Tracking Five Millennia of Horse Management with Extensive Ancient Genome Time Series

Detalhes bibliográficos
Autor(a) principal: Fages, Antoine
Data de Publicação: 2019
Outros Autores: Hanghøj, Kristian, Khan, Naveed, Gaunitz, Charleen, Seguin-Orlando, Andaine, Leonardi, Michela, McCrory Constantz, Christian, Gamba, Cristina, Al-Rasheid, Khaled A.S., Albizuri, Silvia, Alfarhan, Ahmed H., Trbojević Vukičević, Tajana, Trixl, Simon, Tumen, Dashzeveg, Undrakhbold, Sainbileg, Usmanova, Emma, Vahdati, Ali, Valenzuela-Lamas, Silvia, Viegas, Catarina, Wallner, Barbara, Weinstock, Jaco, Allentoft, Morten, Zaibert, Victor, Clavel, Benoit, Lepetz, Sébastien, Mashkour, Marjan, Helgason, Agnar, Stefánsson, Kári, Barrey, Eric, Willerslev, Eske, Outram, Alan K., Librado, Pablo, Alquraishi, Saleh, Orlando, Ludovic, Anthony, David, Baimukhanov, Nurbol, Barrett, James H., Bayarsaikhan, Jamsranjav, Benecke, Norbert, Bernáldez-Sánchez, Eloísa, Berrocal-Rangel, Luis, Biglari, Fereidoun, Boessenkool, Sanne, Boldgiv, Bazartseren, Brem, Gottfried, Brown, Dorcas, Burger, Joachim, Crubézy, Eric, Daugnora, Linas, Davoudi, Hossein, de Barros Damgaard, Peter, de los Ángeles de Chorro y de Villa-Ceballos, María, Deschler-Erb, Sabine, Detry, Cleia, Dill, Nadine, do Mar Oom, Maria, Dohr, Anna, Ellingvåg, Sturla, Erdenebaatar, Diimaajav, Fathi, Homa, Felkel, Sabine, Fernández-Rodríguez, Carlos, García-Viñas, Esteban, Germonpré, Mietje, Granado, José D., Hallsson, Jón H., Hemmer, Helmut, Hofreiter, Michael, Kasparov, Aleksei, Khasanov, Mutalib, Khazaeli, Roya, Kosintsev, Pavel, Kristiansen, Kristian, Kubatbek, Tabaldiev, Kuderna, Lukas, Kuznetsov, Pavel, Laleh, Haeedeh, Leonard, Jennifer A., Lhuillier, Johanna, Liesau von Lettow-Vorbeck, Corina, Logvin, Andrey, Lõugas, Lembi, Ludwig, Arne, Luis, Cristina, Margarida Arruda, Ana, Marques-Bonet, Tomas, Matoso Silva, Raquel, Merz, Victor, Mijiddorj, Enkhbayar, Miller, Bryan K., Monchalov, Oleg, Mohaseb, Fatemeh A., Morales, Arturo, Nieto-Espinet, Ariadna, Nistelberger, Heidi, Onar, Vedat, Pálsdóttir, Albína H., Pitulko, Vladimir, Pitskhelauri, Konstantin, Pruvost, Mélanie, Rajic Sikanjic, Petra, Rapan Papeša, Anita, Roslyakova, Natalia, Sardari, Alireza, Sauer, Eberhard, Schafberg, Renate, Scheu, Amelie, Schibler, Jörg, Schlumbaum, Angela, Serrand, Nathalie, Serres-Armero, Aitor, Shapiro, Beth, Sheikhi Seno, Shiva, Shevnina, Irina, Shidrang, Sonia, Southon, John, Star, Bastiaan, Sykes, Naomi, Taheri, Kamal, Taylor, William, Teegen, Wolf-Rüdiger
Tipo de documento: Artigo
Idioma: eng
Título da fonte: Repositório Científico de Acesso Aberto de Portugal (Repositórios Cientìficos)
Texto Completo: http://hdl.handle.net/10451/42994
Resumo: Horse domestication revolutionized warfare and accelerated travel, trade, and the geographic expansion of languages. Here, we present the largest DNA time series for a non-human organism to date, including genome-scale data from 149 ancient animals and 129 ancient genomes (≥1-fold coverage), 87 of which are new. This extensive dataset allows us to assess the modern legacy of past equestrian civilizations. We find that two extinct horse lineages existed during early domestication, one at the far western (Iberia) and the other at the far eastern range (Siberia) of Eurasia. None of these contributed significantly to modern diversity. We show that the influence of Persian-related horse lineages increased following the Islamic conquests in Europe and Asia. Multiple alleles associated with elite-racing, including at the MSTN "speed gene," only rose in popularity within the last millennium. Finally, the development of modern breeding impacted genetic diversity more dramatically than the previous millennia of human management.
