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4HE /RIGIN OF 4URTLES ! 0ALEONTOLOGICAL 0ERSPECTIVE 7!,4%2 ' */9#% $EPARTMENT OF 'EOSCIENCE 5NIVERSITY OF &RIBOURG &RIBOURG 3WITZERLAND

4HE ORIGIN OF TURTLES AND THEIR UNUSUAL BODY PLAN HAS FASCINATED SCIENTISTS FOR THE LAST TWO CENTURIES /VER THE COURSE OF THE LAST DECADES A BROAD SAMPLE OF MOLECULAR ANALYSES HAVE FAVORED A SISTER GROUP RELATIONSHIP OF TURTLES WITH ARCHOSAURS BUT RECENT STUDIES REVEAL THAT THIS SIGNAL MAY BE THE RESULT OF SYSTEMATIC BIASES AFFECTING MOLECULAR APPROACHES IN PARTICULAR SAMPLING NON RANDOMLY DISTRIBUTED RATE HETEROGENEITY AMONG TAXA AND THE USE AND THE USE OF CONCATENATED DATA SETS -ORPHOLOGICAL STUDIES BY CONTRAST DISFAVOR ARCHOSAURIAN RELATIONSHIPS FOR TURTLES BUT THE PROPOSED ALTERNATIVE TOPOLOGIES ARE POORLY SUPPORTED AS WELL 4HE RECENTLY REVIVED PALEONTOLOGICAL HYPOTHESIS THAT THE -IDDLE 0ERMIAN %UNOTOSAURUS AFRICANUS IS AN INTERMEDIATE STEM TURTLE IS NOW ROBUSTLY SUPPORTED BY NUMEROUS CHARACTERS THAT WERE PREVIOUSLY THOUGHT TO BE UNIQUE TO TURTLES AND THAT ARE NOW SHOWN TO HAVE ORIGINATED OVER THE COURSE OF TENS OF MILLIONS OF YEARS UNRELATED TO THE ORIGIN OF THE TURTLE SHELL !LTHOUGH % AFRICANUS DOES NOT SOLVE THE PLACEMENT OF TURTLES WITHIN !MNIOTA IT SUCCESSFULLY EXTENDS THE STEM LINEAGE OF TURTLES TO THE 0ERMIAN AND HELPS RESOLVE SOME QUESTIONS ASSOCIATED WITH THE ORIGIN OF TURTLES IN PARTICULAR THE NON COMPOSITE ORIGIN OF THE SHELL THE SLOW ORIGIN OF THE SHELL AND THE TERRESTRIAL SETTING FOR THE ORIGIN OF TURTLES

4HE ORIGIN OF TURTLES AND THEIR UNUSUAL BODY PLAN REMAIN SOME OF GIRDLES AND PERHAPS ALSO THE GASTRALIA WITH THE DERMIS AS HAD THE MOST VEXING QUESTIONS IN VERTEBRATE EVOLUTION 4HE DEBATE IS BEEN PREDICTED FOR YEARS BY DEVELOPMENTAL BIOLOGISTS EG UNUSUALLY COMPLEX AS IT INVOLVES SEVERAL INTERRELATED ISSUES IN :ANGERL f "URKE f 'ILBERT ET AL  AND NOT THROUGH THE PARTICULAR THE RELATIONSHIP OF TURTLES RELATIVE TO EXTANT FUSION OF THESE ELEMENTS WITH OSTEODERMS AS HAD LARGELY BEEN EG :ARDOYA AND -EYER f 4ZIKA ET AL  &IELD ET AL  FAVORED BY PALEONTOLOGISTS EG #OX f ,EE f *OYCE ET AL http://doc.rero.ch THE CLOSEST FOSSIL RELATIVES OF TURTLES EG ,AURIN AND 2EISZ f   4HE kNDING OF / SEMITESTACEA FURTHERMORE SPAWNED DE"RAGA AND 2IEPPEL f ,EE f #ARROLL  THE PLACEMENT RENEWED INTEREST IN THE OLD HYPOTHESIS 7ATSON f THAT TURTLES OF THE