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The makeup of the modern horse: a history of the biological changes introduced by human management

Total Cost €


EC-Contrib. €






 PEGASUS project word cloud

Explore the words cloud of the PEGASUS project. It provides you a very rough idea of what is the project "PEGASUS" about.

gather    domestication    modern    animal    epigenomic    impacted    societies    communication    isotopic    diversity    tremendously    transportation    500    ideas    genomic    cavalry    follow    breeding    trajectory    historical    pontic    strategies    underlying    warfare    selective    economical    625    population    genetics    rapid    independently    biological    difficult    intensive    until    ancient    living    track    data    molecules    changed    evolutionary    domesticated    metagenomic    makeup    perhaps    chariotry    characterization    alone    stages    latest    politico    dietary    technologies    preferred    agricultural    archaeology    multiple    natural    horses    dna    creation    caspian    periods    steppe    transformed    physiological    productivity    circulation    reconstruct    transformation    wild    archaeozoological    battlefield    antique    archaeogenetics    starting    genetic    secondary    languages    almost    emergence    stimulated    populations    heavy    breeds    history    religions    unrivaled    individuals    husbandry    behavioural    explore    equestrian    reveal    horse    revolutionizing    iberia    people    trajectories    contexts    domestic   

Project "PEGASUS" data sheet

The following table provides information about the project.


Organization address
address: RUE MICHEL ANGE 3
city: PARIS
postcode: 75794

contact info
title: n.a.
name: n.a.
surname: n.a.
function: n.a.
email: n.a.
telephone: n.a.
fax: n.a.

 Coordinator Country France [FR]
 Total cost 1˙999˙555 €
 EC max contribution 1˙999˙555 € (100%)
 Programme 1. H2020-EU.1.1. (EXCELLENT SCIENCE - European Research Council (ERC))
 Code Call ERC-2015-CoG
 Funding Scheme ERC-COG
 Starting year 2016
 Duration (year-month-day) from 2016-12-01   to  2021-11-30


Take a look of project's partnership.

# participants  country  role  EC contrib. [€] 
2    THE UNIVERSITY OF EXETER UK (EXETER) participant 46˙593.00


 Project objective

The horse provided us with rapid transportation, an almost unrivaled secondary product that tremendously impacted the politico-economical trajectory of our societies, revolutionizing the circulation of ideas, people, languages, religions and communication. Horse chariotry and cavalry also changed warfare and beyond the battlefield new equestrian technologies have stimulated agricultural productivity. However, the 5,500 year long history of horse domestication and management, which transformed the natural evolutionary trajectory of wild horses into the more than 625 domestic breeds living today, is difficult to reconstruct from archaeology, history and modern genetics alone. Yet, with archaeogenetics, one can access the genetic information from past individuals and track in great detail past population trajectories. In this project, I propose to build on the latest advances in the analysis of ancient DNA molecules to gather new genomic, epigenomic and metagenomic information from ancient horses. This will be integrated with archaeozoological, isotopic and historical data to enhance our understanding of the multiple processes underlying the transformation of the animal that perhaps most impacted our history. Starting from the characterization of pre-domestic populations of wild horses, I will evaluate the genomic and dietary impact of early domestication stages and will explore whether horses were independently domesticated in Iberia and the Pontic-Caspian steppe. I will follow how the emergence of chariotry and the development of heavy cavalry impacted the horse’s behavioural, physiological and biological makeup. I will reveal the horse characteristics that were preferred in various historical contexts and will investigate a diversity of management strategies and husbandry conditions to reveal their impact on horses, from classical and late antique periods until the recent creation of modern breeds by means of intensive selective breeding.


