DÉPARTEMENT DE GÉOSCIENCES – SCIENCES DE LA TERRE UNIVERSITÉ DE FRIBOURG (SUISSE) The RuminanTia (mammalia, CeTaRTiodaCTyla) fRom The oligoCene To The eaRly mioCene of WesTeRn euRope: sysTemaTiCs, palaeoeCology and palaeobiogeogRaphy THÈSE présentée à la Faculté des Sciences de l’Université de Fribourg (Suisse) pour l’obtention du grade Doctor rerum naturalium Bastien menneCaRT de Epinay sur Seine, France Thèse N° 1756 Multiprint SA, Fribourg, 2012 Acceptée par la faculté des Sciences de l’Université de Fribourg (Suisse) sur la proposition de : Prof. Jean-Pierre BERGER Université de Fribourg (Fribourg, Suisse) Directeur Prof. André STRASSER Université de Fribourg (Fribourg, Suisse) Directeur Dr. Damien BECKER Paléontologie A16 (Porrentruy, Suisse) Expert Dr. Gertrud RÖSSNER Bavarian State Collections of Palaeontologie and Geology (Munich, Germany) Expert Dr. Grégoire METAIS Museum National d’Histoire Naturelle (Paris, France) Expert Prof. Vincent SERNEELS Université de Fribourg (Fribourg, Suisse) Président de Jury Fribourg, le 1 Juin 2012 Le Doyen: Prof. Rolf INGOLD Directeur de thèse: Prof. Jean-Pierre BERGER (décédé) Prof. André STRASSER Cover pictures left : Litocranius walleri (with the courtesy of Jill Mason) right : Hydropotes inermis (Bastien Mennecart) To Jean-Pierre tAble of contents AbstrAct ...............................................................I ..4.4.2 Taxonomical affinities..................................51 . 4.5.Relative.Warp.Analysis......................................52 Résumé ...............................................................III 4.6.Discussion.......................................................... 54 4.6.1 Biostratigraphy............................................54 Acknowledgements........................................................V ..4.6.2 Phylogenetic implications........................... 55 ..4.6.3 Palaeodiet....................................................56 I.– IntroductIon . 4.7.Conclusions........................................................56 . Acknowledgments................................................... 57 1 – GenerAl IntroductIon . References................................................................57.. 1.1.Aims.of.the.thesis................................................ 3.. 5 - eArly olIGocene europeAn ruMInAntIA (MAMMAlIA): 1.2.Organization.of.the.Dissertation......................... 7 coMMunItIes And evolutIon . References..................................................................8 . 5.1.Introduction........................................................61 2 - MAndIble shApe of ruMInAnts: between phyloGeny And . 5.2.Material.and.methods.........................................62 feedInG hAbIt . 5.3.Systematic.palaeontology.................................. 62 ..5.3.1 “Lophiomeryx” nov. sp............................... 67 Abstract. .................................................................11 5.3.2 “Lophiomeryx” minor.................................68 . 2.1.Introduction........................................................ 11 5.3.3 “Lophiomeryx” gaudryi..............................70 . 2.2.Material.and.Methods........................................14 5.3.4 Lophiomeryx mouchelini.............................71 ..2.2.1 Sampling......................................................14 5.3.5 Lophiomeryx chalaniati...............................76 ..2.2.2 Principal Component Analysis....................14 5.3.6 Iberomeryx minor........................................85 ..2.2.3 Abbreviations...............................................16 5.3.7 Bachitherium curtum...................................90 . 2.3.Results................................................................16 5.3.8 Bachitherium insigne...................................97 . 2.4.Discussion..........................................................18 5.3.9 Bachitherium vireti......................................103 ..2.4.1 Phylogeny................................................... 18 5.3.10 Bachitherium guirountensis...................... 103 2.4.2 Feeding Habit..............................................21 5.3.11 Bachitherium lavocati............................... 103 . 2.5.Conclusion......................................................... 22 5.3.12 Gelocus communis.....................................104 . Acknowledgments................................................... 22 5.3.13 Gelocus laubei...........................................108 . References................................................................22 5.3.14 Gelocus villebramarensis..........................108 5.3.14 Pseudogelocus scotti................................. 109 3 – MAterIAl And Methods . 5.4.Discussion..........................................................109 . 