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spelling Tracking Five Millennia of Horse Management with Extensive Ancient Genome Time SeriesAnimalsAsiaBiological EvolutionBreedingDNA, AncientDomesticationEquidaeEuropeFemaleGenetic variationGenomeHistory, AncientHorsesMalePhylogenyHorse domestication revolutionized warfare and accelerated travel, trade, and the geographic expansion of languages. Here, we present the largest DNA time series for a non-human organism to date, including genome-scale data from 149 ancient animals and 129 ancient genomes (≥1-fold coverage), 87 of which are new. This extensive dataset allows us to assess the modern legacy of past equestrian civilizations. We find that two extinct horse lineages existed during early domestication, one at the far western (Iberia) and the other at the far eastern range (Siberia) of Eurasia. None of these contributed significantly to modern diversity. We show that the influence of Persian-related horse lineages increased following the Islamic conquests in Europe and Asia. Multiple alleles associated with elite-racing, including at the MSTN "speed gene," only rose in popularity within the last millennium. Finally, the development of modern breeding impacted genetic diversity more dramatically than the previous millennia of human management.ElsevierRepositório da Universidade de LisboaFages, AntoineHanghøj, KristianKhan, NaveedGaunitz, CharleenSeguin-Orlando, AndaineLeonardi, MichelaMcCrory Constantz, ChristianGamba, CristinaAl-Rasheid, Khaled A.S.Albizuri, SilviaAlfarhan, Ahmed H.Trbojević Vukičević, TajanaTrixl, SimonTumen, DashzevegUndrakhbold, SainbilegUsmanova, EmmaVahdati, AliValenzuela-Lamas, SilviaViegas, CatarinaWallner, BarbaraWeinstock, JacoAllentoft, MortenZaibert, VictorClavel, BenoitLepetz, SébastienMashkour, MarjanHelgason, AgnarStefánsson, KáriBarrey, EricWillerslev, EskeOutram, Alan K.Librado, PabloAlquraishi, SalehOrlando, LudovicAnthony, DavidBaimukhanov, NurbolBarrett, James H.Bayarsaikhan, JamsranjavBenecke, NorbertBernáldez-Sánchez, EloísaBerrocal-Rangel, LuisBiglari, FereidounBoessenkool, SanneBoldgiv, BazartserenBrem, GottfriedBrown, DorcasBurger, JoachimCrubézy, EricDaugnora, LinasDavoudi, Hosseinde Barros Damgaard, Peterde los Ángeles de Chorro y de Villa-Ceballos, MaríaDeschler-Erb, SabineDetry, CleiaDill, Nadinedo Mar Oom, MariaDohr, AnnaEllingvåg, SturlaErdenebaatar, DiimaajavFathi, HomaFelkel, SabineFernández-Rodríguez, CarlosGarcía-Viñas, EstebanGermonpré, MietjeGranado, José D.Hallsson, Jón H.Hemmer, HelmutHofreiter, MichaelKasparov, AlekseiKhasanov, MutalibKhazaeli, RoyaKosintsev, PavelKristiansen, KristianKubatbek, TabaldievKuderna, LukasKuznetsov, PavelLaleh, HaeedehLeonard, Jennifer A.Lhuillier, JohannaLiesau von Lettow-Vorbeck, CorinaLogvin, AndreyLõugas, LembiLudwig, ArneLuis, CristinaMargarida Arruda, AnaMarques-Bonet, TomasMatoso Silva, RaquelMerz, VictorMijiddorj, EnkhbayarMiller, Bryan K.Monchalov, OlegMohaseb, Fatemeh A.Morales, ArturoNieto-Espinet, AriadnaNistelberger, HeidiOnar, VedatPálsdóttir, Albína