year authors and title journal last update
List of publications.
2019 Ludovic Orlando, Pablo Librado
Origin and Evolution of Deleterious Mutations in Horses
published pages: 649, ISSN: 2073-4425, DOI: 10.3390/genes10090649
Genes 10/9 2019-09-20
2019 Xuexue Liu, Yanli Zhang, Yefang Li, Jianfei Pan, Dandan Wang, Weihuang Chen, Zhuqing Zheng, Xiaohong He, Qianjun Zhao, Yabin Pu, Weijun Guan, Jianlin Han, Ludovic Orlando, Yuehui Ma, Lin Jiang
EPAS1 gain-of-function mutation contributes to high-altitude adaptation in Tibetan horses
published pages: , ISSN: 0737-4038, DOI: 10.1093/molbev/msz158
Molecular Biology and Evolution 2019-08-30
2018 Gabriel Renaud, Bent Petersen, Andaine Seguin-Orlando, Mads Frost Bertelsen, Andrew Waller, Richard Newton, Romain Paillot, Neil Bryant, Mark Vaudin, Pablo Librado, Ludovic Orlando
Improved de novo genomic assembly for the domestic donkey
published pages: eaaq0392, ISSN: 2375-2548, DOI: 10.1126/sciadv.aaq0392
Science Advances 4/4 2019-06-18
2017 Peter D Heintzman, Grant D Zazula, Ross DE MacPhee, Eric Scott, James A Cahill, Brianna K McHorse, Joshua D Kapp, Mathias Stiller, Matthew J Wooller, Ludovic Orlando, John Southon, Duane G Froese, Beth Shapiro
A new genus of horse from Pleistocene North America
published pages: , ISSN: 2050-084X, DOI: 10.7554/eLife.29944
eLife 6 2019-06-18
2017 Mikkel Schubert, Marjan Mashkour, Charleen Gaunitz, Antoine Fages, Andaine Seguin-Orlando, Shiva Sheikhi, Ahmed H. Alfarhan, Saleh A. Alquraishi, Khaled A.S. Al-Rasheid, Richard Chuang, Luca Ermini, Cristina Gamba, Jaco Weinstock, Onar Vedat, Ludovic Orlando
Zonkey: A simple, accurate and sensitive pipeline to genetically identify equine F1-hybrids in archaeological assemblages
published pages: 147-157, ISSN: 0305-4403, DOI: 10.1016/j.jas.2016.12.005
Journal of Archaeological Science 78 2019-06-18
2019 Lucie Zinger, Aurélie Bonin, Inger G. Alsos, Miklós Bálint, Holly Bik, Frédéric Boyer, Anthony A. Chariton, Simon Creer, Eric Coissac, Bruce E. Deagle, Marta De Barba, Ian A. Dickie, Alex J. Dumbrell, Gentile Francesco Ficetola, Noah Fierer, Luca Fumagalli, M. Thomas P. Gilbert, Simon Jarman, Ari Jumpponen, Håvard Kauserud, Ludovic Orlando, Johan Pansu, Jan Pawlowski, Leho Tedersoo, Philip F
DNA metabarcoding—Need for robust experimental designs to draw sound ecological conclusions
published pages: 1857-1862, ISSN: 0962-1083, DOI: 10.1111/mec.15060
Molecular Ecology 28/8 2019-08-05
2019 Antoine Fages, Kristian Hanghøj, Naveed Khan, Charleen Gaunitz, Andaine Seguin-Orlando, Michela Leonardi, Christian McCrory Constantz, Cristina Gamba, Khaled A.S. Al-Rasheid, Silvia Albizuri, Ahmed H. Alfarhan, Morten Allentoft, Saleh Alquraishi, David Anthony, Nurbol Baimukhanov, James H. Barrett, Jamsranjav Bayarsaikhan, Norbert Benecke, Eloísa Bernáldez-Sánchez, Luis Berrocal-Rangel, Fereid
Tracking Five Millennia of Horse Management with Extensive Ancient Genome Time Series
published pages: 1419-1435.e31, ISSN: 0092-8674, DOI: 10.1016/j.cell.2019.03.049
Cell 177/6 2019-08-05
2019 Kristian Hanghøj, Gabriel Renaud, Anders Albrechtsen, Ludovic Orlando
DamMet: ancient methylome mapping accounting for errors, true variants, and post-mortem DNA damage
