"ULL &LA -US .AT (IST .%7 ,%/.4).)$)$3 #,!33 -!--!,)! /2$%2 ./4/5.'5,!4! &!-),9,%/.4).))$!% &2/- 4(% 3!,,!"%$3 /& "/,)6)! $%3%!$!. ,!4%/,)'/#%.% "RUCE * 3HOCKEY ! REVISED DIAGNOSIS OF THE ,EONTINIIDAE #LASS -AMMALIA /RDER.OTOUNGULATA IS PROVIDED AND LEONTINIIDS OF THE $ESEADAN 3OUTH !MERICAN ,AND -AMMAL hAGEv 3!,-! LATE /LIGOCENE ARE SUMMARIZED 4WO NEW SPECIES OF LEONTINIDS FROM THE $ESEADAN 3ALLA "EDS OF "OLIVIA ARE DESCRIBED AND PLACED WITHIN THE NEW GENUS !NAYATHERIUM THE SMALLER! EKECOA AND A MUCH LARGER! FORTIS ! PHYLOGENETIC ANALYSIS SUGGESTS THAT THESE SPECIES ARE CLOSELY RELATED TO THE DERIVED 0ATAGONIAN GENERA !NCYLOCOELUS AND #OLPODON WITH THE LOSS OF THE CANINE SERVING AS A PUTATIVE SYNAPOMORPHY UNITING THESE TAXA !LTHOUGH LEONTINIIDS ARE THE MOST FREQUENTLY ENCOUNTERED TAXA AT MANY OTHER $ESEADAN LOCALITIES THEY ARE EXCEEDINGLY RARE AT 3ALLA 4HIS SCARCITY THE SCARCITY OF OTHER UNGULATES WITH LOW CROWNED CHEEK TEETH AND THE HEAVY TOOTH WEAR OF A RELATIVELY YOUNG INDIVIDUAL OF! EKECOA SUGGEST THAT UNGULATES WITH LOW CROWNED TEETH WERE AT A SELECTIVE DISADVANTAGE AT 3ALLA +EY 7ORDS .OTOUNGULATA ,EONTINIIDAE 3ALLA $ESEADAN NEW TAXA ).42/$5#4)/. +AY ET AL RATHER THAN EARLY /LIGOCENE 4HE $ESEADAN LATE /LIGOCENE 3OUTH !MERICAN ,AND 3IMPSON OR MUCH OLDER #RETACEOUS PER!MEGHINO -AMMAL h!GEv 3!,-! HAS BEEN OF CONSIDERABLE 3ALLA RECORDS ABOUT SPECIES OF MAM INTEREST TO STUDENTS OF 3OUTH !MERICAN PALEONTOLOGY FOR MALS IN OVER FAMILIES FROM OR ORDERS OF MAMMALS OVER A CENTURY 4HIS INTEREST HAS BEEN STIMULATED BY THE 3HOCKEY A GREAT DIVERSITY OF $ESEADAN MAMMALS INCLUDING THE hIM ,EONTINIIDS WERE MEDIUM SIZED TO LARGE MIGRANTv TAXA RODENTS AND MONKEYS !MEGHINO NOTOUNGULATES KNOWN FROM THE -USTERSAN 3!,-! 'AUDRY ,OOMIS (OFFSTETTER "OND ,OPEZ EARLY /LIGOCENE SEE +AY ET AL 0ATTERSON 0ASCUAL (ISTORICALLY THE MASSIVE TO THE MIDDLE -IOCENE OF ,A 6ENTA #OLOMBIA 0YROTHERIUM AND LEONTINIIDS HAVE SERVED AS hGUIDE FOS 6ILLARROEL #OLWELL $ANIS 4HEIR GREATEST DI SILSv TO INDICATE $ESEADAN HORIZONS 4HESE LARGE MAM VERSITY WAS BY FAR IN THE $ESEADAN 3!,-! WHERE MALS WERE THE MOST CONSPICUOUS FAUNAL ELEMENTS FOR FIVE GENERA ARE HERE REGARDED AS VALID SEE ALSO 3IMPSON THE EARLY STUDENTS OF THE $ESEADAN !MEGHINO -C+ENNA "ELL !LTHOUGH LEONTINIIDS ARE 'AUDRY ,OOMIS 3IMPSON COMMON MEMBERS OF MANY $ESEADAN FAUNAS OF 3ALLA PRONOUNCED SÖL YA BY THE LOCAL !YMARAN )N 0ATAGONIA EG ,A &LECHA ;!MEGHINO 'AUDRY DIANS "OLIVIA CONTAINS $ESEADAN BEDS THAT HAVE RE = THE LOCALITY NOW KNOWN AS #ABEZA "LANCA CEIVED MUCH ATTENTION IN THE LAST YEARS 4HIS IS DUE ;!MEGHINO ,OOMIS = AND 3CARRITT 0OCKET LARGELY TO THE DISCOVERY OF THE PRIMATE "RANISELLA ;#HAFFEE = AND OF THE TROPICAL 4REMEMB£ &ORMA (OFFSTETTER THE EARLIEST KNOWN OF MONKEY IN 3OUTH TION OF "RAZIL 3ORIA !LVARENGA THEY ARE RE !MERICA AS WELL AS 3ALLAS DIVERSE FAUNA OF MARSUPIALS MARKABLY UNCOMMON AT 3ALLA 4HE TWO SPECIMENS DE RODENTS AND UNGULATES SEE FAUNAL LIST IN 3HOCKEY A SCRIBED BELOW HOWEVER ARE COMPLETE ENOUGH TO GREATLY &ORTUITOUSLY 3ALLA IS INTERBEDDED WITH SEVERAL LAYERS OF IMPROVE OUR UNDERSTANDING OF THE LEONTINIIDS OF 3ALLA VOLCANIC ASH WHICH HAVE HELPED TO FIRMLY ESTABLISH THE 0RIOR TO THIS WORK ONLY FRAGMENTARY REMAINS HAD BEEN $ESEADAN AS BEING LATE /LIGOCENE -AC&ADDEN ET AL NOTED IN PUBLISHED ACCOUNTS OF 3ALLA )N HIS FIRST PUBLICATION REGARDING 3ALLA (OFFSTETTER &LORIDA -USEUM OF .ATURAL (ISTORY AND $EPARTMENT OF :OOL MENTIONED THE PROBABLE PRESENCE OF LEONTINIIDS OGY 5NIVERSITY OF &LORIDA 'AINESVILLE &, THERE BUT NOTED THAT THE REMAINS WERE RARE AND TOO FRAG #%./:/)# 6%24%"2!4%3 0APERS TO (ONOR 3 $AVID 7EBB MENTARY TO RULE OUT THE POSSIBILITY THAT THEY WERE B !N hEXHAUSTIVE SEARCHv WAS EMPLOYED TO GEN HOMALODONTOTHERIIDS (OFFSTETTER NOTED THE AB ERATE HYPOTHETICAL PHYLOGENETIC TREES 4HE MOST PARSI SENCE OF HOMALODONTOTHERIIDS ON A REVISED FAUNAL LIST AND MONIOUS HYPOTHESIS IS REPORTED AND DISCUSSED GAVE(ENRICOFILHOLIA AS THE GENUS OF LEONTINIID PRESENT -AC&ADDEN ET AL DID NOT INCLUDE LEONTINIIDS ON 3934%-!4)# 0!,%/.4/,/'9 THEIR FAUNAL LIST BUT MENTIONED LEONTINIID GENERA IE /RDER ./4/5.'5,!4! 2OTH (ENRICOFILHOLIA ,EONTINIA AND !NCYLOCOELUS IN THEIR 3UBORDER 4/8/$/.4)! /WEN DISCUSSION OF ISOTEMNIDS SUGGESTING THAT THESE LEONTINIID &AMILY ,%/.4).))$!%!MEGHINO GENERA WERE ERRONEOUSLY CONSIDERED TO BE ISOTEMNIDS 2EVISED $IAGNOSIS"RACHYDONT TO MESODONT 3UBSEQUENT TO (OFFSTETTERS EXPEDITIONS TO 3ALLA 4OXODONTIA WITH REDUCED CANINES #ANINES INCISIFORM TEAMS LED BY "RUCE * -AC&ADDEN OF THE 5NIVERSITY OF OR ABSENT 4ENDENCY FOR ENLARGEMENT OF AN UPPER INCI &LORIDA IN COLLABORATION WITH '%/"/, 3ERVICIO SOR TO CANINIFORM OR TUSK LIKE FORM -OLAR PROTOLOPHS 'EOLGICO DE "OLIVIA ,A 0AZ "OLIVIA AND -ASANARU MUCH GREATER LENGTH THAN METALOPH !NTEROPOSTERIORLY 4AKAI OF THE 0RIMATE )NSTITUTE OF +YOTO *APAN IN COL ORIENTED MEDIAN VALLEY CONTAINING SIMPLE CRISTAE FORM LABORATION WITH &EDERICO !NAYA THEN WITH THE -.