutionary biology, their application is not so straightforward and can be theoretically problematic depending on the level of analysis considered (Love 2007). Moreover, in recent years this topic has gained a larger amount of attention due to insights obtained from Evo/Devo approaches (Minelli 2003; West-Eberhard 2003). The overmentioned distinction is relevant also among cognitive strains of research that ought to describe and explain in evolutionary terms complex and species-specific behaviours adopting the comparative method. The case of language faculty is, in this respect, paradigmatic. The present work aims at understanding language as a complex biological function while defending the necessity to pay attention to structural elements and ontogenetic processes to properly characterize it and analyse it from an evolutionary point of view. Most of the hypothesis formulated in this area suffer of a functionalist bias, often neglecting the role that morphological structures as well as ontogenetic processes exert on the emergence, development and evolution of animal behaviors, thus offering at best only a partial view. The interplay of these structural elements is modulated in and by the socio-ecological context in which the organism is to live in, constraining, while at the same time providing, a range of functional possibilities. That is to say functions are both epistemologically and chronologically dependent on forms and structures. This aspect has been heavily recognised among new embodied approaches to cognition (Shapiro 2004), in contrast to genetic and cerebrocentric/computational ways of thinking about human cognitive functions (Berwick & Chomsky 2016).

134 P37 Teleology and evolution Rasmus Sandnes Haukedal Durham University, UK

In Organisms, Agency, and Evolution (2015), D. M. Walsh argues that a new synthesis – which goes beyond the Modern Synthesis (MS) – cannot merely be about extending the scope of what evolution is about but must thoroughly reformulate its main tenets. It is thus wrong to assume that the basic framework of the MS could be left intact by a new model. We must formulate a comprehensive new theory, in which history is reintroduced into evolutionary thinking – not merely add factors with evolutionary significance to a pre-existing framework. In Walsh’s view, what is at stake in this reorientation is defining what counts as ‘evolutionary’ in the first place. He proposes an ‘a posteriori’ approach, in which evolutionary significant factors are not given but established retroactively. This means that such factors are transhistorical: while occurring at a specific historical point, they change the trajectory and dynamics of evolution. Furthermore, Walsh states that a new evolutionary synthesis demands the restoration of some sort of teleology – which he associates with phenotypic plasticity. We find the same idea in Scott Turner’s Purpose & Desire (2017), where it is also linked up with homeostasis and self-organization. They both argue that teleological explanations are indispensable to biology. I will examine what this move entails and apply it as an analytic tool to judge different attempts at transgressing the MS framework. Evidently, rethinking individuality is required, as is the inclusion of more factors than genetics. It is, however, not clear how Walsh’s ‘situated Darwinism’ differs from other attempts at going beyond MS, and whether these proposals are complementary or not. They all share an emphasis on the developmental effects on evolution, linked up with the question of how variation is produced. It seems the question of teleology might ultimately be what separates them.

135 P38 The evolution of adaptive epigenetic inheritance Zuzana Sekajová, Elena Rosa, Irja Ida Ratikainen, Martin Lind Uppsala University, Sweden

Environmental heterogeneity can influence the evolution of inheritance systems. If the environment is stable, genetic specialization should be favored by selection. If the environment is slowly fluctuating (cycle length spanning several generations), then the parental environment represents a reliable cue to the offspring and epigenetic inheritance is predicted to evolve. In the contrast, if the environment is fluctuating fast and unpredictably, there is no correlation between parental and offspring environment and phenotypic plasticity, bet hedging and/or generalism are predicted to be adaptive. My overall aim is to investigate the role of environmental heterogeneity for the evolution of inheritance systems with special focus on epigenetic inheritance. I will test this using experimental evolution in nematode worm Caenorhabditis remanei by manipulating temperature environment. Worms adapted to 20°C were selected in four regimes differing in cycle length for 113 non-overlapping generations: Warm with constant 25°C, where 25°C is a novel and mildly stressful environment, Cold with constant 20°C, Cyclic where temperature cycles between 20°C and 25°C every 8 generations and Random where temperature fluctuates randomly between 20°C and 25°C. I will conduct trans-generational phenotypic assays in a fully factorial design (parental temperature x offspring temperature) in which I will test worms for the key life history traits as reproduction and growth. I predict the evolution of epigenetic inheritance in worms from cyclic environments, specialization in worms from constant environments and the evolution of plasticity/bet hedging in worms evolving in random environment. In addition, I will investigate the molecular basis and regulation of plastic, epigenetic and genetic trait induction by small RNA sequencing which is the main epigenetic mechanism in the nematode worms.