published pages: , ISSN: 2047-217X, DOI: 10.1093/gigascience/giz025
GigaScience 8/4 2019-08-05
2019 Gabriel Renaud, Kristian Hanghøj, Thorfinn Sand Korneliussen, Eske Willerslev, Ludovic Orlando
Joint Estimates of Heterozygosity and Runs of Homozygosity for Modern and Ancient Samples
published pages: genetics.302057., ISSN: 0016-6731, DOI: 10.1534/genetics.119.302057
Genetics 2019-08-05
2018 Saskia Wutke, Edson Sandoval-Castellanos, Norbert Benecke, Hans-Jürgen Döhle, Susanne Friederich, Javier Gonzalez, Michael Hofreiter, Lembi Lõugas, Ola Magnell, Anna-Sapfo Malaspinas, Arturo Morales-Muñiz, Ludovic Orlando, Monika Reissmann, Alexandra Trinks, Arne Ludwig
Decline of genetic diversity in ancient domestic stallions in Europe
published pages: eaap9691, ISSN: 2375-2548, DOI: 10.1126/sciadv.aap9691
Science Advances 4/4 2019-05-25
2018 Charleen Gaunitz, Antoine Fages, Kristian Hanghøj, Anders Albrechtsen, Naveed Khan, Mikkel Schubert, Andaine Seguin-Orlando, Ivy J. Owens, Sabine Felkel, Olivier Bignon-Lau, Peter de Barros Damgaard, Alissa Mittnik, Azadeh F. Mohaseb, Hossein Davoudi, Saleh Alquraishi, Ahmed H. Alfarhan, Khaled A. S. Al-Rasheid, Eric Crubézy, Norbert Benecke, Sandra Olsen, Dorcas Brown, David Anthony, Ken Massy, Vladimir Pitulko, Aleksei Kasparov, Gottfried Brem, Michael Hofreiter, Gulmira Mukhtarova, Nurbol Baimukhanov, Lembi Lõugas, Vedat Onar, Philipp W. Stockhammer, Johannes Krause, Bazartseren Boldgiv, Sainbileg Undrakhbold, Diimaajav Erdenebaatar, Sébastien Lepetz, Marjan Mashkour, Arne Ludwig, Barbara Wallner, Victor Merz, Ilja Merz, Viktor Zaibert, Eske Willerslev, Pablo Librado, Alan K. Outram, Ludovic Orlando
Ancient genomes revisit the ancestry of domestic and Przewalski’s horses
published pages: 111-114, ISSN: 0036-8075, DOI: 10.1126/science.aao3297
Science 360/6384 2019-05-25
2019 Heidi M. Nistelberger, Albína Hulda Pálsdóttir, Bastiaan Star, Rúnar Leifsson, Agata T. Gondek, Ludovic Orlando, James H. Barrett, Jón Hallsteinn Hallsson, Sanne Boessenkool
Sexing Viking Age horses from burial and non-burial sites in Iceland using ancient DNA
published pages: 115-122, ISSN: 0305-4403, DOI: 10.1016/j.jas.2018.11.007
Journal of Archaeological Science 101 2019-05-25
2018 Jan E. Janečka, Brian W. Davis, Sharmila Ghosh, Nandina Paria, Pranab J. Das, Ludovic Orlando, Mikkel Schubert, Martin K. Nielsen, Tom A. E. Stout, Wesley Brashear, Gang Li, Charles D. Johnson, Richard P. Metz, Al Muatasim Al Zadjali, Charles C. Love, Dickson D. Varner, Daniel W. Bellott, William J. Murphy, Bhanu P. Chowdhary, Terje Raudsepp
Horse Y chromosome assembly displays unique evolutionary features and putative stallion fertility genes
published pages: , ISSN: 2041-1723, DOI: 10.1038/s41467-018-05290-6
Nature Communications 9/1 2019-05-25
2016 Saskia Wutke, Norbert Benecke, Edson Sandoval-Castellanos, Hans-Jürgen Döhle, Susanne Friederich, Javier Gonzalez, Jón Hallsteinn Hallsson, Michael Hofreiter, Lembi Lõugas, Ola Magnell, Arturo Morales-Muniz, Ludovic Orlando, Albína Hulda Pálsdóttir, Monika Reissmann, Matej Ruttkay, Alexandra Trinks, Arne Ludwig
Spotted phenotypes in horses lost attractiveness in the Middle Ages
published pages: , ISSN: 2045-2322, DOI: 10.1038/srep38548
Scientific Reports 6/1 2019-05-25
