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! 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 SCARCITYOF 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 "LANCA THE MOST COMMONLY ENCOUNTERED ANI SPECIES SPECIES THEN REFERRED TO (ENRICOFILHOLIA ARE MAL THERE HE INFERRED THAT THERE WAS BUT ONE SEXUALLY NOW REFERRED TO BOTH !STRAPOTHERIA AND ,EONTINIIDAE DIMORPHIC SPECIES REGARDING THOSE WITH LARGE CANINIFORM SEE BELOW BUT TENTATIVELY REFERRED 3CARRITTIA TO THE INCISORS AS BEING MALES AND THOSE WITH SMALLER ONES , ,EONTINIIDAE OXYRHYNCHA !MEGHINO FEMALES (E BELIEVED THAT #HAFFEE REVISED THE DIAGNOSIS OF3CARRITTIA THE SEXUAL DIMORPHISM AND CHANGES IN THE APPEARANCE AND INDICATED THAT IT WAS hVERY CLOSE TO,EONTINIA v OF THE TEETH THAT OCCUR WITH WEAR ACCOUNTED FOR MUCH OF DIFFERING ONLY IN THAT IT WAS THE ) OF 3CARRITTIA THAT THE VARIATION SEEN WITHIN THIS SPECIES WAS CANINIFORM ITS LOWER INCISORS WERE MORE PROCUM 3OON AFTER THE DISCOVERY OF 3CARRITT 0OCKET BENT AND 0 LACKED LINGUAL GROOVES ON THE PROTOCONES 3IMPSON NAMED AND HASTILY DESCRIBED THE hMOST (E PROPOSED AN INFORMAL GROUPING OF THESE TWO GENERA COMMON QUARRY SPECIESv FROM THAT LOCALITY 7ITHOUT AND SUGGESTED THAT3CARRITTIA MAY BE FOUND TO BE A SUB ADEQUATE EXPLANATION HE DESIGNATED hA SPECIMEN NOT GENUS OF ,EONTINIA COLLECTED AND NOW UNIDENTIFIABLE 3IMPSON P v 4HE POORLY KNOWN AND POORLY DEFINED GENUS AS THE HOLOTYPE FOR 3CARRITTIA CANQUELENSIS DESPITE THE (ENRICOFILHOLIA !MEGHINO SHOULD BE DISCUSSED FACT THAT MANY SPECIMENS WERE AT HAND THOUGH UNPRE SINCE IT HAS APPEARED ON FAUNAL LISTS OF 3ALLA AS A LEONTINIID PARED )N NAMING 3CARRITTIA 3IMPSON DID NOT (OFFSTETTER AND AS AN ISOTEMNID -AC&ADDEN ET #%./:/)# 6%24%"2!4%3 0APERS TO (ONOR 3 $AVID 7EBB
&IGURE -OLAR DIMENSIONS WIDTHS VS LENGTHS OF - AND - OF SELECTED $ESEADAN LEONTINIIDS 3(/#+%9 .EW ,EONTINIIDS &ROM 3ALLA "OLIVIA
&IGURE 6IEWS OF THE HOLOTYPE OF!NAYATHERIUM EKECOA N GEN SP -.(. "OL 6 ! OCCLUSAL VIEW OF RIGHT ) OR # 0 n - AND " LATERAL VIEW OF LEFT ROSTRUM ) ) ) OR # POSITION OF 0 NOT PRESERVED AND 0
AL 4HE GENUS WAS BASED LARGELY ON INADEQUATE GENUS IN THE ,EONTINIIDAE THE HEADING WAS ERRONEOUSLY TYPE MATERIAL AND THE RESULTING CHAOS ILLUSTRATES THE FUTIL PRINTED AS h)SOTEMNIDAEv 4HIS hTYPOv IS THE APPARENT ITY OF NAMING FRAGMENTARY REMAINS SEE ALSO 0ASCUAL SOURCE OF ERRONEOUS ASSIGNMENT OF (ENRICOFILHOLIA TO (ENRICOFILHOLIA !MEGHINO WAS FIRST AP )SOTEMNIDAE BY LATER WORKERS IE ,OOMIS PLIED TO 0ARASTRAPOTHERIUM CINGULATUM !MEGHINO -AC&ADDEN ET AL !MEGHINOS INCLUSION OF WITH !MEGHINO EXPLICITLY DESIGNATING (ENRICOFILHOLIA AND#OLPODON IN THE ,EONTINIIDAE WAS 0ARASTRAPOTHERIUM CINGULATUM AS THE TYPE SPECIES TENTATIVE AS HE ADDED THAT THE DEFINITIVE ALLOCATION DE FOR (ENRICOFILHOLIA !MEGHINO (E SIMULTA PENDED UPON THE FAMILIAL PLACEMENT OF#OLPODON 4HIS NEOUSLY NAMED TWO NEW SPECIES OF(ENRICOFILHOLIA ALL hCOLOCACIN DEFINITIVAv APPEARED SOON AFTER !MEGHINO OF WHICH