Actinopterygii: Neopterygii) Et De Certains Taxons Fossiles Halécomorphes Basaux : Morphologie, Taxonomie Et Relations Phylogénétiques

THÈSE

Pour l'obtention du grade de DOCTEUR DE L'UNIVERSITÉ DE POITIERS UFR des sciences fondamentales et appliquées Institut de paléoprimatologie, paléontologie humaine : évolution et paléoenvironnements - IPHEP (Poitiers) (Diplôme National - Arrêté du 7 août 2006)

École doctorale : Sciences pour l'environnement - Gay Lussac (La Rochelle) Secteur de recherche : Terre solide et enveloppes superficielles

Présentée par : Giselle Ribeiro de Paula Machado

Révision des Ionoscopiformes (Actinopterygii: Neopterygii) et de certains taxons fossiles halécomorphes basaux : morphologie, taxonomie et relations phylogénétiques

Directeur(s) de Thèse : Olga Otero, Paulo Brito

Soutenue le 25 mars 2016 devant le jury

Jury :

Président Didier Bouchon Professeur des Universités, Université de Poitiers

Rapporteur François Meunier Professeur émérite, Musem national d'histoire naturelle

Rapporteur Gaël Clément Professeur des Universités, Museum national d'histoire naturelle

Membre Olga Otero Maître de conférences, Université de Poitiers

Membre Paulo Brito Professor, Rio de Janeiro universitário federal, Brasil

Membre Lionel Cavin Conservateur, Muséum d'histoire naturelle de Genève, Suisse

Membre Diogo de Mayrinck Professor assistente, Rio de Janeiro universitário federal, Brasil

Pour citer cette thèse : Giselle Ribeiro de Paula Machado. Révision des Ionoscopiformes (Actinopterygii: Neopterygii) et de certains taxons fossiles halécomorphes basaux : morphologie, taxonomie et relations phylogénétiques [En ligne]. Thèse Terre solide et enveloppes superficielles. Poitiers : Université de Poitiers, 2016. Disponible sur l'Intranet de l'Université de Poitiers 8QLYHUVLWpGH3RLWLHUV )DFXOWpGHV6FLHQFHV)RQGDPHQWDOHVHW$SSOLTXpHV

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Ophiopsis attenuata

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Ophiopsis penicillata

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Ophiopsis breviceps

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7DEHOD 'DGRV FRPSDUDWLYRV GRV HVSpFLPHV PHOKRU SUHVHUYDGRV GH 2SKLRSVLV 9HU OLVWDGHDEUHYLDWXUDVQDSiJ&RQWLQXDQDSUy[LPDSiJLQD (VSpFLHQRPLQDO /RFDOLGDGH &7 &3 &3$ &$ &33 &3Y &3' LGHQWLILFDomRQDHWLTXHWD Ophiopsis procera 1+08.393 $OHPDQKD.HOKHLQ " " " " " " " 1+08.393 $OHPDQKD.HOKHLQ " " " " " 1+08.393 $OHPDQKD.HOKHLQ " " " " " " " 1+08.393 $OHPDQKD(LFKVWlWW " 1+08.393 $OHPDQKD.HOKHLQ 1+08.393 $OHPDQKD(LFKVWlWW " " %63*$69,,, $OHPDQKD.HOKHLQ " " " %63*$6;,; $OHPDQKD.HOKHLQ -0(626 $OHPDQKD.HOKHLQ -0(626 $OHPDQKD.HOKHLQ " " -0(626 $OHPDQKD.HOKHLQ " " " " " " " -0(626 $OHPDQKD.HOKHLQ " " " -0(626 $OHPDQKD.HOKHLQ " " -0(626 $OHPDQKD.HOKHLQ " Ophiopsis attenuata 1+08.393 $OHPDQKD.HOKHLQ " " 1+08.393 )UDQoD&HULQ 1+08.393 $OHPDQKD.HOKHLQ " " 1+08.393 $OHPDQKD(LFKVWlWW " " " 0+1/ )UDQoD&HULQ " " " " " " " 0+1/ )UDQoD&HULQ " " " 0+1/ )UDQoD&HULQ " 0+1/ )UDQoD&HULQ " 0+1/ )UDQoD&HULQ " 0+1/ )UDQoD&HULQ 0+1/ )UDQoD&HULQ " " " " " " 0+1/ )UDQoD&HULQ " " 0+1/ )UDQoD&HULQ " " 0+1/ )UDQoD&HULQ " " 0+1/ )UDQoD&HULQ " " " " " " " 0+1/ )UDQoD&HULQ 0+1/ )UDQoD&HULQ " " " " " " " %63*;;,,,DHE $OHPDQKD.HOKHLQ " " " " " " %63*;;;;DHE $OHPDQKD.HOKHLQ " " -0(626E $OHPDQKD.HOKHLQ " " " -0(626 $OHPDQKD.HOKHLQ " " -0(626 $OHPDQKD.HOKHLQ " OphiopsisVSRXQmR LGHQWLILFDGRV %63*9 $OHPDQKD.HOKHLQ (77 $OHPDQKD(WWOLQJ " " " " " " " (77 $OHPDQKD(WWOLQJ " " (77 $OHPDQKD(WWOLQJ Ophiopsis penicillata 1+08.393 ,QJODWHUUD'RUVHW Ophiopsis breviceps 1+08.393 ,QJODWHUUD7LVEXU\ " 1+08.393 ,QJODWHUUD7LVEXU\ " " Ophiopsis dorsalis 1+08.393 ,QJODWHUUD'RUVHW 1+08.393 ,QJODWHUUD'RUVHW " " Ophiopsis montsechensis 01+106(D (VSDQKD0RQWVHF "