136 P39 Unique evolutionary properties of plastic traits: an empirical study of erosion of phenotypic plasticity under relaxed selection Helen Spence-Jones, Michael M Webster, Kevin N Laland University of St Andrews, UK

While there are many hypotheses concerning the dynamics of plasticity which have proof-of-concept (Baldwin effect, plasticity-first, plasticity-mediated persistence etc), few of them have been found to be universal rules. I suggest that this is because plasticity is such a broad topic; dynamics are specific to subtypes of plasticity. Predicting the evolutionary dynamics of a plastic trait, and the applicability of hypotheses concerning this, requires understanding of when and how the underlying mechanisms of plasticity affect trait evolution. While many aspects of plasticity are only quantitively different from non-plastic traits, there are some unique properties - generally to do with decoupling of genotypic & phenotypic variation and the development of novel traits. These unique properties of phenotypic plasticity come together in the response of a population to a novel environment (one which has been under relaxed selection): here there is the potential for extrapolation of reaction norms to generate novel traits, and/or release of previously-selection-shadowed cryptic genetic variation. Here I present the results from my research into erosion of salinity-based plasticity under relaxed selection in threespine stickleback populations, in an attempt to investigate the dynamics of plastic traits under relaxed selection and potentially link this to the mechanism of plasticity of the traits concerned. I show that trait plasticity does appear to erode under relaxed selection, but that patterns of erosion are not consistent between populations.

137 P40 A culturally driven plasticity hypothesis for the evolution of human life course Francesco Suman Research Unit in Evolutionary Biology, University of Padua, Italy

Homo sapiens' life history pattern possesses both fast and slow components, in a combination that is unique among the extant great apes. The role played by phenotypic plasticity as a non-genetic means of adaptation to evolutionary challenges is still debated today. While life history parameters are shaped at a species level by genetic adaptations via natural selection, they remain very sensitive to changes in the environment. Relying on evidence from primates, extinct hominins and extant humans and on key explanatory tools of the Extended Evolutionary Synthesis (phenotypic plasticity, niche construction and inclusive inheritance), it is likely that environmentally induced plasticity led the way in human life history evolution, promoting subsequent genetic accommodation (Kuzawa and Bragg, 2012). It will be shown that the increase in brain size (modified brain growth and developmental rates) in hominin evolution was dependent on the appearance of certain life-history traits, which are in turn dependent on modified extrinsic mortality rates (Robson and Wood, 2008; Hublin et al., 2015). To the extent that culturally transmitted behaviors altered the selective pressures that acted on life history traits across generations during human evolution, a culturally driven plasticity dynamic shaping human life history traits can be identified. Two case studies in particular will be discussed: the human adaptations to the domestication of fire and the self-domestication hypothesis. The relationship between a relaxation of the selective regime and the emergence of plastic responses to changing environment will be also discussed. Finally, given this activity of cultural mitigation of selection altering environmental mortality rates is powered by an enlarged and reorganized brain, it is argued that the causal relationships in play in this evolutionary dynamic is to be conceived as reciprocal rather than linear (Laland et al., 2015).

138 P41 Unique, essential, and unknown: the enigma of taxonomically restricted, essential genes of unknown function Change Tan, Andrew Jones, PA Nelson Department of Biology, University of Missouri, USA

The advent of automated DNA sequencing in the mid-1990s enabled the development shortly thereafter of transposon mutagenesis screens of entire genomes, to determine what genes (and their protein or functional RNA products) were essential for cell viability under laboratory conditions. Two decades later, these experiments, as well as systematic or targeted gene deletion experiments, have consistently returned an unexpected result: many essential genes are (1) taxonomically restricted in their distribution, not universally, or even widely, shared, but also (2) when annotated, these genes are most often classified as "unknown function." Evident already in the initial transposon mutagenesis screens of Mycoplasma genitalium (Hutchison et al. 1999), where approximately one third of the essential genes were listed as "unknown function," the same "unique, essential, and unknown function" signal has been found in all three domains of life, e.g., in the proteobacterium Caulobacter crescentus (Christen et al. 2011), the archaean Sulfolobus islandicus (Zhang et al. 2018), and the eukaryote Drosophila melanogaster (Chen et al. 2010). We synthesize the findings of all whole genome transposon mutagenesis and targeted deletion experiments to date, with respect to their fraction of "unique, essential, and unknown" sequences, and consider the theoretical consequences of this pattern for theories of cell function and evolution.