HE CONSIDERED AS BELONGING TO /RDER AND (ENRICOFILHOLIA WAS PLACED IN THE !STRAPOTHERIA ,YDEKKER #OLPODONTIDAE WHICH IN TURN WAS INCLUDED IN THE ORDER ,ATER !MEGHINO B INDICATED THAT (IPPOIDEA (IPPOIDEA WAS COMPOSED OF hCOLPODONTIDSv (ENRICOFILHOLIA WAS CLOSER TO #OLPODON THAN NOTOHIPPIDS AND EQUIDS !STRAPOTHERIUM AND INCLUDED IT AMONG THE ,EONTINIIDAE 7ITH ONE EXCEPTION SPECIES OF(ENRICOFILHOLIA 4HOUGH HE CLEARLY STATED IN THE TEXT THAT HE PLACED THE WERE BASED UPON ONE OR TWO ISOLATED TEETH #HAFFEE #%./:/)# 6%24%"2!4%3 0APERS TO (ONOR 3 $AVID 7EBB
4ABLE $IMENSIONS IN MM OF UPPER TEETH OF THE HOLOTYPES OF !NAYATHERIUM FORTISAND !NAYATHERIUM EKECOA TWO NEW SPECIES OF LEONTINIIDS FROM 3ALLA "OLIVIA
!NAYATHERIUM FORTIS N SP!NAYATHERIUM EKECOA N SP 5!4& 6 -.(. "OL 6
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REGARDED( INTERCINCTA AND ( LEMOINEI AS SYN SPECIES &IG "RYAN 0ATTERSON REGARDED ONYMS FOR 0ARASTRAPOTHERIUM EPHEBICUM AN (ENRICOFILHOLIA INAEQUILATERA AS A JUNIOR SYNONYM OF ASTRAPOTHERE NOT A NOTOUNGULATE /THER SPECIES OF! FREQUENS IN HIS UNPUBLISHED CATALOG OF SPECI (ENRICOFILHOLIA ARE BASED ON SUCH INADEQUATE TYPE MENS IN THE !MEGHINO #OLLECTION MATERIAL THAT THEY ARE BEST IGNORED 4HE EXCEPTION IS 3UMMARY AND REVISION OF $ESEADAN SPECIES OF (ENRICOFILHOLIA INAEQUILATERA !MEGHINO WHICH LEONTINIIDS RECOGNIZED IN THIS WORK ARE PRESENTED IN 4ABLE IS BASED UPON A NEARLY COMPLETE CHEEK TOOTH ROW 6ILLARROEL AND #OLWELL $ANIS PRESENTED A RE -!#. ! 0 - !MEGHINO AFIG CENT AND CREDIBLE PHYLOGENTETIC ANALYSIS OF LEONTINIID GEN )T IS INDISTINGUISHABLE FROM THE LEONTINIID!NCYLOCOELUS ERA 3YNAPOMORPHIES FOR THE ,EONTINIDAE SUGGESTED BY FREQUENS SHOWING THE SAME SUITE OF CHARACTERS SEEN IN THE PHYLOGENETIC ANALYSIS INCLUDE HYPERTROPHIED I AND ! FREQUENS AND IS WELL WITHIN THE SIZE RANGE OF THIS DEMARCATED hLEONTINIID BASINv SENSU 6ILLARROEL 3(/#+%9 .EW ,EONTINIIDS &ROM 3ALLA "OLIVIA
#OLWELL $ANIS OF THE UPPER PREMOLARS 0AMPA IN THE 3ALLA "EDS IN 5NIT JUST BELOW THE %L 0LANIMIENTO A CARBONATE HORIZON THAT IMMEDIATELY UN !.!9!4(%2)5- N GEN DERLIES THE "RANISELLA ,EVEL OF 5NIT 4HE HOLOTYPE 4YPE 3PECIES!NAYATHERIUM EKECOA N GEN WAS FOUND DURING A -.(. "OL EXPEDITION SPONSORED SP BY -ASANARU 4AKAI OF THE 0RIMATE )NSTITUTE OF +YOTO 2EFERRED 3PECIES!NAYATHERIUM FORTIS N GEN *APAN 4HE HOLOTYPE OF ! EKECOA IS THE ONLY KNOWN SP SPECIMEN 'ENERIC $IAGNOSIS3NOUT SHORT WITH CORRESPOND %TYMOLOGY!FTER %KECO A DIMINUTIVE DEITY TO ING LOSS OF AN ANTERIOR UPPER TOOTH APPARENTLY BUT NOT WHOM!YMARAN PEOPLE GIVE MINIATURE OFFERINGS 4HIS IS DEFINITELY THE CANINE ) GREATER THAN ) 0 0 WITH IN REFERENCE TO THE SMALL SIZE OF THE ANIMAL AND TO THE VERTICAL LINGUAL GROOVES AS IN,EONTINIA BUT INDISTINCT GEOGRAPHIC LOCATION OF THE TYPE LOCALITY NOW OCCUPIED OR LACKING IN 0 $IFFERS FROM,EONTINIA BY INCISIFORM BY !NDEAN .ATIVE !MERICANS RATHER THAN CANINIFROM ) AND APPARENT LOSS OF CANINE $ESCRIPTION4HE HOLOTYPE -.