7DEHOD'DGRVFRPSDUDWLYRVGRVHVSpFLPHVPHOKRUSUHVHUYDGRVGH2SKLRSVLV9HU OLVWDGHDEUHYLDWXUDVQDSiJ&RQWLQXDQDSUy[LPDSiJLQD (VSpFLHQRPLQDO &'&3 &' && &&&7 &&&3 $& && $0& LGHQWLILFDomRQDHWLTXHWD Ophiopsis procera 1+08.393 " " " " " " " 1+08.393 " " " " " " 1+08.393 " " " " " " 1+08.393 1+08.393 1+08.393 %63*$69,,, " %63*$6;,; " " " " " " -0(626 -0(626 -0(626 " " " " " " " " -0(626 -0(626 -0(626 Ophiopsis attenuata 1+08.393 " " " 1+08.393 " " 1+08.393 1+08.393 " " 0+1/ " " " " " " " 0+1/ " " 0+1/ 0+1/ 0+1/ 0+1/ 0+1/ " " " " " " 0+1/ 0+1/ 0+1/ " " 0+1/ " " " " " " " " 0+1/ 0+1/ " " " " " " " " %63*;;,,,DHE " " " " " " " " %63*;;;;DHE " -0(626E -0(626 -0(626 OphiopsisVSRXQmR LGHQWLILFDGRV %63*9 (77 " " " " " " (77 (77 Ophiopsis penicillata 1+08.393 Ophiopsis breviceps 1+08.393 1+08.393 " " " Ophiopsis dorsalis 1+08.393 1+08.393 Ophiopsis montsechensis 01+106(D "

7DEHOD'DGRVFRPSDUDWLYRVGRVHVSpFLPHVPHOKRUSUHVHUYDGRVGH2SKLRSVLV9HU OLVWDGHDEUHYLDWXUDVQDSiJ&RQWLQXDQDSUy[LPDSiJLQD (VSpFLHQRPLQDO 1RGHHVFDPDVDR 1RGHHVFDPDVQD 1RGHHVFDPDVQR LGHQWLILFDomRQDHWLTXHWD ORQJRGDOLQKD SDUWHPDLVDOWDGR SHG~QFXORFDXGDO ODWHUDO FRUSR Ophiopsis procera 1+08.393 " " 1+08.393 " 1+08.393 " DSUR[ 1+08.393 1+08.393 DSUR[ 1+08.393 " " " %63*$69,,, DSUR[ " %63*$6;,; -0(626 -0(626 -0(626 " " " -0(626 " " -0(626 RX -0(626 Ophiopsis attenuata 1+08.393 " 1+08.393 1+08.393 DSUR[ DSUR[ 1+08.393 " " 0+1/ " " 0+1/ " " 0+1/ j 0+1/ 0+1/ 0+1/ 0+1/ " " " 0+1/ DSUR[ 0+1/ PDLVTXH DSUR[ 0+1/ " 0+1/ " " " 0+1/ " " " 0+1/ " " " %63*;;,,,DHE " " " %63*;;;;DHE DRPHQRV DSUR[ -0(626E -0(626 -0(626 OphiopsisVSRXQmR LGHQWLILFDGRV %63*9 " (77 (77 (77 Ophiopsis penicillata 1+08.393 DRPHQRV DSUR[ Ophiopsis breviceps 1+08.393 " " " 1+08.393 " " " Ophiopsis dorsalis 1+08.393 " " " 1+08.393 " " " Ophiopsis montsechensis 01+106(D "