139 P42 The evolution of relatedness and evolutionary transitions in individuality Christoph Thies, Markus Brede, Frederick J Nash, Richard A Watson University of Southampton, UK

Fraternal transitions in individuality, such as the transition from unicellular to multicellular organisms, are central to the evolution of complex life. Such a transition involves and depends on an increase in the genetic relatedness of the particles within collectives, such as cells within organisms. Most models of social evolution take relatedness to be a parameter defined by exogenous factors. However, this assumption precludes analysis of how this crucial population property arises as a product of the evolutionary process on the particle level. For such an analysis, relatedness must be endogenised. Although some previous models have studied mechanisms, such as the evolution of group size or inbreeding, where relatedness is modified by individual traits under selection, general conditions for the evolution of relatedness have not been identified. Here we show that no net change in relatedness occurs when the interaction between particles is described as additive pairwise game with two strategies, i.e. a game without synergistic effects such as a standard Prisoner’s Dilemma. In additive games, selection for beneficial organisation has opposite effects for each of the two strategies. While this result requires some assumptions, it illustrates the intuitive idea that a stable organisation of particles in collectives requires the alignment of benefits from that organisation. We show that the synergistic effects between particle types in non-additive games, such as coordination/anti-coordination games, cause such an alignment and facilitate the evolution of relatedness in a population. Focusing on anti- coordination games as models of functional complementarity between particle traits, we study how plastic trait expression evolves in concert with relatedness. This allows us to identify conditions under which particles organise into the genetically homogeneous and phenotypically heterogeneous collectives that are the characteristic consequence of a fraternal transition in individuality.

140 P43 Deconstructing reef complexity: relating coral diversity, rugosity and fractal dimension Damaris Torres-Pulliza, Maria Dornelas, Joshua S Madin Macquarie University, Australia

The relationship between habitat structural complexity and biodiversity is foundational to understanding community ecology and informing conservation management strategies. For habitat engineering organisms, like the reef-building corals, the direction of the complexity-biodiversity relationship is not clear, because the organisms that create structural complexity often require complexity to settle as larvae. Meanwhile, studies looking at complexity-biodiversity relationships typically revolve around one metric, rugosity, and the assumption is that higher levels of rugosity lead to higher abundances and richness of reef organisms. Here, we test the strength of complexity- biodiversity relationships using two metrics that capture different aspects of 3D complexity (rugosity and fractal dimension) as well as explore reciprocal causation of the relationships. Metrics of structural complexity were quantified from photogrammetric surveys at 21 reef sites encircling Lizard Island in the Great Barrier Reef. We assessed the strengths of relationships between these two metrics in 336 two-by-two metre reef plots and the abundance and richness of coral species in the same plots. The results are nuanced: structural complexity indeed constructs niches for coral species at the same time as specific species growth forms drive habitat structural complexity. Rugosity and fractal dimension captured different elements of the reef complexity and produced contrasting associations with coral species richness and abundance. We discuss the implications for niche diversity and the expected size distributions of reef-associated organisms, like the reef fishes. A better understanding of the role of reef structural complexity on reef biodiversity is important to guide management efforts as reefs continue to rapidly change.

141 P44 Cultural evolution and neuroscience: a pathway to integration Ryutaro Uchiyama, Michael Muthukrishna London School of Economics and Political Science, UK

Humans have a remarkable capacity to accumulate adaptive behaviours in the form of cumulative cultural knowledge, and to transmit these traits across generations in parallel with genetic transmission. This notion of culture as a second system of inheritance has become increasingly influential in recent years, and is frequently proposed as an explanation of our ecological success. The usual subtext is that whereas genetic information is stored in DNA, cultural information is stored within brains and the machinery of high-fidelity social learning enables this information to be replicated across brains. While this picture is not incorrect, it fails to capture much of what makes the human brain–culture interface so effective as an adaptive mechanism. In order to rectify this, we integrate evidence from neuroscience and cultural evolution, and lay out the following series of arguments: (1) Although neural plasticity is usually discussed in the context of how the nervous system responds to deviations from normative input, we should instead view it in terms of the space of neurophenotypic variation that it enables. (2) Culture exploits neural plasticity to explore this neurophenotypic space. (3) In most species, neurophenotypic variation is driven predominantly by environmental variation (via learning), but the intrinsic evolutionary dynamics of cumulative culture allow it to explore neurophenotypes not only in the absence of environmental change, but also beyond the range of variation that could plausibly be achieved by environmental input. Culture thus realizes functional fit between brains and environments in an indirect manner, by serving as a rapidly evolvable surrogate of environmental input. (4) In order to understand human brain function, we need to look at extant cultural variation as well as cultural evolutionary dynamics.