SHOULDER GIRDLE .AGASHIMA ET AL   ,YSON AND MAY BE RELATED TO THE 0ERMIAN TAXON %UNOTOSAURUS AFRICANUS *OYCE  THE ORIGIN OF THE TURTLE SHELL EG "URKE f #ARROLL  ,YSON ET AL  A   (OWEVER WHILE .AGASHIMA ET AL  3CHEYER ET AL  *OYCE ET AL  THE HYPOTHESIS THAT THE -IDDLE 4RIASSIC / SEMITESTACEA IS A ,YSON ET AL A THE SPEED OF THE ORIGIN OF THE TURTLE SHELL EG DERIVED STEM TURTLE HAS BEEN UNIVERSALLY ACCEPTED EG 2IEPPEL  *OYCE ET AL  AND THE ECOLOGICAL SETTINGS FOR .AGASHIMA ET AL  +URATANI ET AL  !NQUETIN  THE ORIGIN OF TURTLES EG 2IEPPEL AND 2EISZ f *OYCE AND .AGASHIMA ET AL   (IRASAWA ET AL   LIKELY 'AUTHIER  3CHEYER AND 3ANDER   4HE RECENT DISCOVERY BECAUSE OF THE PRESENCE OF A FULLY DEVELOPED PLASTRON THE OF THE INTERMEDIATE FOSSIL TAXON /DONTOCHELYS SEMITESTACEA FROM REINTERPRETATION OF THE ,ATE 0ERMIAN % AFRICANUS AS AN THE ,ATE 4RIASSIC OF #HINA ,I ET AL  SHEDS MUCH LIGHT ON SOME ASPECTS OF THIS DISCUSSION AS IT CLEARLY IS SITUATED OUTSIDE THE PREVIOUSLY RECOGNIZED TURTLE CLADE 4ESTUDINATA HEREIN DEkNED AS #ORRESPONDENCE TO 7ALTER ' *OYCE $EPARTMENT OF 'EOSCIENCE THE CLADE ORIGINATING FROM THE kRST WITH A FULLY 5NIVERSITY OF &RIBOURG &RIBOURG 3WITZERLAND DEVELOPED TURTLE SHELL ;SENSU *OYCE ET AL = CONSISTING OF % MAIL WALTERGJOYCE GMAILCOM INTERLOCKING NEURALS COSTALS PERIPHERALS PYGALS AND PLASTRAL ELEMENTS ;'AFFNEY AND -EYLAN f=  -OST IMPORTANTLY THIS TAXON REVEALS THAT THE ORIGIN OF THE TURTLE SHELL OCCURRED THROUGH THE NOVEL INTERACTION OF THE DORSAL VERTEBRAE AND RIBS THE SHOULDER

1 INTERMEDIATE STEM TURTLE IS LESS QUICKLY GAINING ACCEPTANCE EG -/,%#5,!2 2%35,43 -!9 "% !&&%#4%$ "9 +URATANI ET AL  .AGASHIMA ET AL   (IRASAWA 3934%-!4)# ")!3%3 ET AL    4HIS RELUCTANCE LIKELY STEMS FROM A SERIES OF 4HE DISCUSSION REGARDING THE ORIGIN OF TURTLES HAS BEEN DOMINATED MISCONCEPTIONS ABOUT AMNIOTE SYSTEMATICS AND THE FOSSIL RECORD OVER THE COURSE OF THE LAST  YEARS BY PHYLOGENETIC HYPOTHESES IN PARTICULAR THE REMAINING AMBIGUITY REGARDING THE PHYLOGE OBTAINED FROM MOLECULAR DATA AND ) HEREIN ONLY INTEND TO PROVIDE NETIC POSITION OF TURTLES 4HE PURPOSE OF THIS CONTRIBUTION IS A BRIEF OVERVIEW 4HE kRST DECADE OF MOLECULAR RESEARCH DID NOT THEREFORE TO CLEAR UP THESE MISCONCEPTIONS AND TO SUMMARIZE THE PROVIDE CONSISTENT RESULTS FOR AMNIOTE PHYLOGENY AND TURTLES WERE MOUNTING EVIDENCE IN FAVOR OF % AFRICANUS AS AN INTERMEDIATE VARIOUSLY PLACED AS SISTER TO !VES 0OLLOCK ET AL  #OTTON AND STEM TURTLE 0AGE  #ROCODILIA (EDGES AND 0OLING f -ANNEN AND ,I f #AO ET AL  $IAPSIDA EG #ASPERS ET AL f 3TRIMMER 4(% 42!