7DEHOD'DGRVFRPSDUDWLYRVGRVHVSpFLPHVPHOKRUSUHVHUYDGRVGH2SKLRSVLV9HU OLVWDGHDEUHYLDWXUDVQDSiJ&RQWLQXDQDSUy[LPDSiJLQD (VSpFLHQRPLQDO 1RGHHOHPHQWRVGD 1RGHUDLRVGD )RUPDGDGRUVDO LGHQWLILFDomRQDHWLTXHWD QDGDGHLUDGRUVDO QDGDGHLUDDQDO Ophiopsis procera 1+08.393 " " " 1+08.393 EHPIDOFDGD USUSU " 1+08.393 " USUSU " 1+08.393 EHPIDOFDGD USUSU " 1+08.393 EHPIDOFDGD USRXUSU DRPHQRV 1+08.393 " FHUFDGH " %63*$69,,, " " %63*$6;,; EHPIDOFDGD " -0(626 EHPIDOFDGD -0(626 EHPIDOFDGD " -0(626 EHPIDOFDGD " " -0(626 EHPIDOFDGD USUSUIE " -0(626 EHPIDOFDGD " " -0(626 EHPIDOFDGD " " Ophiopsis attenuata 1+08.393 " " DRPHQRV 1+08.393 " " " 1+08.393 " USUSU " 1+08.393 " " " 0+1/ " " " 0+1/ " " " 0+1/ " " DRPHQRV 0+1/ " " " 0+1/ " " " 0+1/ EHPIDOFDGD USUSU " 0+1/ " " " 0+1/ " " 0+1/ " " " 0+1/ " DRPHQRV " 0+1/ " " " 0+1/ " " " 0+1/ " " " %63*;;,,,DHE " " " %63*;;;;DHE " " " -0(626E SRXFRIDOFDGD USUSU " -0(626 PHGLDQDPHQWHIDOFDGD " -0(626 SRXFRIDOFDGD " OphiopsisVSRXQmR LGHQWLILFDGRV %63*9 " FHUFDGH " (77 SRXFRIDOFDGD (77 SRXFRIDOFDGD USUSUIE " (77 SRXFRIDOFDGD USUSUIE UWIE Ophiopsis penicillata 1+08.393 SRXFRIDOFDGD USRXUSU " Ophiopsis breviceps 1+08.393 " " " 1+08.393 " DRPHQRV " Ophiopsis dorsalis 1+08.393 SRXFRIDOFDGDjUHWD FHUFDGH RXUS 1+08.393 " " " Ophiopsis montsechensis 01+106(D SRXFRIDOFDGDj USUSU UW UHWDGHFUHVFHQWH

7DEHOD'DGRVFRPSDUDWLYRVGRVHVSpFLPHVPHOKRUSUHVHUYDGRVGH2SKLRSVLV9HU OLVWDGHDEUHYLDWXUDVQDSiJ&RQWLQXDQDSUy[LPDSiJLQD (VSpFLHQRPLQDO 1RGHUDLRVSULQFLSDLVGDQDGDGHLUD 1RGHUDLRVGDQDGDGHLUD LGHQWLILFDomRQDHWLTXHWD FDXGDO SHLWRUDO Ophiopsis procera 1+08.393 " " 1+08.393 DSUR[ " 1+08.393 " SUHVHUYDGRV 1+08.393 DSUR[WRWDLV USU 1+08.393 i 1+08.393 " %63*$69,,, " " %63*$6;,; " -0(626 -0(626 DSUR[ -0(626 " " -0(626 -0(626 " -0(626 " Ophiopsis attenuata 1+08.393 " 1+08.393 1+08.393 YLVtYHLV 1+08.393 " " 0+1/ " 0+1/ " 0+1/ DSUR[ 0+1/ 0+1/ 0+1/ 0+1/ " " 0+1/ YLVtYHLV 0+1/ " 0+1/ " " 0+1/ " " 0+1/ " " 0+1/ " " %63*;;,,,DHE " " %63*;;;;DHE " " -0(626E YLVtYHLV -0(626 DSUR[ DSUR[YLVtYHLVHPFDGD -0(626 YLVtYHLV OphiopsisVSRXQmR LGHQWLILFDGRV %63*9 " (77 (77 WRWDLV (77 USUSUIE Ophiopsis penicillata 1+08.393 FRQWiYHLV Ophiopsis breviceps 1+08.393 " " 1+08.393 " " Ophiopsis dorsalis 1+08.393 " " 1+08.393 " " Ophiopsis montsechensis 01+106(D " WRWDLV