(. ING SMALL FOSSETTES IN SOME SPECIES ,INGUAL WALL OF "OL HAVE RECOVERED MATERIAL THAT PERMITS A MUCH IM MOLAR ECTOLOPHS WITH LONGITUDINAL GROOVES OBSCURED EARLY PROVED UNDERSTANDING OF LEONTINIIDS OF 3ALLA 4HIS PRESENT WITH WEAR 4HIRD LOWER INCISOR ENLARGED OR CANINIFORM WORK DESCRIBES TWO SPECIMENS EACH REPRESENTING A NEW "ROAD ROBUST ENTOLOPHID OF MOLARS CONTAINING EARLY SPECIES REFERRED TO THE NEW GENUS !NAYATHERIUM FORMED FOSSETTID THAT IS NOT THE RESULT OF A UNION OF ENTOLOPHID AND HYPOLOPHID (YPOLOPHID OF M LESS THAN -!4%2)!,3 -%4(/$3 !""2%6)!4)/.3 HALF THE TOTAL LENGTH OF THE TALONID DIAGNOSIS MODIFIED 3PECIMENS OF LEONTINIIDS WERE STUDIED IN THE FOLLOWING AFTER #HAFFEE COLLECTIONS GIVEN WITH THEIR ABBREVIATIONST 5!4& $ISCUSSION&LORENTINO !MEGHINO HONORED 5NIVERSIDAD!UTNOMA h4HOMAS &RIASv 0OSTOS¤ "OLIVIA HIS WIFE ,EONTINA BY NAMING THE GENOTYPE OF THIS FAMILY -.(. "OL THE DEPARTMENT OF PALEONTOLOGY -USEO ,EONTINIA SEE 3IMPSON )N THE SAME PUBLICATION .ACIONAL DE (ISTORIA .ATURAL ,A 0AZ "OLIVIA5& 6ER HE ALSO NAMED THREE OTHER GENERA OF LEONTINIIDS TEBRATE 0ALEONTOLOGY #OLLECTION &LORIDA -USEUM OF !NCYLOCOELUS 2ODIOTHERIUM AND ,OXOCOELUS THE LAT .ATURAL (ISTORY 5NIVERSITY OF &LORIDA 'AINESVILLE TER TWO OF WHICH WERE REPRESENTED BY MERE FRAGMENTS &LORIDA 53! !-.( THE !MERICAN -USEUM OF 4HREE SPECIES OF,EONTINIA WERE NAMED IN THIS FIRST PAPER .ATURAL (ISTORY .EW 9ORK 53!&-.( &IELD -U REGARDING THE $ESEADAN !MEGHINO ,EONTINIA SEUM OF .ATURAL (ISTORY #HICAGO )LLINOIS 53! AND GAUDRYI , LAPIDOSA AND , GARZONI,EONTINIA !#- THE 0RATT -USEUM AT !MHERST #OLLEGE !MHERST GAUDRYI IS REGARDED AS THE TYPE FOR THE GENUS WHEREAS -ASSACHUSETTS 53! , LAPIDOSA WAS BASED ON A MANDIBULAR FRAGMENT WITH $ENTAL TERMINOLOGY FOLLOWS THE CONVENTION OF US A SINGLE MOLAR FROM AN ANIMAL ABOUT THE SAME SIZE AS, ING UPPER CASE LETTERS FOR THE UPPER INCISORS CANINES GAUDRYI ,OOMIS REGARDED , LAPIDOSA AS A JUNIOR SYN PREMOLARS AND MOLARS IE ) # 0 - AND LOWER CASE ONYM OF , GAUDRYI ,EONTINIA GARZONI HOWEVER IS FOR THE LOWER SERIES $ECIDUOUS TEETH ARE PREFIXED WITH CLEARLY DIFFERENTIATED FROM , GAUDRYI BY ITS DISTINCTLY hDv SMALL SIZE BUT WAS CONSIDERED INDISTINGUISHABLE FROM 4HE PHYLOGENETIC POSITION OF !NAYATHERIUM WAS !NCYLOCOELUS FREQUENS !MEGHINO BY 0ATTERSON ESTIMATED BY EVALUATING ITS CHARACTERS IN TERMS OF THOSE IN HIS UNPUBLISHED CATALOG OF THE !MEGHINO COLLECTION IN DEFINED BY 6ILLARROEL AND #OLWELL $ANIS FOR THEIR THE -!