$)4)/.!, #,!33)&)#!4)/. /& &/33), 2%04),%3 AND VON (AESELER f ,EPIDOSAURIA (EDGES f 4HECODONTIA 53).' 4%-0/2!, &%.%342!% )3 /54 /& $!4% IE -AMMALIA þ !RCHOSAURIA 'ARDINER f OR !RCHOSAURIA 7ILLISTON f PUBLISHED A FUNDAMENTAL TREATMENT OF THE PHYLOGENY EG 0LATZ AND #ONLON f #AO ET AL f :ARDOYA AND -EYER AND CLASSIkCATION OF AMNIOTES BASED ON /SBORNfS @ WORK ON f +UMAZAWA AND .ISHIDA f 2EST ET AL   /VER THE TEMPORAL FENESTRATION THAT WAS HIGHLY INlUENTIAL FOR MANY DECADES COURSE OF THE SECOND DECADE HOWEVER THE VAST MAJORITY OF 4HE BASIS OF THIS CLASSIkCATION WAS THE NUMBER OF OPENINGS IN THE ANALYSES UTILIZING MOLECULAR DATA SUPPORTED A SISTER GROUP TEMPORAL REGION OF THE SKULL !CCORDING TO 7ILLISTONfS CLASSIkCATION RELATIONSHIP OF TURTLES TO ARCHOSAURS USING A BROAD RANGE OF TAXA THAT LACK TEMPORAL FENESTRAE ARE GROUPED AS !NAPSIDA DATA AND ANALYTICAL TECHNIQUES EG )WABE ET AL  (UGALL EG TURTLES VARIOUS BASAL TAXA WITH A PAIR OF UPPER ET AL  3HEN ET AL  #HIARI ET AL  #RAWFORD ET AL TEMPORAL FENESTRAE AS 0ARAPSIDA AKA %URYAPSIDA EG ICHTHYO  &ONG ET AL  !BRAMYAN ET AL  3CHAFFER ET AL SAURS AND SQUAMATES TAXA WITH A PAIR OF LOWER TEMPORAL FENESTRAE  7ANG ET AL  ALTHOUGH OCCASIONAL SUPPORT WAS ALSO AS 3YNAPSIDA EG MAMMALS AND TAXA WITH TWO PAIRS OF TEMPORAL FOUND FOR A SISTER GROUP RELATIONSHIP WITH CROCODILIANS EG FENESTRAE ARE GROUPED AS $IAPSIDA EG CROCODILIANS 3PHENODON  3HEDLOCK ET AL  4ZIKA ET AL   4HE ONLY MAJOR MOLECULAR 4HE MOST INlUENTIAL NEW STATEMENT UNDER 7ILLISTONfS MODEL IS THAT STUDY TO ARRIVE AT A TRULY CONlICTING RESULT IN THE LAST  YEARS WAS THE ANAPSID CONDITION IS PRIMITIVE AND THAT TURTLES ARE CONSEQUENTLY THE STUDY OF ,YSON ET AL  UTILIZING MICRO$.! DATA BUT THE TO BE CONSIDERED THE MOST PRIMITIVE OF LIVING AMNIOTES -OST RECENT EXPANSION AND REANALYSES OF THIS DATA SET BY &IELD ET AL SUBSEQUENT AUTHORS FOLLOWED 7ILLISTONfS MODEL AND PLACED  AND 4HOMSON ET AL  RETRIEVED ARCHOSAURIAN AFkNITIES gANAPSIDu FROM THE 0ERMIAN AND 4RIASSIC AS THE gDIRECT AS WELL -OLECULAR ANALYSES ARE THEREFORE CONVERGING UPON A ANCESTORSu TO TURTLES SUCH AS 'REGORY f DIADECTIDS TURTLE AND ARCHOSAUR RELATIONSHIP /LSON f OR gCAPTORHINOMORPHSu #ARROLL f  3OME OF THE GREATEST