7DEHOD'DGRVFRPSDUDWLYRVGRVHVSpFLPHVPHOKRUSUHVHUYDGRVGH2SKLRSVLV9HU OLVWDGHVLJODVHDEUHYLDWXUDVQDVSiJLQDVH

(VSpFLHQRPLQDO LGHQWLILFDomR R 1 GHUDLRVGDQDGDGHLUDSpOYLFD &DQDOQDQDGDGHLUDFDXGDO QDHWLTXHWD Ophiopsis procera 1+08.393 " VLP DEDL[RGRUDLR 1+08.393 QmRSUHVHUYDGD 1+08.393 " QmRSUHVHUYDGD 1+08.393 US VLP DEDL[RGRUDLR 1+08.393 VLP DEDL[RGRUDLR 1+08.393 " " %63*$69,,, " %63*$6;,; QmRYLVtYHO -0(626 YLVtYHLV VLP DEDL[RGRUDLR -0(626 YLVtYHLV VLP DEDL[RGRUDLR -0(626 " " -0(626 " " -0(626 " -0(626 IE " Ophiopsis attenuata 1+08.393 " 1+08.393 " VLP DEDL[RGRUDLR 1+08.393 " VLP DEDL[RGRUDLR 1+08.393 " " 0+1/ " VLP DEDL[RGRUDLR 0+1/ " " 0+1/ QmRYLVtYHO 0+1/ VLP DEDL[RGRUDLR 0+1/ " VLP DEDL[RGRUDLR 0+1/ " VLP DEDL[RGRUDLR 0+1/ " " 0+1/ YLVtYHLV QmRYLVtYHO 0+1/ " QmRYLVtYHO 0+1/ " 0+1/ " " 0+1/ " " 0+1/ " " %63*;;,,,DHE " " %63*;;;;DHE " " -0(626E " VLP DEDL[RGRUDLR -0(626 "YLVtYHLV " -0(626 YLVtYHLV " OphiopsisVSRXQmR LGHQWLILFDGRV %63*9 "SUHVHUYDGRV VLP DEDL[RGRUDLR (77 (77 US VLP DEDL[RGRUDLR (77 USUSU VLP DEDL[RGRUDLR Ophiopsis penicillata 1+08.393 " " Ophiopsis breviceps 1+08.393 " " 1+08.393 " " Ophiopsis dorsalis 1+08.393 " " 1+08.393 " " Ophiopsis montsechensis 01+106(D UW VLP

7DEHOD&RPSDUDomRGHDOJXPDVFDUDFWHUtVWLFDVGH$SKDQHS\JXVFRPRVJrQHURVGH 2SKLRSVLGDH7HRLFKWK\VH3ODFLGLFKWK\V

Aphanepygus Teoichthys Placidichthys VXSUDRUELWDLV RUJDQL]DGRVHPGXDV RUJDQL]DGRVHPXPD RUJDQL]DGRVHPXPDILOHLUD ILOHLUDV ILOHLUD GHUPRHVIHQyWLFR ID]SDUWHGDyUELWD FRPSOHWDPHQWHDQH[DGR FRPSOHWDPHQWHDQH[DGRDR DRWHWRFUDQLDQR WHWRFUDQLDQR DXWRHVIHQyWLFR DXVHQWH SRVLFLRQDGRQRFDQWR SRVLFLRQDGRQRFDQWR VXSHULRUGDyUELWD VXSHULRUGDyUELWD H[WUDHVFDSXODU IRUPDGRSRUSHTXHQDV XPSDUGHRVVRV XPSDUGHRVVRV SODFDVFDGDXPDFRP DSUR[LPDGDPHQWH DSUR[LPDGDPHQWH RUQDPHQWDomRVHUULOKDGD WULDQJXODUHVSRUWDQGRD WULDQJXODUHVFRP QDERUGDSRVWHULRU FRPLVVXUD RUQDPHQWDomRVHUULOKDGDQD FRPLVVXUDVXSUDWHPSRUDO VXSUDWHPSRUDO ERUGDSRVWHULRUHSRUWDQGRD SUHVHQWH FRPLVVXUDVXSUDWHPSRUDO SyVWHPSRUDO XPSDUUHGX]LGRV XPSDUDPSORV XPSDUDPSORV PD[LOD VHPGHQWH FRPGHQWH FRPGHQWH VXSUDPD[LOD DXVHQWH SUHVHQWH SUHVHQWH DUWLFXODomR " GXSOD GXSOD PDQGLEXODU VXERUELWDLV UHJLmRUHFREHUWDSRU RUJDQL]DGRVHP RUJDQL]DGRVHPVpULH HVTXDPXODV PRVDLFR SODFDJXODU DXVHQWH SUHVHQWH SUHVHQWH SUpRSpUFXOR HPSODFDV SHoD~QLFDDORQJDGDH SHoD~QLFDDORQJDGDHVWUHLWD FUHVFHQWH HFUHVFHQWH QDGDGHLUDGRUVDO ORQJDGLYLGLGDRXQmR ORQJDRXFXUWD ORQJDHGLYLGLGD QDGDGHLUD ERUGDSRVWHULRUFRQYH[D ERUGDSRVWHULRU ERUGDSRVWHULRUELIXUFDGD FDXGDO ELIXUFDGD