(. "OL 6 ,IKE 3CARRITTIA ) IS LARGER THAN ) BUT DIFFERS FROM &IG PRESERVES NEARLY THE ENTIRE UPPER DENTI 3CARRITTIA BY PRESENCE OF THE GROOVED PREMOLAR TION OF THIS SMALL LEONTINIID /NLY THE RIGHT ) ) AND LEFT PROTOCONES 0 ARE MISSING ) IS CANINIFORM AND SIGNIFICANTLY LARGER 'EOGRAPHIC AND 4EMPORAL 2ANGE4WO SPECIES THAN THE OTHER INCISORS AND APPEARS TO HAVE ONLY RECENTLY OF!NAYATHERIUM ARE DESCRIBED BELOW !NAYATHERIUM ERUPTED PRIOR TO THE DEATH OF THE ANIMAL )T HAS A WEAK EKECOA IS FROM 5NIT OF 3ALLA IMMEDIATELY BELOW THE EXTERNAL CINGULUM AND A ROBUST ROOT WHICH IS BROKEN BUT CARBONATE LAYER THAT UNDERLIES THE "RANISELLA LEVEL AND DOES NOT SHOW ANY SIGN OF TAPERING ) AND ) # ARE ! FORTIS IS FROM THE GEOLOGICALLY OLDER 5NIT 4HE BEST SIGNIFICANTLY SMALLER THAN ) ARE ANTEROPOSTERIORLY COM FIT OF PALEOMAGNETIC HORIZONS TO THOSE CONTAINING RADIO PRESSED AND HAVE STRONG CINGULAE 4HE CANINE OR ) ISOTOPICALLY DATED ASHES OF +AY ET AL WOULD PLACE APPEARS TO BE ABSENT THE EXACT DETERMINATION OF THE THE PALEOMAGNETIC HORIZON FROM WHICH THE HOLOTYPE OF ABSENT TOOTH IS DIFFICULT BECAUSE THE PREMAXILLA AND MAX ! EKECOA CAME AS BEING IN #NN BETWEEN AND ILLA ARE SOLIDLY FUSED #RACKS AND SCRATCHES ON THE SPECI +AY ET AL 4HE HOLOTYPE OF ! FORTIS IS MEN ARE MORE CONSPICUOUS THAN ANY SUTURE THAT MAY FROM 5NIT REGARDED AS BEING #HRON NN +AY ET AL REMAIN /NE PALATAL SUTURE LIKE LINE FOLLOWS IMMEDIATELY WHICH HAS AN AGE OF ABOUT -A "ERGGREN ET FROM THE ANTERIOR ROOT OF THE FIRST PREMOLAR SUGGESTING AL !NAYATHERIUM IS ONLY KNOWN FROM 3ALLA THAT THE ABSENT TOOTH IS THE CANINE 4HIS FEATURE HOW "OLIVIA EVER MAY BE A CRACK AND NOT A SUTURE )N ANY EVENT THE %TYMOLOGY! TRIBUTE TO &EDERICO !NAYA $AZA ANIMAL LACKS A COMPLETE UPPER DENTITION 4HE FORMULA FOR HIS HEROIC EFFORTS TO ADVANCE THE SCIENCE OF PALEON OF THE UPPER DENTAL SERIES IS LIKELY BUT MAY BE TOLOGY IN "OLIVIA 4HE LOWER SERIES IS UNKNOWN 4HE FIRST PREMOLAR LIKELY AN UN REPLACED D0 WHICH IS CHARACTERISTICALLY NOT REPLACED IN NOTOUNGULATES IS !.!9!4(%2)5- %+%#/! N SP MUCH SMALLER THAN THE FOLLOWING PREMOLARS 4ABLE AND (OLOTYPE-.(. "OL 6 CAST 5& IS PRESERVED ONLY ON THE RIGHT SIDE )T IS SO WORN THAT ITS UPPER DENTITION WITHIN FRAGMENTARY MAXILLAE AND CROWN IS ESSENTIALLY FEATURELESS EXCEPT FOR A SHALLOW PREMAXILLAE CONTAINING LEFT ) ) AND 0 - AND RIGHT ) FOSSETTE ! DEEP CONSPICUOUS CINGULUM IS PRESENT ON OR # - AND COMPLETE UPPER LEFT DENTITION &IG THE EXTERNAL SURFACE 4HE TOOTH HAS WORN DOWN NEARLY 4ABLE TO THE EXTERNAL CINGULUM AND HAS WORN TO SUCH AN EXTENT $IAGNOSIS#ONSIDERABLY SMALLER THAN THAT THE ANTEROLINGUAL CINGULUM HAS FORMED A FOSSETTE !NAYATHERIUM FORTIS ,EONTINIA GAUDRYI AND 4HOUGH HEAVILY WORN 0 0 ARE COMPARABLE TO 3CARRRITTIA CANQUELENSIS 3IMILAR SIZE TO !NCYLOCOELUS THOSE OF &-.