CONCERNS FOR ALL TYPES OF SYSTEMATICS ARE !LTHOUGH THE TEMPORAL REGION PROVIDES IMPORTANT PHYLOGE REFERRED TO AS gSYSTEMATIC BIASESu /NE OF THE kRST SUCH BIASES TO BE NETIC INFORMATION FOR AMNIOTE PHYLOGENY THE LAST DECADES OF DISCOVERED IS ALSO ONE OF THE MOST INTUITIVE TO UNDERSTAND RESEARCH HAVE SHOWN THAT 7ILLISTONfS @ SYSTEM IS FAR TOO &ELSENSTEIN @ NOTED THAT PARSIMONY METHODS WILL RETRIEVE THE http://doc.rero.ch SIMPLISTIC BECAUSE MANY FORMERLY HOMOGENOUS GROUPS WERE WRONG RESULT IN A FOUR TAXON SYSTEM IF TWO UNRELATED BRANCHES SHOWN TO EXHIBIT MULTIPLE CONDITIONS EG $ILKES f 2IEPPEL ARE SIGNIkCANTLY LONGER THAN THE OTHERS BECAUSE RANDOMLY B 4SUJI AND -ÌLLER  AND BECAUSE PHYLOGENETIC ACCUMULATED HOMOPLASY IN THE LONG BRANCHES WILL WRONGFULLY ANALYSES HAVE SHOWN THAT THE NUMBER OF TEMPORAL FENESTRAE SERVE AS SYNAPOMORPHIES IE SECONDARY HOMOLOGIES SENSU DE CHANGED MULTIPLE TIMES THROUGHOUT AMNIOTE EVOLUTION EG 0INNA f AN EFFECT REFERRED TO AS LONG BRANCH ATTRACTION -OST -ÌLLER   4HE POSSIBLE PLACEMENT OF ANAPSID TURTLES WITHIN VEXINGLY THE ADDITION OF DATA DOES NOT SOLVE THIS PROBLEM BUT CROWN $IAPSIDA ONLY UNDERLINES THIS POINT AS IT IMPLIES THE RATHER AMPLIkES IT THEREBY PRODUCING GREATER CONkDENCE FOR THE SECONDARY LOSS OF BOTH TEMPORAL FENESTRAE 2IEPPEL AND DE"RAGA WRONG RESULT &ELSENSTEIN f  4AKEN TO THE EXTREME THE USE OF  2IEPPEL A A CONCLUSION ALREADY PROPOSED BY EARLY THE TOTAL GENOMES WILL NOT SOLVE PROBLEMS ASSOCIATED WITH LONG OPPONENTS TO THE 7ILLISTON MODEL "ROOM f 'OODRICH f BRANCH ATTRACTION BUT ONLY COMPOUND THEM DE"EER f AND READILY ACCEPTED BY THOSE FAVORING THIS !LTHOUGH THE CONSISTENT SIGNAL BEING RETRIEVED FROM INCREAS PHYLOGENETIC ARRANGEMENT (OWEVER EVEN IF TURTLES EVOLVED INGLY DIVERSE SETS OF MOLECULAR DATA IS IMPRESSIVE THE PERSISTENT FROM DIAPSID ANCESTORS THERE IS NO REASON TO BELIEVE A PRIORI THAT MORPHOLOGICAL SIGNAL AGAINST THE ARCHOSAURIAN RELATIONSHIPS OF THE LOSS OF BOTH TEMPORAL FENESTRATIONS OCCURS IN SYNCHRONY WITH TURTLES EG 2EISZ AND ,AURIN f DE"RAGA AND 2IEPPEL f ,EE THE ACQUISITION OF THE TURTLE SHELL 4HE EARLY STEM LINEAGE OF TURTLES f -ÌLLER  ,YSON ET AL  ,EE  SEE BELOW IS THEREFORE EXPECTED TO POTENTIALLY BE POPULATED WITH TAXA WITH COMBINED WITH THE COMPLETE LACK OF A MORPHOLOGICAL SIGNAL IN THE DIAPSID CONDITION &ENESTRATION ALONE IS THEREFORE NOT A SUPPORT OF THE ARCHOSAUR HYPOTHESIS 2IEPPEL A ,YSON ET AL STRONG ARGUMENT FOR OR AGAINST THE PLACEMENT OF %UNOTOSAURUS  DESPITE EARNEST ATTEMPTS TO kND SOME "HULLAR AND "EVER AFRICANUS OR ANY OTHER TAXON ALONG THE STEM LINEAGE OF TURTLES  GIVES REASONS TO SPECULATE IF MOLECULAR HYPOTHESIS ARE