7DEHOD&RPSDUDomRGHDOJXPDVFDUDFWHUtVWLFDVGH0DFURVHPLLGDHH2SKLRSVLGDH (VWUXWXUD 0DFURVHPLLGDH 2SKLRSVLGDH 6LPLODULGDGHV *DQRLGHVJHUDOPHQWHVHUULOKDGDVQD *DQRLGHVJHUDOPHQWHVHUULOKDGDVQDPDUJHP (VFDPDV PDUJHPSRVWHULRUIRUPDWRYDULiYHO SRVWHULRUIRUPDWRYDULiYHODUWLFXODomR DUWLFXODomRSHJ VRFNHW SHJ VRFNHW 1DGDGHLUDGRUVDO /RQJDHRXGLYLGLGD /RQJDHGLYLGLGDRXFXUWD 'LIHUHQoDV $UWLFXODomRVLPSOHV DSHQDVR 'XSODDUWLFXODomR TXDGUDGRHVLPSOpWLFR $UWLFXODomRPDQGLEXODU TXDGUDWRMXJDODUWLFXODFRPRRVVRDUWLFXODU DUWLFXODPFRPRRVVRDUWLFXODU 3RVLomRGDDUWLFXODomR $QWHULRUjyUELWD 3RVWHULRURXQDDOWXUDGDyUELWD PDQGLEXODU 6XSUDPD[LOD $XVHQWH 6LPSOHV $QHOFLUFXPRUELWDO ,QFRPSOHWR &RPSOHWR ,QIUDRUELWDLV 0DLVGHFLQFRHVWUHLWRV $WpTXDWURPDLVDOWRVTXHFRPSULGRV $UWLFXODGRFRPRWHWRFUDQLDQRPDVDLQGD 'HUPRHVIHQyWLFR &RPSOHWDPHQWHVXWXUDGRDRWHWRFUDQLDQR ID]SDUWHGDyUELWD $XWRHVIHQyWLFR SRUomR $XVHQWH HVIHQyWLFRQmRH[SRVWR 3RVLFLRQDGRQRFDQWRVXSHULRUGDyUELWD H[SRVWDGRHVIHQyWLFR 3UHVHQWHQDSRUomRSRVWHULRUGRSDULHWDO 3UHVHQWHQDOLQKDPHGLDQDGRVRVVRV &RPLVVXUD H[WUDHVFDSXODUHVUHGX]LGRVHH[FOXtGRVGD H[WUDHVFDSXODUHV HVWHVRVVRVQmRVmRUHGX]LGRV VXSUDWHPSRUDO OLQKDPpGLDGRWHWRFUDQLDQR HVHHQFRQWUDPQDOLQKDPpGLDGRWHWRFUDQLDQR 6XERUELWDLV $XVHQWHV 3UHVHQWHV )XOFUDVEDVDLVH 3UHVHQWHVDSHQDVQDQDGDGHLUDFDXGDO 3UHVHQWHVWDPEpPQDVQDGDGHLUDVPpGLDV PDUJLQDLV

7DEHOD5HVXPRGDVSULQFLSDLVYDULDo}HVHQWUHDVHVSpFLHVQRPLQDLVGH,RQRVFRSXV

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Ionoscopus 3LHWUDURMD,WiOLD RXPDLV DUUHGRQGDGD FRPHOHYDomRSUp pietraroiea GRUVDOEHP GHVHQYROYLGD

Ionoscopus desori 6ROQKRIHQ j j j DUUHGRQGDGD FRPHOHYDomRSUp $OHPDQKDH&HULQ GRUVDOEHPPHGLDQD )UDQoD

Ionoscopus 6ROQKRIHQ j RX j DUUHGRQGDGD FRPHOHYDomRSUp muensteri $OHPDQKD GRUVDOEHPPHGLDQD

Ionoscopus 6ROQKRIHQ DUUHGRQGDGD FRPHOHYDomRSUp striatissimus $OHPDQKD GRUVDOEHPPHGLDQD

Ionoscopus 6ROQKRIHQ RX RX DUUHGRQGDGD FRPHOHYDomRSUp analibrevis (nomen $OHPDQKD GRUVDOEHPPHGLDQD nudum)

Ionoscopus 6ROQKRIHQ RX RX DORQJDGDHP VHPHOHYDomRSUp cyprinoides $OHPDQKD IRUPDGHRYR GRUVDO

7DEHOD&RPSDUDomRGRQ~PHURGHHVFDPDVHQWUH+HWHUROHSLGRWXVODWXVHGRLVSHL[HV6HPLRQRWLIRUPHV

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Heterolepidotus latus

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Lepidotes notopterus &HUFDGH