( 0 AN UNNAMED SPECIES DESCRIBED FREQUENS BUT DISTINCT FROM !NCYLOCOELUS FREQUENS BY AND FIGURED BY 0ATTERSON AFIG C AND REFERRED THE PRESENCE OF GROOVED PROTOCONES OF 0 0 AND THE TO,EONTINIA SP 4HE EXTERNAL ENAMEL UNDULATES AROUND LACK OF WELL DEFINED AND PERSISTENTCRISTA THAT GIVES THE A DISTINCT PARASTYLE AND PARACONE RIDGE THEN FOLLOWS A BRANCHED APPEARANCE TO THE MAJOR FOSSETTE OF THE UPPER SMOOTHER PATH TO A SHALLOW METACONE RIDGE ! DISTINCT MOLARS OF! FREQUENS EXTERNAL CINGULUM IS PRESENT ON 0 0 BUT ABSENT ON THE 4YPE ,OCALITY4HE HOLOTYPE WAS FOUND AT 4APIAL MOLARS 4HE PROTOCONE IS NOT CONNECTED AT THE OCCLUSAL #%./:/)# 6%24%"2!4%3 0APERS TO (ONOR 3 $AVID 7EBB
&IGURE /CCLUSAL VIEW OF THE HOLOTYPE OF !NAYA THERIUM FORTIS N GEN SP 5!4& 6 WITH PARTIAL DENTITION ALVEOLI OF RIGHT ) RIGHT 0 - AND LEFT ) ALVEOLUS OF ) AND 0
LEVEL TO THE PARASTYLE EXCEPT ON THE MORE HEAVILY WORN !.!9!4(%2)5- &/24)3 N SP 0 !N ANTEROLINGUAL CINGULUM DEEP TO THE LEVEL OF OC (OLOTYPE5!4& 6 CAST 5& CLUSION CONTACTS THE PROTOCONE WITH THE PROTOLOPH 4HE 2IGHT MAXILLA WITH 0 - OF WHICH 0 AND - ARE POORLY PROTOCONES OF 0 0 HAVE LONGITUDINAL LINGUAL GROOVES PRESERVED RIGHT AND LEFT PREMAXILLAE WITH LEFT ) AND AS IN , GAUDRYI AND THE UNNAMED 0 AND UNLIKE ROOTS OF RIGHT ) AND ROOTS OF BOTH )S AND LEFT MAXILLA 3CARRITTIA CANQUELENSIS ;#HAFFEE = CONTAINING FRAGMENTS OF 0 &IG 4ABLE #ROWNS 4HE MOLARS HAVE THE SAME GENERAL APPEARANCE AS OF INCISOR AND PREMOLARS ARE DAMAGED THOSE OF,EONTINIA AND 3CARRITTIA - IS ROUGHLY SQUARE $IAGNOSIS,ARGER THAN ! EKECOA SIMILAR SIZE IN OCCLUSAL VIEW BUT HAS A PROJECTION OF THE ANTEROLABIAL TO ,EONTINIA GAUDRYI 3CARRITTIA CANQUELENSISAND CORNER SUCH THAT IT IMBRICATES THE 0 EVIDENT ON LEFT 0 4AUBATHERIUM MAJOR -UZZLE SHORTER THAN THOSE OF, - NOT FIGURED 4HOUGH THE ) AND - HAVE NOT COME INTO COMPLETE OCCLUSION THE - IS WORN TO THE LEVEL OF GAUDRYIAND 3 CANQUELENSIS !LVEOLUS OF )LARGER THE INTERNAL CINGULUM 4HE - PROTOLOPH IS NARROW LABI THAN THAT OF ) ) NOT GREATLY ENLARGED BUT SIMILAR TO ) ALLY BUT BROADENS RAPIDLY TO FORM A BULBOUS STRUCTURE AT OF , GAUDRY $IFFERS FROM , GAUDRYI BY SMALL SIZE OF THE PROTOCONE 4HE VALLEY BETWEEN THE PROTOLOPH AND ) AND REDUCTION OF ANTERIOR DENTITION NO APPARENT CA ECTOLOPH IS NARROW AND LACKS BRANCHES OR FOSSETTES - NINE $IFFERS FROM 3 CANQUELENSIS BY SMALLER ) AND LACKS A POSTERIOR FOSSETTE 4HE - IS RELATIVELY AND AB PRESENCE OF LONGITUDINAL LINGUAL GOOVE IN PROTOCONE OF SOLUTELY LONGER THAN THE - HAS A NARROWER PROTOLOPH UPPER PREMOLARS LINGUAL PROTOCONE GROOVE IS ABSENT IN AND A NARROW TRANSVERSELY ORIENTED POSTERIOR FOSSETTE 3 CANQUELENSIS -OLARS RELATIVELY NARROWER THAN THOSE - IS INCOMPLETELY ERUPTED AND BADLY DAMAGED ! DIS OF3 CANQUELENSIS AND , GAUDRYI ,ACKS THE DISTINCT TINCTIVE INTERNAL CINGULUM IS PRESENT ON IT AND ALL THE MOLAR CRISTAE OF 4 MAJOR 4HE REFERRAL OF THIS SPECIES MOLARS BUT NONE OF THESE UPPER MOLARS HAVE EXTERNAL TO!NAYATHERIUM IS TENTATIVE AS IT IS BASED UPON A SHARED CINGULA SUIT OF CHARACTERS PREMOLAR GROOVES OF 0 AND EN 3(/#+%9 .EW ,EONTINIIDS &ROM 3ALLA "OLIVIA