2 PERHAPS CONVERGING UPON THE SAME RESULT BECAUSE OF A SYSTEMATIC SEPARATELY AND ARRIVED AT THE SURPRISING CONCLUSION THAT THE BIAS 4HREE PARTIALLY INTERRELATED BIASES COME TO MIND THAT MIGHT THREE PRIMARY PLACEMENTS OF TURTLES IE AS SISTER TO $IAPSIDA BE RELEVANT TO THIS PARTICULAR PHYLOGENETIC CONUNDRUM SAMPLING ,EPIDOSAURIA OR !RCHOSAURIA ARE SUPPORTED BY ROUGHLY THE SAME RATE HETEROGENEITY AND THE USE OF CONCATENATED DATA SETS NUMBER OF GENES AND THAT AN ARCHOSAURIAN SIGNAL EMERGES ONLY 3AMPLING IS OF ENDURING IMPORTANCE IN SYSTEMATICS EG THROUGH THE CONCATENATION OF THE DATA &ROM A MOLECULAR ,ECOINTRE AND 0HILIPPE f 'RAYBEAL f (ILLIS f 2ANNALA PERSPECTIVE TURTLES ARCHOSAURS AND LEPIDOSAURS THEREFORE ET AL f BECAUSE THE INCLUSION OF INTERMEDIATE MORPHOLOGIES OR SEEM TO FORM A HARD POLYTOMY ,U ET AL  LIKELY DUE TO GENOTYPES HELPS AVOID A BROAD SET OF SYSTEMATIC BIASES INCLUDING THEIR RAPID DIVERSIkCATION AND THE RECENT TREND TOWARDS THE ABOVE MENTIONED LONG BRANCH ATTRACTION 'AUTHIER ET AL RECOVERING ARCHOSAURIAN SISTER GROUP RELATIONSHIPS FOR TURTLES fB $ONOGHUE ET AL f  (OWEVER WHEREAS PALEONTOLOGISTS AT MAY BE THE RESULT OF RATE HETEROGENEITY IN COMBINATION WITH THE LEAST HAVE THE POTENTIAL TO SAMPLE ALL OF LIFE DESPITE MASSIVE USE OF CONCATENATED DATA TAPHONOMIC kLTERS "ENTON ET AL  MOLECULAR SYSTEMATISTS 4HERE ONLY IS ONE TREE OF LIFE AND IT IS APPARENT THAT EITHER THE CAN ONLY SAMPLE EXTANT TAXA OR AT BEST NEWLY EXTINCT TAXA MOLECULAR OR THE MORPHOLOGICAL SIGNAL IS WRONG !S OUTLINED .ICHOLLS   4HIS ISSUE IS COMPOUNDED BY THE FACT THAT EXTANT ABOVE THERE ARE GOOD REASONS TO SERIOUSLY ENTERTAIN THE IDEA THAT TAXA DO NOT SAMPLE THE !MNIOTE TREE AT RANDOM BUT RATHER AS THE MOLECULAR DATA MAY POSSIBLY BE CONVERGING ON THE WRONG EIGHT DISCRETE CLADES IE !VES #ROCODILIA -ONOTREMATA ANSWER DUE TO SYSTEMATIC BIASES ASSOCIATED WITH SAMPLING RATE -ARSUPIALIA 0LACENTALIA 3QUAMATA 3PHENODON AND 4ESTUDINES HETEROGENEITY AND THE USE OF CONCATENATED DATA &UTURE THAT DIVERSIkED WITHIN THE LAST  -A AND THAT ARE INTERCONNECTED MOLECULAR STUDIES SHOULD THEREFORE FOCUS THEIR EFFORTS