Tlayuamichin itztli &HUFDGH

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2SKLRSVLGDH 2SKLRSVLVSURFHUD -0(626 -XUiVVLFR6XSHULRU6ROQKRIHQ$OHPDQKD 2SKLRSVLGDH 2SKLRSVLVSURFHUD -0(626 -XUiVVLFR6XSHULRU6ROQKRIHQ$OHPDQKD 2SKLRSVLGDH 2SKLRSVLVSURFHUD -0(626 -XUiVVLFR6XSHULRU6ROQKRIHQ$OHPDQKD 2SKLRSVLGDH 2SKLRSVLVSURFHUD -0(626 -XUiVVLFR6XSHULRU6ROQKRIHQ$OHPDQKD 2SKLRSVLGDH 2SKLRSVLVSURFHUD -0((77 -XUiVVLFR6XSHULRU6ROQKRIHQ$OHPDQKD 2SKLRSVLGDH 2SKLRSVLVDWHQXDWWD 2SKLRSVLVSURFHUD -0(626DHE -XUiVVLFR6XSHULRU6ROQKRIHQ$OHPDQKD 2SKLRSVLGDH 2SKLRSVLVDWHQXDWWD 2SKLRSVLVSURFHUD -0(626DHE -XUiVVLFR6XSHULRU6ROQKRIHQ$OHPDQKD 2SKLRSVLGDH 2SKLRSVLVDWHQXDWWD 2SKLRSVLVSURFHUD -0(626 -XUiVVLFR6XSHULRU6ROQKRIHQ$OHPDQKD 2SKLRSVLGDH 2SKLRSVLVDWHQXDWWD 2SKLRSVLVSURFHUD -0(626DHE -XUiVVLFR6XSHULRU6ROQKRIHQ$OHPDQKD 2SKLRSVLGDH 2SKLRSVLVDWHQXDWWD 2SKLRSVLVSURFHUD -0(626 -XUiVVLFR6XSHULRU6ROQKRIHQ$OHPDQKD 2SKLRSVLGDH 2SKLRSVLVDWHQXDWWD 2SKLRSVLVSURFHUD -0(626 -XUiVVLFR6XSHULRU6ROQKRIHQ$OHPDQKD 2SKLRSVLGDH 2SKLRSVLVDWHQXDWWD 2SKLRSVLVSURFHUD -0(626 -XUiVVLFR6XSHULRU6ROQKRIHQ$OHPDQKD 2SKLRSVLGDH 2SKLRSVLVVS 2SKLRSVLVSURFHUD -0((77 -XUiVVLFR6XSHULRU6ROQKRIHQ$OHPDQKD 2SKLRSVLGDH 2SKLRSVLVVS 2SKLRSVLVSURFHUD -0((77 -XUiVVLFR6XSHULRU6ROQKRIHQ$OHPDQKD 2SKLRSVLGDH 2SKLRSVLVVS 2SKLRSVLVSURFHUD -0((77 -XUiVVLFR6XSHULRU6ROQKRIHQ$OHPDQKD 2SKLRSVLGDH 2SKLRSVLVDWHQXDWWD 2SKLRSVLVSURFHUD 1+1/ .LPHULGLDQR,QIHULRU&HULQ)UDQoD 2SKLRSVLGDH 2SKLRSVLVDWHQXDWWD 2SKLRSVLVSURFHUD 1+1/ .LPHULGLDQR,QIHULRU&HULQ)UDQoD 2SKLRSVLGDH 2SKLRSVLVDWHQXDWWD 2SKLRSVLVSURFHUD 1+1/ .LPHULGLDQR,QIHULRU&HULQ)UDQoD 2SKLRSVLGDH 2SKLRSVLVDWHQXDWWD 2SKLRSVLVSURFHUD 1+1/ .LPHULGLDQR,QIHULRU&HULQ)UDQoD 2SKLRSVLGDH 2SKLRSVLVDWHQXDWWD 2SKLRSVLVSURFHUD 1+1/ .LPHULGLDQR,QIHULRU&HULQ)UDQoD 2SKLRSVLGDH 2SKLRSVLVDWHQXDWWD KROyWLSR 2SKLRSVLVSURFHUD 1+1/ .LPHULGLDQR,QIHULRU&HULQ)UDQoD 2SKLRSVLGDH 2SKLRSVLVDWHQXDWWD 2SKLRSVLVSURFHUD 1+1/ .LPHULGLDQR,QIHULRU&HULQ)UDQoD 2SKLRSVLGDH 2SKLRSVLVDWHQXDWWD 2SKLRSVLVSURFHUD 1+1/ .LPHULGLDQR,QIHULRU&HULQ)UDQoD 2SKLRSVLGDH 2SKLRSVLVDWHQXDWWD 2SKLRSVLVSURFHUD 1+1/ .LPHULGLDQR,QIHULRU&HULQ)UDQoD 2SKLRSVLGDH 2SKLRSVLVDWHQXDWWD 2SKLRSVLVSURFHUD 1+1/ .LPHULGLDQR,QIHULRU&HULQ)UDQoD 2SKLRSVLGDH 2SKLRSVLVDWHQXDWWD 2SKLRSVLVSURFHUD 1+1/ .LPHULGLDQR,QIHULRU&HULQ)UDQoD 2SKLRSVLGDH 2SKLRSVLVDWHQXDWWD 2SKLRSVLVSURFHUD 1+1/ .LPHULGLDQR,QIHULRU&HULQ)UDQoD 2SKLRSVLGDH 2SKLRSVLVDWHQXDWWD 2SKLRSVLVSURFHUD 1+1/ .LPHULGLDQR,QIHULRU&HULQ)UDQoD 2SKLRSVLGDH 2SKLRSVLVSURFHUD 1+08.393 -XUiVVLFR6XSHULRU6ROQKRIHQ$OHPDQKD

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Crâne d’Ionoscopus desori (M HNL 20015.090) en lumière UV.