LARGED ) RATHER THAN AN UNAMBIGUOUS SYNAPOMORPHY AGE OF THIS TOOTH SINCE THE POSTERIOR CINGULUM OCCURS 4YPE ,OCALITY&ROM 5NIT AT 0ASTO 'RANDE 3ALLA CLOSER TO THE OCCLUSAL LEVEL THAN THE ANTERIOR CINGULUM "EDS "OLIVIA "OTH )S ARE MISSING BUT THE ALVEOLI AND ROOTS OF ) ARE %TYMOLOGY&ORTIS ,ATIN FOR hSTRONG v IN REFER NOT ENLARGED AS IN , GAUDRYI SEE ,OOMIS AND ENCE TO THE ROBUST NATURE OF THE BEAST AND TO THE CHARAC DISCUSSION BELOW 4HE SPECIMEN DOES NOT PRESERVE ) TER OF THE NAMESAKE & !NAYA OF THE GENUS 0 AND IS SO POORLY PRESERVED IN THE REGION THAT NOTHING $ESCRIPTION4HE HOLOTYPE 5!4& 6 IS CAN BE SAID OF THEM NOT EVEN WHETHER OR NOT ANY OR ALL THE ONLY KNOWN SPECIMEN SO THE FOLLOWING DESCRIPTION IS OF THESE TEETH WERE PRESENT BASED SOLELY ON THIS SPECIMEN 4HE FIRST PREMOLARS ARE NOT PRESERVED AND 0 IS 4HE ) HAS THE SAME GENERAL FORM AS THAT OF BADLY DAMAGED 0 IS NOT HEAVILY WORN AND PRESERVES ,EONTINIA GAUDRYI !N EXTERNAL CINGULUM IS PRESENT SOME OF THE CROWN FEATURES 4HE PROTOCONES OF 0 0 BUT INDISTINCT 4HE CROWN OF ) IS BROKEN .O POSTERIOR HAVE THE VERTICAL LINGUALGROOVES AS SEEN IN , GAUDRYI CINGULUM IS SEEN BUT THIS MAY HAVE BEEN LOST IN THE BREAK AND ,EONTINIA SP OF 0ATTERSON AFIG C 4HE
&IGURE 0HLOGENETIC HYPOTHESIS BASED UPON MOST PARSIMONIUS PHYLOGENETIC TREE 4REE ,ENGTH #ONSISTENCY )NDEX (OMOPLASY )NDEX !NALYSIS BASED UPONA TAXONCHARACTER MATRIX MODIFIED FROM 6ILLARROEL #OLWELL $ANIS TO INCLUDE !NAYATHERIUM AND ITS CHARACTER STATES %VALUATED WITH THE hEXHAUSTIVE SEARCHv OF 0!50 VERSION B 0UTATIVE SYNAPOMORPHIES FOR THE ,EONTINIIDAE AND THE hTROPICALv AND h0ATAGONIANv CLADES NUMBERED CHARACTER STATES OF 6ILLARROEL #OLWELL $AVIS ARE AS FOLLOWS .ODE ,EONTINIIDAE "ASINS OF hLEONTINIID DEPRESSIONv INCOMPLETELY SEPARATED I SOMEWHAT HYPERTROPHIED ,OWER MOLAR POST FOSSETID 5 SHAPED ,OWER MOLAR ENTOFLEXID ROUNDED h0ATAGONIANv NODE ,EONTINIID DEPRESSION WELL SEPARATED 0 LABIAL CINGULUM WEAK OR ABSENT 0OSTERIOR ARM OF UPPER PREMOLAR HYPOFOSSETTE WEAKLY DEVELOPED #HEEKTEETH MESODONT h4ROPICALv CLADE 5PPER MOLAR ANTERIOR LINGUALCINGULUM ABSENT 5PPER MOLAR ANTEROLINGUAL ENAMEL BORDER OF HYPFLEXUS CONSPICUOUS IN RELIEF WITH WEAR ,OWER MOLAR LABIAL CINGULUM ABSENT #%./:/)# 6%24%"2!4%3 0APERS TO (ONOR 3 $AVID 7EBB
BADLY DAMAGED 0 LACKS THIS GROOVE /N THE 0 THE KNOWN IN! FORTIS /NLY CHARACTERS COULD BE SCORED OCCLUSAL SURFACE OF THE PROTOCONE IS ISOLATED SAVE FOR A FOR THE SPECIES OF !NAYATHERIUM AS THE LOWER JAW IS CONNECTION TO THE METALOPH !N UNWORN RIDGE CONNECTS UNKNOWN 4HE LOWER JAW CHARACTERS WERE ALL SCORED AS THE PROTOCONE WITH THE POSTERIOR CINGULUM 4HE ANTERIOR h v !N EXHAUSTIVE SEARCH USING 0!50 VERSION B CINGULUM IS WELL REMOVED FROM THE OCCLUSAL SURFACE AND PRODUCED A SINGLE MOST PARSIMONIOUS PHYLOGENETIC HY BECOMES INCORPORATED INTO THE PROTOLOPH 4HE PARASTYLE POTHESES TREE LENGTH CONSISTENCY INDEX IS DISTINCT AND A SHARP GROOVE CUTS INTO THE ECTOLOPH BE HOMOPLASY INDEX 4HIS ANALYSIS PLACED TWEEN THE PARASTYLE AND THE PARACONE RIDGE 0 0 HAVE !NAYATHERIUMWITHIN THE h0ATAGONIANv CLADE MORE SPE AN EXTERNAL CINGULUM CIFICALLY AMONG!NCYLOCOELOUS AND #OLPODON WITH THE 0 IS MORE WORN THANTHE 0 )T ALSO HAS A BETTER LOSS OF THE CANINE AS THE HYPOTHETICAL SYNAPOMORPHY DEVELOPED PROTOLOPH THAT UNITESTHE PROTOCONE WITH THE UNITING THE THREE GENERA &IG 4HE TWO SPECIES OF ECTOLOPH 4HE POSTERIOR LINGUAL CINGULUM FORMS A CUP !NAYATHERIUM APPEAR TO FORM A DISTINCT CLADE 4HE THAT PROJECTS THIS REGION OF THE TOOTHS SURFACE MEDIALLY LOSS OF ENAMEL LINED PITS OF THE UPPER MOLARS APPEARED 4HIS APPEARS RATHER CONSPICUOUS COMPARED TO THE RATHER AS A PUTATIVE SYNAPOMORPHY UNITING THESE TWO SPECIES WEAK PROTOCONE 4HE 3CARRITA ,EONTINIA hSUB GROUPv OF #HAFEE 4HE - IS BUT MODERATELY WORN #INGULA ARE IS SUPPORTED IN THIS AND THE ORIGINAL ANALYSIS OF PRESENT FROM THE ANTEROLINGUAL PORTION OF THE TOOTH TO THE 6ILLARROEL #OLWELL $ANIS )T IS NOTEWORTHY THAT POSTERIOR BORDER !LL ARE REMOVED FROM THE OCCLUSAL LEVEL THE CHARACTERS THAT UNITE THESE TWO GENERA ARE THOSE EXCEPT FOR THE POSTERIOR CINGULUM WHICH HAS COME INTO FROM THE LOWER DENTAL SERIES WHICH IS LACKING IN BOTH WEAR .ONE OF THE MOLARS HAVE A LABIAL CINGULUM 4HE SPECIES OF!NAYTHERIUM THUS COMPARISON IN THIS ANALY CENTRAL VALLEY IS 9 SHAPED WITH THE ANTERIOR ARM BEING SIS WAS IMPOSSIBLE .ONETHELESS !NAYATHERIUM AP CONSPICUOUSLY LONGER 4HIS ANTERIOR ARM LACKS ANY BIFUR PEARS TO BE FIRMLY ESTABLISHED WITHIN THE h0ATAGONIANv CATION SUCH AS IS PRESENT IN!NCYLOCOELUS 4AUBATHERIUM CLADE OF LEONTINIIDS DISTINCT FROM THE TROPICAL GENERA AND #OLPODON 4AUBATHERIUMAND (UILATHERIUM 0UTATIVE 4HE POSTERIOR CINGULUM OF THE - HAD NOT COME SYNAPOMORPHIES FOR THIS CLADE OF 0ATAGONIAN TAXA PLUS INTO WEAR 4HIS TOOTH HAS A SIMILAR FORM TO THE - THOUGH !NAYATHERIUM INCLUDE A WELL DEMARCATED LEONTINIID DE WITH SIGNIFICANTLY LESS WEAR PRESSION OF 0 0 POORLY DEVELOPED LABIAL CINGULUM OF /NLY THE ANTERIOR MOST PORTION OF THE - HAD BRO 0 AND MESODONT DENTITION KEN THE GUM LINE -OST OF THIS TOOTH IS MISSING BUT PART 4HE NESTING OF !NAYATHERIUM WITHIN THE OF THE ECTOLOPH WAS RECOVERED 4HE INTERNAL WALL OF THIS 0ATAGONIAN GENERA IS SIGNIFICANT IN TERMS OF THE BIOGEOG ECTOLOPH DEMONSTRATES THE VERTICAL RIDGES DESCRIBED BY RAPHY OF THE 3ALLA LEONTINIIDS 3UCH A PHYLOGENETIC PLACE 0ATTERSON AFIG B OF AN UNWORN MOLAR OF MENT IS CONSISTENT WITH THE PREVIOUS OBSERVATION THAT MOST ,EONTINIA GAUDRYI &-.( 0 OTHER UNGULATES OF 3ALLA WERE CLOSELY RELATED TO 0ATAGONIAN $ESEADAN TAXA WITHMOST BEING COGENERIC $)3#533)/. THOUGH SPECIFICALLY DISTINCT 3HOCKEY A 0(9,/'%.9 6ILLARROEL AND #OLWELL $ANIS PRESENTED A 0!,%/%#/,/'9 PHYLOGENY OF LEONTINIIDS THAT ILLUSTRATED TWO DISTINCT CLADES !LTHOUGH LEONTINIIDS ARE AMONG THE MOST COMMON FOR THE FAMILY &IG /NE BRANCH CONTAINED 0ATAGONIAN SPECIMENS COLLECTED AT MOST $ESEADAN LOCALITIES SEE GENERA ,EONTINIA 3CARRITTIA !NCYLOCOELUS AND )NTRODUCTION THEY ARE EXCEEDINGLY SCARCE AT 3ALLA 4HE #OLPODON WHEREAS THE OTHER BRANCH CONTAINED THE TWO SPECIMENS DESCRIBED HERE ARE THE ONLY INSTRUCTIVE 4ROPICAL GENERA4AUBATHERIUM AND (UILATHERIUM 4HE SPECIMENS ENCOUNTERED IN LARGE COLLECTIONS OF FOSSILS OF TROPICAL PLUS 0ATAGONIAN CLADES ARE SHOWN AS A MONO 3ALLA -.