H.Pitulko, VladimirPitskhelauri, KonstantinPruvost, MélanieRajic Sikanjic, PetraRapan Papeša, AnitaRoslyakova, NataliaSardari, AlirezaSauer, EberhardSchafberg, RenateScheu, AmelieSchibler, JörgSchlumbaum, AngelaSerrand, NathalieSerres-Armero, AitorShapiro, BethSheikhi Seno, ShivaShevnina, IrinaShidrang, SoniaSouthon, JohnStar, BastiaanSykes, NaomiTaheri, KamalTaylor, WilliamTeegen, Wolf-Rüdiger2020-04-21T08:49:37Z20192019-01-01T00:00:00Zinfo:eu-repo/semantics/publishedVersioninfo:eu-repo/semantics/articleapplication/pdfhttp://hdl.handle.net/10451/42994engFages, A., Hanghoj, K., Khan, N., Gaunitz, C., Seguin-Orlando, A., Leonardi, M., . . . Orlando, L. (2019). Tracking Five Millennia of Horse Management with Extensive Ancient Genome Time Series. Cell, 177(6) 1419-1435 e1431. doi: 10.1016/j.cell.2019.03.0491097-417210.1016/j.cell.2019.03.049metadata only accessinfo:eu-repo/semantics/openAccessreponame:Repositório Científico de Acesso Aberto de Portugal (Repositórios Cientìficos)instname:Agência para a Sociedade do Conhecimento (UMIC) - FCT - Sociedade da Informaçãoinstacron:RCAAP2023-11-08T16:43:12Zoai:repositorio.ul.pt:10451/42994Portal AgregadorONGhttps://www.rcaap.pt/oai/openaireopendoar:71602024-03-19T21:55:54.115258Repositório Científico de Acesso Aberto de Portugal (Repositórios Cientìficos) - Agência para a Sociedade do Conhecimento (UMIC) - FCT - Sociedade da Informaçãofalse
dc.title.none.fl_str_mv Tracking Five Millennia of Horse Management with Extensive Ancient Genome Time Series
title Tracking Five Millennia of Horse Management with Extensive Ancient Genome Time Series
spellingShingle Tracking Five Millennia of Horse Management with Extensive Ancient Genome Time Series
Fages, Antoine
Animals
Asia
Biological Evolution
Breeding
DNA, Ancient
Domestication
Equidae
Europe
Female
Genetic variation
Genome
History, Ancient
Horses
Male
Phylogeny
title_short Tracking Five Millennia of Horse Management with Extensive Ancient Genome Time Series
title_full Tracking Five Millennia of Horse Management with Extensive Ancient Genome Time Series
title_fullStr Tracking Five Millennia of Horse Management with Extensive Ancient Genome Time Series
title_full_unstemmed Tracking Five Millennia of Horse Management with Extensive Ancient Genome Time Series
title_sort Tracking Five Millennia of Horse Management with Extensive Ancient Genome Time Series
author Fages, Antoine
author_facet Fages, Antoine
Hanghøj, Kristian
Khan, Naveed
Gaunitz, Charleen
Seguin-Orlando, Andaine
Leonardi, Michela
McCrory Constantz, Christian
Gamba, Cristina
Al-Rasheid, Khaled A.S.
Albizuri, Silvia
Alfarhan, Ahmed H.
Trbojević Vukičević, Tajana
Trixl, Simon
Tumen, Dashzeveg
Undrakhbold, Sainbileg
Usmanova, Emma
Vahdati, Ali
Valenzuela-Lamas, Silvia
Viegas, Catarina
Wallner, Barbara
Weinstock, Jaco
Allentoft, Morten
Zaibert, Victor
Clavel, Benoit
Lepetz, Sébastien
Mashkour, Marjan
Helgason, Agnar
Stefánsson, Kári
Barrey, Eric
Willerslev, Eske
Outram, Alan K.
Librado, Pablo
Alquraishi, Saleh
Orlando, Ludovic
Anthony, David
Baimukhanov, Nurbol
Barrett, James H.