142 P45 Lost (and found) in translation: circulation of EES-related concepts in two scientific communities in Mexico Francisco Vergara-Silva, Alejandro Fábregas-Tejeda Instituto de Biología, Universidad Nacional Autónoma de México, Mexico

Academic institutions in the United States of America (USA) concentrate most of the criticism to the Extended Evolutionary Synthesis (EES) from the Standard Evolutionary Theory (SET). Given that the hegemonic version of biological evolutionism practiced and taught in the rest of the American continent –including Canada and the assemblage of (mostly) Spanish- and Portuguese-speaking countries usually identified as ‘Latin America’– is deeply influenced by SET-aligned research from the USA, inquiry into the reception of the core proposals of the EES outside of that country is of potential interest to evaluate the standing of alternative evolutionary perspectives at local/regional and international scales. Informed by debates on the ‘circulation of scientific knowledge’ held mostly in history of science circles, in this work we present a comparison of responses from two research/teaching communities in Mexico to the central tenets of the EES. The two communities addressed here correspond to separate academic subdisciplines with legitimate links to evolutionary theory –namely, biological anthropology and (as expected) evolutionary biology. Although in both cases we identify a polarization of positions –i.e., researchers place themselves in favor or against the EES in both Mexican communities– we identify that EES-related ideas have circulated with less resistance and/or have been received more sympathetically by biological anthropologists than by evolutionary biologists. We discuss epistemic and sociological explanations for these results, ranging from the entrenchment of adaptationist/selectionist thinking and the hardening of population genomics (both in ‘acclimatized’ versions) to the power structure in institutionalized life sciences in Mexico, where prominent SET-sympathetic evolutionary biologists have held high-level positions for decades. Finally, we argue that ethnography-laden, locally focused historiography of science studies could foster collaborations with EES-oriented scholars –including practicing evolutionists, as well as historians, philosophers and sociologists of science– currently working in traditional ‘metropolitan’ centers of knowledge production relevant to contemporary evolutionism.

143 P46 Testing the flexible stem hypothesis in stickleback fish: is ancestral morphological plasticity adaptive? Matthew A Wund, Nirmiti Borkhetaria, Rajath Kenath Department of Biology, The College of New Jersey, USA

One way in which phenotypic plasticity might play a constructive role in evolutionary processes is by influencing the pattern of phenotypic diversification during adaptive radiation (the “flexible stem hypothesis,” sensu West-Eberhard). If an ancestral stem group repeatedly colonizes alternative environments, then phenotypic plasticity repeatedly produces alternative phenotypes, which can then be refined by natural selection. This scenario would contribute to the evolution of parallel ecotypic variation within an adaptive radiation. The threespine stickleback radiation is ideally suited for testing the flexible stem hypothesis because we can compare patterns of phenotypic plasticity in the extant, oceanic ancestor to the phenotypes of derived, freshwater ecotypes. We have previously documented patterns of ancestral morphological plasticity consistent with the flexible stem hypothesis. Our goal in this experiment is to determine whether that plasticity confers an advantage in simulated open and shallow water environments, which would further support the hypothesis that ancestral patterns of plasticity biased the repeated evolution of limnetic and benthic ecotypes. We reared oceanic stickleback in simulated open and shallow water environments, respectively. After five months, we tagged and weighed all fish, switched half of the individuals into the alternate treatment, and reduced rations to promote competition. After seven weeks under reduced rations, we again weighed all individuals, predicting that fish who remained in their original treatment would outcompete those with a mismatched morphology. We found general support for our hypothesis: fish who remained in their original developmental environment grew more than those who were switched to the alternative environment, and we could identify specific differences in body shape that were most successful under each condition. Finally, fish that moved to the alternative environment continued to exhibit plasticity, such that after five months, they closely resembled fish that had developed exclusively in those environments.