Cette illustration (Fig. 13 de la thèse) de l’intérêt de la lumière UV pour observer les détails de l’anatomie de certains fossiles rend compte aussi du travail d’anatomie effectué. Légende : ang, angulaire; ao, antorbital; ar, articulaire; asp, autosphénotique; bpt, processus basiptérigoïde; cdsp, canal dermosphénotique; ch, céra- tohyal; de, dentaire; dpt, dermo- ptérotique; dsp, dermosphé- notique; ext, extrascapulaire; fr, frontal; g, plaque gulaire; h, hyomandibulaire; io, infraorbital; mx, maxillaire; na, nasal; pa, pariétal; pas, parasphénoïde; pmx, prémaxillaire; pop, préopercule; q, carré; ro, rostral; sang, supra- angulaire; smx, supra-maxillaire; su, supraorbital; suo, suborbital; sym, symplectique. Echelle = 20mm. Photo : G. M achado.

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Crâne de Heterolepidotus rhombifer (NHM UKPVP 2009) en lumière UV. Cette reconstruction précise l’anatomie crânienne d’une des rares espèces considérées valides d’Heterolepidotus (Fig. 25 de la thèse). Légende : ang, angulaire; ar, articulaire; asp, autosphénotique; br, rayons bran- chiostèges; ch, cératohial; cl, cleithrum; de, dentaire; dpt, dermoptérotique; dsp, dermosphénotique; ext, extrascapulaire; fr, frontal; h, hyomandibulaire; io, infraorbital; iop, interoperculaire; l, lacrymal; le, ethmoide latéral; mx, maxillaire; na, nasal; op, operculaire; pa, pariétal; pcl, post- cleithrum et plaques post-cleithrales; pop, pré-operculaire; pt, post-temporal; q, carré; ro, rostral; sang, supra-angulair; scl, supracleithrum; smx, supramaxillaire; sop, sous-opercule; suo, sous-orbital. Echelle = 20mm. Photo : G. M achado.