(. "OL -.(. 0ARIS &,-.( AND 05 PHYLETIC GROUP WITH THE ISOTEMNID0LEUROSTYLODON AS THE 4HIS SCARCITY OF LEONTINIIDS SUGGESTS SOME ECOLOGICAL DIF SISTER TAXON 6ILLARROEL #OLWELL $ANIS FERENCE BETWEEN 3ALLA AND OTHER WELL KNOWN $ESEADAN 5SING THE CHARACTERS AND SCORING CRITERIA OF LOCALITIES WHERE LEONTINIIDS ARE COMMON 6ILLARROEL AND #OLWELL $ANIS APPENDIX 3ALLA IS AT A SIGNIFICANTLY LOWER LATITUDE THAN THE !NAYATHERIUM EKECOA WAS SCORED AS FOLLOWS CLASSICAL $ESEADAN LOCALITIES OF 0ATAGONIA SO ONE MIGHT AND ! FORTIS WAS SCORED AS CONSIDER THE POSSIBILITY THERE WAS A REGIONAL EFFECT THAT SINCE THE PRESENCE ABSENCE OF THE CANINE IS UN IS CONDITIONS WERE MORE FAVORABLE FOR LEONTINIIDS IN HIGHER 3(/#+%9 .EW ,EONTINIIDS &ROM 3ALLA "OLIVIA
LATITUDES THAN THE MORE TROPICAL LATITUDE OF 3ALLA 4HE ARE MORE CLOSELY RELATED TO 0ATAGONIAN TAXA THAN THEY RELATIVE ABUNDANCE OF LEONTINIIDS IN THE $ESEADAN OF ARE TO OTHER LOW LATITUDE LEONTINIIDS 4HOUGH PHENETICALLY "RAZIL HOWEVER DOES NOT SUPPORT THE NOTION THAT THERE SIMILAR TO 3CARRITTIAAND ,EONTINIA !NAYATHERIUM IS WAS A GROSS REGIONAL EFFECT 3ORIA AND !LVARENGA MORE DERIVED BY ITS APPARENT LOSS OF THE CANINE 4HIS NOTED THAT LEONTINIIDS WERE THE MOST COMMONLY ENCOUN LOSS OF CANINE MAY REPRESENT A SYNAPOMORPHY UNITING TERED FOSSILS IN THE 4AUBAT£ "ASIN WHICH LIES NEAR THE !NAYATHERIUM WITH !NCYLOCOELUSAND #OLPODON 4ROPIC OF #APRICORN ONLY ABOUT SOUTH OF 3ALLA !LSO 4HE SCARCITY OF LEONTINIIDS IS LIKELY DUE TO LOCAL CON THE FACT THAT LEONTINIIDS SURVIVED INTO THE MIDDLE -IOCENE DITIONS THAT WERE MORE FAVORABLE TO HIGH CROWN UNGULATE AT ,A 6ENTA #OLUMBIA 6ILLARROEL 'UERRERO $IAZ TAXA TYPOTHERES AND NOTOHIPPIDS THAN TO LOWER CROWNED -A &LYNN ET AL BUT BECAME EXTINCT AT HERBIVORES )NDEED THE EXTREME WEAR OF THE TEETH OF THE END OF THE #OLHUEHUAPIAN IN 0ATAGONIA ALSO ARGUES THE HOLOTYPE OF! EKECOA A YOUNG ANIMAL AT THE TIME OF AGAINST ANY NOTION THAT TROPICAL REGIONS WERE UNFAVOR DEATH IS VIVID TESTIMONY REGARDING THE EFFECT OF ABRA ABLE TO THEM SIVES IN OR ON THE VEGETATION AT 3ALLA ! CURIOUS FACT REGARDING THE UNGULATE FAUNA OF 3ALLA IS THAT HIGH CROWNED NOTOUNGULATES MESOTHERES !#+./7,%$'%-%.43 INTERATHERIIDS ARCHAEOHYRACIDS HEGETOTHERIIDS AND ) AM GRATEFUL TO THE EDITORS OF THIS VOLUME FOR INVITING ME NOTOHIPPIDS ARE MUCH MORE COMMON THAN ANY UNGULATE TO CONTRIBUTE TO THIS TRIBUTE TO 3 $AVID 7EBB )T IS AN WITH LOWER CROWNS LITOPTERNS PYROTHERES ASTRAPOTHERES HONOR TO SALUTE $R 7EBB IN THIS MANNER 4HE PRESENT /THER LOW CROWNED NOTOUNGULATES IE WORK BEGAN AS PART OF MY DISSERTATION AT THE 5NIVERSITY HOMALODONTOTHERIIDS AND ISOTEMNIDS APPEAR TO BE AB OF &LORIDA AND AS SUCH IT BENEFITED FROM THE CRITICISM OF SENT FROM THE FAUNA 4HE ABUNDANCE OF HYPSODONT TAXA MY COMMITTEE MEMBERS " * -AC&ADDEN #HAIR 3 SUGGESTS THAT THE VEGETATION OF 3ALLA WAS ABRASIVE EI $ 7EBB * !NDERSON $ *ONES AND 2 7OLFF $ #ROFT THER EXTERNAL GRIT OR BIOGENIC SILICATES SEE 3HOCKEY B AND 2 (ITZ PROVIDED CRITICAL REVIEWS AND % 3ARGIS PRO AND REFERENCES THEREIN %ITHER SOURCE OF THE ABRASIVES VIDED HELPFUL COMMENTS 3TUDIES AT THE !