Bayarsaikhan, Jamsranjav
Benecke, Norbert
Bernáldez-Sánchez, Eloísa
Berrocal-Rangel, Luis
Biglari, Fereidoun
Boessenkool, Sanne
Boldgiv, Bazartseren
Brem, Gottfried
Brown, Dorcas
Burger, Joachim
Crubézy, Eric
Daugnora, Linas
Davoudi, Hossein
de Barros Damgaard, Peter
de los Ángeles de Chorro y de Villa-Ceballos, María
Deschler-Erb, Sabine
Detry, Cleia
Dill, Nadine
do Mar Oom, Maria
Dohr, Anna
Ellingvåg, Sturla
Erdenebaatar, Diimaajav
Fathi, Homa
Felkel, Sabine
Fernández-Rodríguez, Carlos
García-Viñas, Esteban
Germonpré, Mietje
Granado, José D.
Hallsson, Jón H.
Hemmer, Helmut
Hofreiter, Michael
Kasparov, Aleksei
Khasanov, Mutalib
Khazaeli, Roya
Kosintsev, Pavel
Kristiansen, Kristian
Kubatbek, Tabaldiev
Kuderna, Lukas
Kuznetsov, Pavel
Laleh, Haeedeh
Leonard, Jennifer A.
Lhuillier, Johanna
Liesau von Lettow-Vorbeck, Corina
Logvin, Andrey
Lõugas, Lembi
Ludwig, Arne
Luis, Cristina
Margarida Arruda, Ana
Marques-Bonet, Tomas
Matoso Silva, Raquel
Merz, Victor
Mijiddorj, Enkhbayar
Miller, Bryan K.
Monchalov, Oleg
Mohaseb, Fatemeh A.
Morales, Arturo
Nieto-Espinet, Ariadna
Nistelberger, Heidi
Onar, Vedat
Pálsdóttir, Albína H.
Pitulko, Vladimir
Pitskhelauri, Konstantin
Pruvost, Mélanie
Rajic Sikanjic, Petra
Rapan Papeša, Anita
Roslyakova, Natalia
Sardari, Alireza
Sauer, Eberhard
Schafberg, Renate
Scheu, Amelie
Schibler, Jörg
Schlumbaum, Angela
Serrand, Nathalie
Serres-Armero, Aitor
Shapiro, Beth
Sheikhi Seno, Shiva
Shevnina, Irina
Shidrang, Sonia
Southon, John
Star, Bastiaan
Sykes, Naomi
Taheri, Kamal
Taylor, William
Teegen, Wolf-Rüdiger
author_role author
author2 Hanghøj, Kristian
Khan, Naveed
Gaunitz, Charleen
Seguin-Orlando, Andaine
Leonardi, Michela
McCrory Constantz, Christian
Gamba, Cristina
Al-Rasheid, Khaled A.S.
Albizuri, Silvia
Alfarhan, Ahmed H.
Trbojević Vukičević, Tajana
Trixl, Simon
Tumen, Dashzeveg
Undrakhbold, Sainbileg
Usmanova, Emma
Vahdati, Ali
Valenzuela-Lamas, Silvia
Viegas, Catarina
Wallner, Barbara
Weinstock, Jaco
Allentoft, Morten
Zaibert, Victor
Clavel, Benoit
Lepetz, Sébastien
Mashkour, Marjan
Helgason, Agnar
Stefánsson, Kári
Barrey, Eric
Willerslev, Eske
Outram, Alan K.
Librado, Pablo
Alquraishi, Saleh
Orlando, Ludovic
Anthony, David
Baimukhanov, Nurbol
Barrett, James H.
Bayarsaikhan, Jamsranjav
Benecke, Norbert
Bernáldez-Sánchez, Eloísa
Berrocal-Rangel, Luis
Biglari, Fereidoun
Boessenkool, Sanne
Boldgiv, Bazartseren
Brem, Gottfried
Brown, Dorcas
Burger, Joachim
Crubézy, Eric
Daugnora, Linas
Davoudi, Hossein
de Barros Damgaard, Peter
de los Ángeles de Chorro y de Villa-Ceballos, María
Deschler-Erb, Sabine
Detry, Cleia
Dill, Nadine
do Mar Oom, Maria
Dohr, Anna
Ellingvåg, Sturla
Erdenebaatar, Diimaajav
Fathi, Homa
Felkel, Sabine
Fernández-Rodríguez, Carlos
García-Viñas, Esteban
Germonpré, Mietje
Granado, José D.
Hallsson, Jón H.