5.4.1 Lophiomerycidae.........................................109 . 3.1.Material..............................................................25 5.4.2 Tragulidae................................................... 110 . 3.2.Locality.distribution...........................................25 5.4.3 Bachitheriidae............................................. 111 ..3.2.1 Oligocene.................................................... 25 5.4.4 Gelocidae.................................................... 112 ..3.2.2 Early Miocene............................................. 32 5.4.5 Biostratigraphy and migration/extinction . 3.3.Stratigraphic.correlations...................................32 events. ................................................................ 112 . 3.4.Systematics........................................................ 33 . 5.5.Conclusions........................................................ 114 ..3.4.1.Systematic palaeontology............................33 . Bibliography............................................................ 114 ..3.4.2.Synonymy abbreviations..............................33 ..3.4.3.Anatomical abbreviations............................33 6 - mosaIcomeryx quercyI, new coMb. ruMInAnt froM the olIGocene . 3.5.Specific.methods................................................33 of europe, And the sIGnIfIcAnce of the “GelocIds” ..3.5.1 Geometric morphometrics...........................34 . 3.5.2 Ecomorphology........................................... 34 Abstract. ................................................................ 119 ..3.5.3 Cenogram....................................................34 . 6.1.Introduction........................................................ 119 . References................................................................34 . 6.2.Material,.methods,.and.abbreviations................120 . ..6.2.1 Systematic palaeontology............................120 II.– eArly And MId-olIGocene . 6.2.2 Abbreviations...............................................120 ..6.2.3 Institutional abbreviations.......................... 120 4 - Iberomeryx mInor (MAMMAlIA, ArtIodActylA) froM the eArly . 6.3.Systematic.palaeontology.................................. 121 olIGocene of soulce (cAnton JurA, nw swItzerlAnd): . 6.3.1 Mosaicomeryx quercyi................................ 121 systeMAtIcs And pAlAeodIet ..6.3.2 Prodremotherium elongatum.......................127 . 6.4.Cladistic.analysis............................................... 130 Abstract. ................................................................41 6.5.Discussion.......................................................... 131 . 4.1.Introduction........................................................42 6.5.1 Phylogeny....................................................131 . 4.2.Geological.setting.and.taphonomy.................... 43 . 6.5.2 Biostratigraphy............................................134 . 4.3.Material,.methods,.and.abbreviations................44 . 6.5.3 Evolution of the family................................ 135 ..4.3.1 Systematic palaeontology............................ 44 . 6.7.Conclusion......................................................... 135 ..4.3.2 Relative Warp Analysis................................44 . Acknowledgments................................................... 135 . 4.3.3 Abbreviations...............................................46 . References................................................................136 . 4.4.Systematic.palaeontology.................................. 46 . Annexe1. ................................................................139 ..4.4.1 Description..................................................47 III.– lAtest olIGocene . 10.1.Introduction......................................................208 . 10.2.Geological.setting.and.taphonomy.................. 208 7 - lArGe MAMMAls froM rIckenbAch (swItzerlAnd, reference . 10.3.Material.and.Methods......................................210 locAlIty Mp29, lAte olIGocene). bIostrAtIGrAphIc And . 10.4.Systematic.Palaeontology................................210 pAlAeoenvIronMentAl IMplIcAtIons ..10.4.1 Historic level............................................. 210 . .10.4.1.1.Diaceratherium.lemanense................................210 Abstract. ................................................................143 .. 10.4.1.2.Diaceratherium.aginense..................................213 . 7.1.Introduction........................................................143 .. 10.4.1.3.Diaceratherium.spp...........................................214 . 7.2.Historical.Background.......................................144 .. 10.4.1.4.Andegameryx.cf..laugnacensis.........................216 . 7.3.Material.and.Methods........................................146