144 P47 Quantifying niche constructing traits in reef corals Kyle JA Zawada, Joshua S Madin, Maria Dornelas University of St Andrews, UK

Morphology constrains how all organisms interact with their environment and other organisms, conditioning the environmental and biological contexts in which they are successful. For autogenic ecosystem engineers such as reef corals, morphological traits determine not only their fitness, but also the ways in which they modify the environment. Hence, morphology determines niche construction processes. Here, we identify quantitative traits that capture the three main axes of variation in morphology of reef building corals: colony volume compactness, surface area complexity and top- heaviness. We posit how variation in coral colony morphology translates to variation in ecosystem function and niche construction processes important for both corals and reef-associated taxa. These morphological traits can be used to test hypotheses about the ecological and evolutionary effects of coral niche construction.

145 P48 Two senses of biological possibility Jason Zinser University of Wisconsin - Stevens Point, USA

Modal concepts are often employed in the evolutionary biological literature despite the lack of analysis of how these concepts function in the biological domain. What does it mean to say, for example, that something is biologically possible or impossible? Is there a sense of modality that is unique to biology? I argue that there are at least two senses of possibility in biology: a broad sense and a restricted sense. Dennett (1995) described a broad sense biological possibility where to be possible is to be genetically accessible: if X is possible, then there is a genotype that codes for X. Among other things, this sense is not epistemically fruitful (and Dennett, it seems, ultimately agrees). Fortunately, I argue that there is a second, restricted sense of biological possibility that is both metaphysically grounded and interesting. This restricted sense ties possibility to the variation potential of actual or historical lineages. I will rely on the homeostatic property cluster kind conception of essentialism in order to ground lineages to a set of essential property clusters. Once lineages have an essence, then possibility can be restricted to what variation is possible in that lineage. In other words, it becomes empirically interesting what variations are restricted to a particular lineage (e.g., how developmental constraints restrict variation). I will defend this view against challenges, for example Forber (2009), that argue that biological possibility is circumscribed by evolutionary forces, such as natural selection and drift. Finally, I will gesture toward the implications of this restricted sense of possibility for other philosophical disputes, such as biological explanations, evolvability, and novelty.

146 P49 Plasticity in mouthpart length in response to developmental diet in leaf-footed bugs Sara Zlotnik, Pablo E Allen, Christine W Miller University of Florida, USA

Invasive species have become increasingly common worldwide. One reason why some species become invasive while others cannot may be their capacity for developmental plasticity. Species with high plasticity may be able to tolerate a range of novel environmental conditions, including new dietary challenges. We are beginning a series of projects to examine the role of developmental plasticity in the invasion process. Here, we report our findings on the first project, where we examine developmental plasticity in mouthpart length in the invasive leaf-footed bug, Leptoglossus zonatus (Hemiptera: Coreidae). This insect feeds on an astounding array of plants from multiple families and may grow longer or shorter mouthparts during development to make the most of the available food type. Such a capacity may contribute to its success as an invader. To investigate this diet-induced plasticity, we raised insects on multiple food types and measured their resulting mouthpart lengths. Our study provides insight into how invasive animals cope with dynamic environmental conditions, such as changing food availability, while colonizing and spreading through their nonnative range.

147 P50 Evolution and development of polyphenisms: a mechanistic model based on bistable switches Roman Zug, Tobias Uller Lund University, Sweden

Polyphenisms represent a form of adaptive phenotypic plasticity that gives rise to discrete alternative phenotypes, based on a developmental switch. However, existing models for the evolution of polyphenisms do not consider the role of development and therefore cannot explain how the switching behavior emerges and evolves. Our aim here is to reveal how polyphenisms evolve by modeling the evolution of their underlying developmental regulatory architecture. In particular, we hypothesize that polyphenisms are based on bistable switches in the underlying gene regulatory network (GRN), which usually requires positive feedback and ultrasensitivity. To test this hypothesis, we develop a theoretical mechanistic model that focuses on the evolution of the GRN responsible for the polyphenism switch. Using individual-based simulations, we explore the conditions (in terms of timescale and predictability of environmental variation) under which both positive feedback and ultrasensitivity emerge in the GRN, thus giving rise to a polyphenism switch. We will also present preliminary findings on the implications of the regulatory architecture for the evolution of polyphenisms. Our model can be applied to a wide range of polyphenisms and will eventually allow us to address critical yet unexplored questions, and make testable predictions, about the evolution of polyphenisms and how it is contingent upon underlying regulatory conditions.

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