#. !MEGHINOS INITIAL DESCRIPTION OF THE FAMILY RECENT PHYLOGENETIC ANALYSIS OF THE ,EONTINIIDAE /NLY WAS BASED ON SOMEWHAT FRAGMENTARY MATERIAL LEADING THE CHARACTERS REGARDING THE UPPER TEETH COULD BE HIM TO ERRONEOUSLY REPORT THAT,EONTINIA LACKED CANINES SCORED FOR!NAYATHERIUM SINCE THE LOWER JAWS AND TEETH AND THAT THE CANINIFORM INCISORS WERE ) AND I !FTER REMAIN UNKNOWN 4YPOGRAPHIC ERRORS OF 6ILLARROEL RECEIVING AND STUDYING SUPERIOR MATERIAL OF ,EONTINIA #OLWELL $ANIS WERE CORRECTED CHARACTER SCORES SPP COLLECTED BY HIS BROTHER #ARLOS AT #ABEZA "LANCA OF hv WERE PRINTED BUT WERE NOT DEFINED 4HIS REVISED HE CORRECTED THESE ERRORS !MEGHINO NOTING THAT TAXONCHARACTER MATRIX WITH!NAYATHERIUM AND ITS CHAR THE CANINIFORM INCISORS WERE ) AND I AND THAT,EONTINIA ACTER SCORES ADDED WAS ANALYZED IN 0!50 VERSION HAD BOTH UPPER AND LOWER INCISIFORM CANINES )N THIS 3(/#+%9 .EW ,EONTINIIDS &ROM 3ALLA "OLIVIA 4ABLE 3YNONYMY OF $ESEADAN LEONTINIIDS BASED ON ,OOMIS #HAFFEE 0ATTERSONS UNPUBLISHED CATALOG AND THIS STUDY !NAYATHERIUM GEN NOV !NAYATHERIUM FORTIS SP NOV !NAYATHERIUM EKECOA SP NOV !NCYLOCOELUS !MEGHINO !NCYLOCOELUS FREQUENS !MEGHINO (ENRICOFILHOLIA INAEQUILATERA !MEGHINO ,EONTINIA GARZONI !MEGHINO 2ODIOTHERIUM ARMATUM !MEGHINO ,EONTINIA !MEGHINO , GAUDRYI !MEGHINO , LAPIDOSA !MEGHINO , OXYRHYNCHA !MEGHINO , STENOGNATHA !MEGHINO , FISSICOLIS !MEGHINO 3CARRITTIA 3IMPSON 3 CANQUELENSIS 3IMPSON 3 ROBUSTA 0EREA "OND 4AUBATHERIUM 3ORIA !LVARENGA 4 MAJOR 3ORIA !LVARENGA 4 PAULACOUTTAI 3ORIA !LVARENGA SECOND WORK ON THE $ESEADAN !MEGHINO CLOSELY COMPARE IT TO ,EONTINIA /F IT HE WROTE THAT !MEGHINO NAMED THREE ADDITIONAL SPECIES OF,EONTINIA hTHERE IS NO REASONABLE POSSIBILITY OF VERY CLOSE RELATION ALL OF WHICH WERE THE SAME SIZE AS , GAUDRYI SHIPv 3IMPSON BUT INDICATED THAT IT WAS SIMILAR ,OOMIS SYNONYMIZED SEVERAL SPECIES IN FOUR IN THE REDUCTION OF THE CANINE AND hSOME OTHER DETAILSv GENERA OF LEONTINIIDS REGARDING THESE AS ,EONTINIA WHICH HE DID NOT SPECIFY (E BELIEVED THAT 3CARRITTIA GAUDRYI "ASED ON HIS LARGE SAMPLE OF LEONTINIIDS FROM WAS CLOSEST TO(ENRICOFILHOLIA BUT DID NOT INDICATE WHICH #ABEZA