Hemmer, Helmut
Hofreiter, Michael
Kasparov, Aleksei
Khasanov, Mutalib
Khazaeli, Roya
Kosintsev, Pavel
Kristiansen, Kristian
Kubatbek, Tabaldiev
Kuderna, Lukas
Kuznetsov, Pavel
Laleh, Haeedeh
Leonard, Jennifer A.
Lhuillier, Johanna
Liesau von Lettow-Vorbeck, Corina
Logvin, Andrey
Lõugas, Lembi
Ludwig, Arne
Luis, Cristina
Margarida Arruda, Ana
Marques-Bonet, Tomas
Matoso Silva, Raquel
Merz, Victor
Mijiddorj, Enkhbayar
Miller, Bryan K.
Monchalov, Oleg
Mohaseb, Fatemeh A.
Morales, Arturo
Nieto-Espinet, Ariadna
Nistelberger, Heidi
Onar, Vedat
Pálsdóttir, Albína H.
Pitulko, Vladimir
Pitskhelauri, Konstantin
Pruvost, Mélanie
Rajic Sikanjic, Petra
Rapan Papeša, Anita
Roslyakova, Natalia
Sardari, Alireza
Sauer, Eberhard
Schafberg, Renate
Scheu, Amelie
Schibler, Jörg
Schlumbaum, Angela
Serrand, Nathalie
Serres-Armero, Aitor
Shapiro, Beth
Sheikhi Seno, Shiva
Shevnina, Irina
Shidrang, Sonia
Southon, John
Star, Bastiaan
Sykes, Naomi
Taheri, Kamal
Taylor, William
Teegen, Wolf-Rüdiger
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dc.contributor.none.fl_str_mv Repositório da Universidade de Lisboa
dc.contributor.author.fl_str_mv Fages, Antoine
Hanghøj, Kristian
Khan, Naveed
Gaunitz, Charleen
Seguin-Orlando, Andaine
Leonardi, Michela
McCrory Constantz, Christian
Gamba, Cristina
Al-Rasheid, Khaled A.S.
Albizuri, Silvia
Alfarhan, Ahmed H.
Trbojević Vukičević, Tajana
Trixl, Simon
Tumen, Dashzeveg
Undrakhbold, Sainbileg
Usmanova, Emma
Vahdati, Ali
Valenzuela-Lamas, Silvia
Viegas, Catarina
Wallner, Barbara
Weinstock, Jaco
Allentoft, Morten
Zaibert, Victor
Clavel, Benoit
Lepetz, Sébastien
Mashkour, Marjan
Helgason, Agnar
Stefánsson, Kári
Barrey, Eric
Willerslev, Eske
Outram, Alan K.
Librado, Pablo
Alquraishi, Saleh
Orlando, Ludovic
Anthony, David
Baimukhanov, Nurbol
Barrett, James H.
Bayarsaikhan, Jamsranjav
Benecke, Norbert
Bernáldez-Sánchez, Eloísa
Berrocal-Rangel, Luis
Biglari, Fereidoun
Boessenkool, Sanne
Boldgiv, Bazartseren
Brem, Gottfried
Brown, Dorcas
Burger, Joachim
Crubézy, Eric
Daugnora, Linas
Davoudi, Hossein
de Barros Damgaard, Peter
de los Ángeles de Chorro y de Villa-Ceballos, María
Deschler-Erb, Sabine
Detry, Cleia
Dill, Nadine
do Mar Oom, Maria
Dohr, Anna
Ellingvåg, Sturla
Erdenebaatar, Diimaajav
Fathi, Homa
Felkel, Sabine
Fernández-Rodríguez, Carlos
García-Viñas, Esteban
Germonpré, Mietje
Granado, José D.
Hallsson, Jón H.
Hemmer, Helmut
Hofreiter, Michael
Kasparov, Aleksei
Khasanov, Mutalib
Khazaeli, Roya
Kosintsev, Pavel
Kristiansen, Kristian
Kubatbek, Tabaldiev
Kuderna, Lukas
Kuznetsov, Pavel
Laleh, Haeedeh
Leonard, Jennifer A.
Lhuillier, Johanna
Liesau von Lettow-Vorbeck, Corina
Logvin, Andrey
Lõugas, Lembi
Ludwig, Arne
Luis, Cristina
Margarida Arruda, Ana
Marques-Bonet, Tomas
Matoso Silva, Raquel
Merz, Victor
Mijiddorj, Enkhbayar
Miller, Bryan K.