-.( WERE SUGGESTS A MORE OPEN HABITAT HAVING LITTLE IN THE WAY OF MADE POSSIBLE BY 2 4EDFORD $ "RYANT - .ORELL AS NON ABRASIVE VEGETATION TO SUPPORT A PUTATIVE BROWSER WELL AS A #OLLECTIONS 3TUDY 'RANT FROM THAT INSTITUTION LIKE !NAYATHERIUM 4HE EXTREME WEAR OF THE CHEEK *OHN &LYNN " 3IMPSON - !LVERY ALONG WITH SUPPORT TEETH PRIOR TO THE ERUPTION OF ) AND - IN THE HOLOTYPE OF THE $EE &UND MADE WORK AT THE &,-.( POSSIBLE OF ! EKECOASUGGEST THAT THE ANIMAL HAD SOME ABRASIVE AND ENJOYABLE - #OOMBS AND , 4HOMAS HELPED WITH FOODS IN ITS DIET (AVING THE CHEEK TEETH WEAR SO MUCH THE STUDY OF THE SPECIMENS OF ,EONTINIA AT THE !#- AT A RELATIVELY YOUNG AGE LIKELY HAD A NEGATIVE EFFECT ON AND # DE -UIZON FACILITATED STUDY OF THE 3ALLA COLLECTION THE FITNESS OF THESE ANIMALS AT THE -.(( 0ARIS !DDITIONAL SUPPORT FOR THIS WORK %COLOGICAL FACTORS RELATING TO BODY SIZE MAY ALSO CAME FROM A .ATIONAL 3CIENCE &OUNDATION )NTERNATIONAL BE A FACTOR IN THE SCARCITY OF LEONTINIIDS AT 3ALLA 4HOUGH 2ESEARCH &ELLOWSHIP 2 -AC#LEAN PRODUCED THE FIG A RIGOROUS ECOMORPHOLOGICAL COMPARISON OF THE 3ALLA URES OF THE HOLOTYPES IN THIS STUDY &INALLY A SPECIAL FAUNA WITH THOSE OF OTHER $ESEADAN LOCALITIES IS WANT THANKS TO &EDERICO !NAYA $AZA WHO HAS HELPED SO MUCH ING THE SCARCITY OF LARGE ANIMALS AT 3ALLA IS FAIRLY CON IN THE FIELD AND IN THE LAB AND EVEN PUT ME UP IN HIS SPICUOUS &OR EXAMPLE ASTRAPTHERES ARE EXCEEDING RARE AT 3ALLA KNOWN ONLY FROM A FEW FRAGMENTS AND THE COM HOME WHEN ) WORKED IN "OLIVIA MON PYROTHERE OF 3ALLA 0YROTHERIUM MACFADDENI IS MUCH SMALLER THAN0 ROMEROI OF 0ATAGONIA 3HOCKEY 2%&%2%.#%3 #)4%$ !NAYA !MEGHINO & %NUM£RATION SYNOPTIQUE DES ESPáCES DE MAMMIFáRES FOSSILS DES FORMATIONS £OCáNES DE 0ATAGOINIE #/.#,53)/.3 "OLET¤N DE LA !CADEMIA .ACIONAL DE #IENCIAS DE #RDOBA 5NLIKE $ESEADAN LOCALITIES OF 0ATAGONIA AND EVEN "RA !MEGHINO & 0REMI£RE CONTRIBUTION LA CONNAISSANCE DE ZIL LEONTINIIDS ARE SCARCE AT 3ALLA 4WO REASONABLY WELL LA FAUNA MAMMALOGIQUE DE COUCHES Ü 0YROTHERIUM PRESERVED AND INSTRUCTIVE SPECIMENS HOWEVER ARE SUF "OLETIN )NSTITUTO 'EOGRÖFICO !RGENTINO FICIENT TO ILLUSTRATE THAT THE LEONTINIIDS OF 3ALLA REPRESENT !MEGHINO & -AMMIFáRES CR£TAC£S DE L!RGENTINE A DISTINCT GENUS OF TWO SPECIES DESCRIBED HERE $EUXIáME CONTRIBUTION Ü LA CONNAISSANCE DE LA FAUNA !NAYATHERIUM EKECOAAND !NAYATHERIUM FORTIS ! MAMMALOGIQUE DE COUCHES Ü 0YROTHERIUM "OLETIN PHYLOGENETIC ANALYSIS SUGGESTS THAT THESE TWO SPECIES )NSTITUTO 'EOGRÖFICO !RGENTINO #%./:/)# 6%24%"2!4%3 0APERS TO (ONOR 3 $AVID 7EBB
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