2017 Christina Warinner, Alexander Herbig, Allison Mann, James A. Fellows Yates, Clemens L. Weiß, Hernán A. Burbano, Ludovic Orlando, Johannes Krause
A Robust Framework for Microbial Archaeology
published pages: 321-356, ISSN: 1527-8204, DOI: 10.1146/annurev-genom-091416-035526
Annual Review of Genomics and Human Genetics 18/1 2019-05-25
2017 Michela Leonardi, Pablo Librado, Clio Der Sarkissian, Mikkel Schubert, Ahmed H. Alfarhan, Saleh A. Alquraishi, Khaled A. S. Al-Rasheid, Cristina Gamba, Eske Willerslev, Ludovic Orlando
Evolutionary Patterns and Processes: Lessons from Ancient DNA
published pages: syw059, ISSN: 1063-5157, DOI: 10.1093/sysbio/syw059
Systematic Biology 2019-05-25
2017 Robert J. Schaefer, Mikkel Schubert, Ernest Bailey, Danika L. Bannasch, Eric Barrey, Gila Kahila Bar-Gal, Gottfried Brem, Samantha A. Brooks, Ottmar Distl, Ruedi Fries, Carrie J. Finno, Vinzenz Gerber, Bianca Haase, Vidhya Jagannathan, Ted Kalbfleisch, Tosso Leeb, Gabriella Lindgren, Maria Susana Lopes, Núria Mach, Artur da Câmara Machado, James N. MacLeod, Annette McCoy, Julia Metzger, Cecilia Penedo, Sagi Polani, Stefan Rieder, Imke Tammen, Jens Tetens, Georg Thaller, Andrea Verini-Supplizi, Claire M. Wade, Barbara Wallner, Ludovic Orlando, James R. Mickelson, Molly E. McCue
Developing a 670k genotyping array to tag ~2M SNPs across 24 horse breeds
published pages: , ISSN: 1471-2164, DOI: 10.1186/s12864-017-3943-8
BMC Genomics 18/1 2019-05-25
2017 Pablo Librado, Cristina Gamba, Charleen Gaunitz, Clio Der Sarkissian, Mélanie Pruvost, Anders Albrechtsen, Antoine Fages, Naveed Khan, Mikkel Schubert, Vidhya Jagannathan, Aitor Serres-Armero, Lukas F. K. Kuderna, Inna S. Povolotskaya, Andaine Seguin-Orlando, Sébastien Lepetz, Markus Neuditschko, Catherine Thèves, Saleh Alquraishi, Ahmed H. Alfarhan, Khaled Al-Rasheid, Stefan Rieder, Zainolla Samashev, Henri-Paul Francfort, Norbert Benecke, Michael Hofreiter, Arne Ludwig, Christine Keyser, Tomas Marques-Bonet, Bertrand Ludes, Eric Crubézy, Tosso Leeb, Eske Willerslev, Ludovic Orlando
Ancient genomic changes associated with domestication of the horse
published pages: 442-445, ISSN: 0036-8075, DOI: 10.1126/science.aam5298
Science 356/6336 2019-05-25
2017 Jessica L. Metcalf, Se Jin Song, James T. Morton, Sophie Weiss, Andaine Seguin-Orlando, Frédéric Joly, Claudia Feh, Pierre Taberlet, Eric Coissac, Amnon Amir, Eske Willerslev, Rob Knight, Valerie McKenzie, Ludovic Orlando
Evaluating the impact of domestication and captivity on the horse gut microbiome
published pages: , ISSN: 2045-2322, DOI: 10.1038/s41598-017-15375-9
Scientific Reports 7/1 2019-05-25
2018 Michela Leonardi, Francesco Boschin, Konstantinos Giampoudakis, Robert M. Beyer, Mario Krapp, Robin Bendrey, Robert Sommer, Paolo Boscato, Andrea Manica, David Nogues-Bravo, Ludovic Orlando
Late Quaternary horses in Eurasia in the face of climate and vegetation change
published pages: eaar5589, ISSN: 2375-2548, DOI: 10.1126/sciadv.aar5589
Science Advances 4/7 2019-05-25
2018 William Taylor, Svetlana Shnaider, Aida Abdykanova, Antoine Fages, Frido Welker, Franziska Irmer, Andaine Seguin-Orlando, Naveed Khan, Katerina Douka, Ksenia Kolobova, Ludovic Orlando, Andrei Krivoshapkin, Nicole Boivin