Monchalov, Oleg
Mohaseb, Fatemeh A.
Morales, Arturo
Nieto-Espinet, Ariadna
Nistelberger, Heidi
Onar, Vedat
Pálsdóttir, Albína H.
Pitulko, Vladimir
Pitskhelauri, Konstantin
Pruvost, Mélanie
Rajic Sikanjic, Petra
Rapan Papeša, Anita
Roslyakova, Natalia
Sardari, Alireza
Sauer, Eberhard
Schafberg, Renate
Scheu, Amelie
Schibler, Jörg
Schlumbaum, Angela
Serrand, Nathalie
Serres-Armero, Aitor
Shapiro, Beth
Sheikhi Seno, Shiva
Shevnina, Irina
Shidrang, Sonia
Southon, John
Star, Bastiaan
Sykes, Naomi
Taheri, Kamal
Taylor, William
Teegen, Wolf-Rüdiger
dc.subject.por.fl_str_mv Animals
Asia
Biological Evolution
Breeding
DNA, Ancient
Domestication
Equidae
Europe
Female
Genetic variation
Genome
History, Ancient
Horses
Male
Phylogeny
topic Animals
Asia
Biological Evolution
Breeding
DNA, Ancient
Domestication
Equidae
Europe
Female
Genetic variation
Genome
History, Ancient
Horses
Male
Phylogeny
description Horse domestication revolutionized warfare and accelerated travel, trade, and the geographic expansion of languages. Here, we present the largest DNA time series for a non-human organism to date, including genome-scale data from 149 ancient animals and 129 ancient genomes (≥1-fold coverage), 87 of which are new. This extensive dataset allows us to assess the modern legacy of past equestrian civilizations. We find that two extinct horse lineages existed during early domestication, one at the far western (Iberia) and the other at the far eastern range (Siberia) of Eurasia. None of these contributed significantly to modern diversity. We show that the influence of Persian-related horse lineages increased following the Islamic conquests in Europe and Asia. Multiple alleles associated with elite-racing, including at the MSTN "speed gene," only rose in popularity within the last millennium. Finally, the development of modern breeding impacted genetic diversity more dramatically than the previous millennia of human management.
publishDate 2019
dc.date.none.fl_str_mv 2019
2019-01-01T00:00:00Z
2020-04-21T08:49:37Z
dc.type.status.fl_str_mv info:eu-repo/semantics/publishedVersion
dc.type.driver.fl_str_mv info:eu-repo/semantics/article
format article
status_str publishedVersion
dc.identifier.uri.fl_str_mv http://hdl.handle.net/10451/42994
url http://hdl.handle.net/10451/42994
dc.language.iso.fl_str_mv eng
language eng
dc.relation.none.fl_str_mv Fages, A., Hanghoj, K., Khan, N., Gaunitz, C., Seguin-Orlando, A., Leonardi, M., . . . Orlando, L. (2019). Tracking Five Millennia of Horse Management with Extensive Ancient Genome Time Series. Cell, 177(6) 1419-1435 e1431. doi: 10.1016/j.cell.2019.03.049
1097-4172
10.1016/j.cell.2019.03.049
dc.rights.driver.fl_str_mv metadata only access
info:eu-repo/semantics/openAccess
rights_invalid_str_mv metadata only access
eu_rights_str_mv openAccess
dc.format.none.fl_str_mv application/pdf
dc.publisher.none.fl_str_mv Elsevier
publisher.none.fl_str_mv Elsevier
dc.source.none.fl_str_mv reponame:Repositório Científico de Acesso Aberto de Portugal (Repositórios Cientìficos)
instname:Agência para a Sociedade do Conhecimento (UMIC) - FCT - Sociedade da Informação
instacron:RCAAP
instname_str Agência para a Sociedade do Conhecimento (UMIC) - FCT - Sociedade da Informação
instacron_str RCAAP
institution RCAAP
reponame_str Repositório Científico de Acesso Aberto de Portugal (Repositórios Cientìficos)
collection Repositório Científico de Acesso Aberto de Portugal (Repositórios Cientìficos)
repository.name.fl_str_mv Repositório Científico de Acesso Aberto de Portugal (Repositórios Cientìficos) - Agência para a Sociedade do Conhecimento (UMIC) - FCT - Sociedade da Informação
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