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Légende : A, photographie du crâne d’A. dorsalis (NHM UKPVOR 103932) (sob luz UV). B, dessin du crâne d’A. dorsalis (NHM UKPVOR 103932) (modifié d’après Bartram, 1977b). C, dessin du crâne de Placidichthys bidorsallis (UERJ-PM B 475). D, dessin du crâne de Teoichthys brevipina (IGM 6744). asp, autosphénotique; dsp, dermosphénotique; esq, écailles; ext, extrascapulaire; m.suo, mosaïque de plaques susorbitaires; mx, maxillaire; pop, préopercule; pt, post-temporal; smx, supramaxillaire; su, supraorbital; suo, suborbital. Echelle = 5mm. Photo et dessins de G. M achado. &RPSDUDLVRQGHPlacidichthys DYHFGHV2SKLRSVLGDH HWGHV0DFURVHPLLGDH /HSODFHPHQWDOWHUQDWLIGH3ODFLGLFKWK\VGDQVFHVGHX[IDPLOOHVVXLYDQWOHVDXWHXUVDDSSHOp FHWWHFRPSDUDLVRQ'HIDLWOHJHQUHSUpVHQWHGHVFDUDFWqUHVUHQFRQWUpVFKH]GHVPHPEUHV GHVGHX[IDPLOOHV HWG¶DXWUHVWD[RQVQpRSWpU\JLHQV PDLVDXVVLGL[FDUDFWqUHVH[FOXVLIVGHV 2SKLRSVLGDH FRPPH SDU H[HPSOH OD GRXEOH DUWLFXODWLRQ GH OD PDQGLEXOH HW XQ DQQHDX FLUFXPRUELWDOFRPSOHW $QDO\VHVFODGLVWLTXHVGHVUHODWLRQVSK\ORJpQpWLTXHV 8QHSUHPLqUHDQDO\VHLQFOXDQWWRXVOHVWD[RQVDSURGXLWDUEUHVSDUFLPRQLHX[GHSDV /H FRQVHQVXV VWULFW ODLVVH OHV 7HOHRVWHL $PLLIRUPHV ,RQRVFRSLGDH 2SKLRSVLGDH H *LQJO\PRGLHQSRO\WRPLHHWVHXOVOHVFODGHV/HSLVRVWHLIRUPHV $WUDFWRVWHXV/HSLVRVWHXV ,RQRVFRSLGDH 2VKXQLD 4XHW]DOLFKWK\V ,RQRVFRSXV HW 7HOHRVWHL (ORSV 3KROLGRSKRUXV VRQWUHWURXYpV (QH[FOXDQW3RO\SWHUXVRUQDWLSLQQLVDUEUHVSDUFLPRQLHX[GHSDVVRQWWURXYpV HWHQUDFLQpVVXUOHVWpOpRVWpHQV 3KROLGRSKRUXV /HV$PLLGDH*LQJO\PRGLHW,RQRVFRSLGDH VRQW HQ SRO\WRPLH PDLV OHV WD[RQV /HSLVRVWHLIRUPHV $WUDFWRVWHXV /HSLVRVWHXV HW ,RQRVFRSLGDH 2VKXQLD4XHW]DOLFKWK\V,RQRVFRSXV VRQWUpVROXV $YHF O¶H[FOXVLRQ G¶2VWHRUDFKLV HW 1HRUKRPEROHSLV TXDWUH DUEUHV SDUFLPRQLHX[ GH SDVVRQWUHWURXYpVDYHF717HWDYHF3$83&HWWHGLIIpUHQFHHVWPDOFRPSULVHHW GRLWrWUHDQDO\VpHSOXVSUpFLVpPHQW1pDQPRLQVRQUHWURXYHV\VWpPDWLTXHPHQW D ODPRQRSK\OLHGHV2SKLRSVLGDHVRLWXQHUHODWLRQSURFKHHQWUH2SKLRSVLVHW3ODFLGLFKWK\V E XQFODGH2SKLRSVLGDH +HWHUROHSLGRWXV)XUR HW5REXVWLFKWK\VOXRSLQJHQVLVFRPPH WD[RQEDVDOjFHFODGH F OD PRQRSK\OLH GHV $PLLIRUPHV ,RQRVFRSLGDH G OH SODFHPHQW GHV 3DUDVHPLRQRWLIRUPHV j OD EDVH GHV +DOHFRPRUSKL H OHV *LQJO\PRGL PRQRSK\OpWLTXHV PDLV OHV UHODWLRQV LQWHUQHV QH VRQW SDV UpVROXHV I OD PRQRSK\OLH GHV +RORVWHL J OH SODFHPHQW G¶$SKDQHS\JXV j OD Arbres de consensus strict obtenus en analyse de parcimonie lorsque Neorhombolepis EDVHGHV1HRSWHU\JLL excelsus et Osteorachis macrocephalus sont exclus (31 espèces retenues). Fig. 40 de la thèse. &HWWH WRSRORJLH HVW JOREDOHPHQW UHWURXYpH ORUVTXH O¶RQ H[FOXW DXVVL 3RO\SWHUXV 3OXV JpQpUDOHPHQW O¶DQDO\VH SK\ORJpQpWLTXH GRLW rWUH DSSURIRQGLH QRWDPPHQW j WUDYHUV O¶LQFOXVLRQ GH QRXYHDX[ FDUDFWqUHV PDLV DXVVL HQ SRXVVDQW OD UpIOH[LRQ VXU O¶HIIHW GH O¶pFKDQWLOORQQDJH VXU OD WRSRORJLH GH O¶DUEUH HW VD UREXVWHVVH(QILQOHVFDXVHVGHV GLIIpUHQFHVHQWUHOHVDQDO\VHVGHSDUFLPRQLHHQWUH3$83HW717GRLYHQWrWUHFRPSULVHV 5pIOH[LRQVXUOD©YDOHXUªGHVFDUDFWqUHVDQDWRPLTXHV 8QGHVSULQFLSDX[FRQVWDWVFRQFHUQHVDQVGRXWHODYDULDELOLWpGHODPRUSKRORJLHVFDODLUHHW FHjGLIIpUHQWVQLYHDX[&HUWDLQVFDUDFWqUHVVRQWYDULDEOHVVXLYDQWODSRVLWLRQGHO¶pFDLOOHVXU OHFRUSVPDLVDXVVLYUDLVHPEODEOHPHQWDXFRXUVGHODYLHGHVLQGLYLGXV,OHQUpVXOWHTXHOD UHFRQQDLVVDQFHGHWD[RQVVXUGHVpFDLOOHVLVROpHVHVWIUDJLOHHWOHXUXWLOLVDWLRQSRXUO¶DQDO\VH SK\ORJpQpWLTXHSUREOpPDWLTXHWDQWTX¶XQHpWXGHDSSURIRQGLHGHFHWWHYDULDELOLWpQHSHUPHW SDV GH FRPSUHQGUH OHV GLIIpUHQWV FDUDFWqUHV GHV pFDLOOHV $ FHW pJDUG GHV pWXGHV RQWRJpQpWLTXHVHWKLVWRORJLTXHVSHUPHWWURQWSUREDEOHPHQWGHPLHX[OHVDSSUpKHQGHUHWGRQF GHPLHX[OHVH[SORLWHU

Ornementation de quelques os du toit crânien (Fig. 42 de la thèse).

Légende:

A, Placidichthys bidorsallis (M PSC-P 775). B, Ophiopsis procera (JME- ETT 3370, en lumière UV). Les flêches rouge indique les serriations sur le bord de l’extrascapulaire et le post-temporal. Echelle = 10mm.

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