Early pastoral economies along the Ancient Silk Road: Biomolecular evidence from the Alay Valley, Kyrgyzstan
published pages: e0205646, ISSN: 1932-6203, DOI: 10.1371/journal.pone.0205646
PLOS ONE 13/10 2019-05-25
2017 Valerie J. McKenzie, Se Jin Song, Frédéric Delsuc, Tiffany L. Prest, Angela M. Oliverio, Timothy M. Korpita, Alexandra Alexiev, Katherine R. Amato, Jessica L. Metcalf, Martin Kowalewski, Nico L. Avenant, Andres Link, Anthony Di Fiore, Andaine Seguin-Orlando, Claudia Feh, Ludovic Orlando, Joseph R. Mendelson, Jon Sanders, Rob Knight
The Effects of Captivity on the Mammalian Gut Microbiome
published pages: 690-704, ISSN: 1540-7063, DOI: 10.1093/icb/icx090
Integrative and Comparative Biology 57/4 2019-05-25
2017 T. Cucchi, A. Mohaseb, S. Peigné, K. Debue, L. Orlando, M. Mashkour
Detecting taxonomic and phylogenetic signals in equid cheek teeth: towards new palaeontological and archaeological proxies
published pages: 160997, ISSN: 2054-5703, DOI: 10.1098/rsos.160997
Royal Society Open Science 4/4 2019-05-25
2018 Peter de Barros Damgaard, Rui Martiniano, Jack Kamm, J. Víctor Moreno-Mayar, Guus Kroonen, Michaël Peyrot, Gojko Barjamovic, Simon Rasmussen, Claus Zacho, Nurbol Baimukhanov, Victor Zaibert, Victor Merz, Arjun Biddanda, Ilja Merz, Valeriy Loman, Valeriy Evdokimov, Emma Usmanova, Brian Hemphill, Andaine Seguin-Orlando, Fulya Eylem Yediay, Inam Ullah, Karl-Göran Sjögren, Katrine Højholt Iversen, Jeremy Choin, Constanza de la Fuente, Melissa Ilardo, Hannes Schroeder, Vyacheslav Moiseyev, Andrey Gromov, Andrei Polyakov, Sachihiro Omura, Süleyman Yücel Senyurt, Habib Ahmad, Catriona McKenzie, Ashot Margaryan, Abdul Hameed, Abdul Samad, Nazish Gul, Muhammad Hassan Khokhar, O. I. Goriunova, Vladimir I. Bazaliiskii, John Novembre, Andrzej W. Weber, Ludovic Orlando, Morten E. Allentoft, Rasmus Nielsen, Kristian Kristiansen, Martin Sikora, Alan K. Outram, Richard Durbin, Eske Willerslev
The first horse herders and the impact of early Bronze Age steppe expansions into Asia
published pages: eaar7711, ISSN: 0036-8075, DOI: 10.1126/science.aar7711
Science 360/6396 2019-05-25
2018 Ludovic Orlando
Late Bronze Age cultural origins of dairy pastoralism in Mongolia
published pages: 12083-12085, ISSN: 0027-8424, DOI: 10.1073/pnas.1817559115
Proceedings of the National Academy of Sciences 115/48 2019-05-25
2018 Theodore S. Kalbfleisch, Edward S. Rice, Michael S. DePriest, Brian P. Walenz, Matthew S. Hestand, Joris R. Vermeesch, Brendan L. O′Connell, Ian T. Fiddes, Alisa O. Vershinina, Nedda F. Saremi, Jessica L. Petersen, Carrie J. Finno, Rebecca R. Bellone, Molly E. McCue, Samantha A. Brooks, Ernest Bailey, Ludovic Orlando, Richard E. Green, Donald C. Miller, Douglas F. Antczak, James N. MacLeod
Improved reference genome for the domestic horse increases assembly contiguity and composition
published pages: , ISSN: 2399-3642, DOI: 10.1038/s42003-018-0199-z
Communications Biology 1/1 2019-05-25
2018 Pablo Librado, Ludovic Orlando
Detecting Signatures of Positive Selection along Defined Branches of a Population Tree Using LSD
published pages: 1520-1535, ISSN: 0737-4038, DOI: 10.1093/molbev/msy053
Molecular Biology and Evolution 35/6 2019-05-25

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