ON BY LINEAGES SPANNING MANY HUNDREDS OF MILLIONS OF YEARS EG IDENTIFYING SEQUENCE DATA RELEVANT TO RESOLVING DIVERGENCES 7ANG ET AL   3IGNIkCANT PORTIONS OF THE TREE IN PARTICULAR THAT OCCURRED DURING THE LATE 0ALEOZOIC AND CALIBRATE RATE THE STEM LINEAGES OF VARIOUS CLADE COMBINATIONS ARE THEREFORE HETEROGENEITY THROUGH THE USE OF FOSSILS 4HE TRADITIONAL FOCUS INHERENTLY OUT OF REACH FOR MOLECULAR SYSTEMATISTS AND MUST BE ON OBTAINING EVER INCREASING AMOUNTS OF SEQUENCE DATA FROM MODELED 4HE FACT THAT THE THREE PRIMARY LINEAGES OF REPTILES LIKELY EXTANT TAXA IS NOT SUFkCIENT ANY LONGER 0HILIPPE ET AL   DIVERSIkED FROM ONE ANOTHER WITHIN A SHORT TIME IN THE MIDDLE TO 5NTIL THESE POINTS HAVE BEEN ADDRESSED THE DEBATE REGARDING THE LATE 0ERMIAN 7ANG ET AL  DOES NOT MAKE MATTERS EASIER FOR PLACEMENT OF TURTLES WITHIN !MNIOTA REMAINS FAR FROM OVER SYSTEMATISTS USING NEONTOLOGICAL DATA ALONE 6ARYING RATES OF MOLECULAR EVOLUTION MAY BE THE BASIS OF YET 34%- 4524,%3 #!. "% )$%.4)&)%$ 7)4(/54 ANOTHER SYSTEMATIC BIAS !LTHOUGH RATES OF MOLECULAR EVOLUTION 5.$%234!.$).' 4(% 0,!#%-%.4 /& 4524,%3 HAVE BEEN SHOWN TO BE VARIABLE ACROSS THE TREE ALL MOLECULAR )N CONTRAST TO MOLECULAR SYSTEMATISTS PALEONTOLOGISTS NOT ONLY METHODS RELY AT LEAST PARTIALLY ON A CERTAIN AMOUNT OF RATE HAVE THE ABILITY TO INVESTIGATE THE PLACEMENT OF TURTLES AMONG HOMOGENEITY AMONG LINEAGES EG 0HILIPPE ET AL   EXTANT AMNIOTES BUT ALSO THE ORGANISMS THAT POPULATE THE STEM 5NFORTUNATELY RATES DO NOT VARY AT RANDOM ACROSS THE TREE BUT LINEAGE OF TURTLES IE THE PHYLOGENETIC LINEAGE OF EXTINCT RATHER HAVE A STRONG TAXONOMIC SIGNAL AS WELL &OR INSTANCE THE ORGANISMS THAT LEADS TO THE MODERN TURTLE CLADE  (OWEVER IT IS http://doc.rero.ch STUDY OF (UGALL ET AL  REVEALED THAT CROCODILIANS AND TURTLES NOT NECESSARY TO UNDERSTAND THE PLACEMENT OF TURTLES WITHIN THE SHOWED THE SLOWEST RATE OF MOLECULAR EVOLUTION AMONG TETRAPODS AMNIOTE SYSTEM TO ASSIGN FOSSILS TO THEIR STEM LINEAGE WITH WHILE PASSERINE BIRDS AND SNAKES AMONG OTHERS SHOW SOME OF THE CONkDENCE &OR INSTANCE RECENT CLADISTIC ANALYSES HAVE ASSIGNED A FASTEST RATES )T IS THEREFORE POSSIBLE THAT THE RATE AVERAGING EFFECT BROAD SAMPLE OF SHELLED AMNIOTES FROM THE ,ATE 4RIASSIC EG OF VARIOUS MOLECULAR METHODS PULLS TAXA WITH SLOW RATES TOWARDS 0ROTEROCHERSIS ROBUSTA 0ROGANOCHELYS QUENSTEDTI 0ALAEOCHERSIS THE TIPS OF THE TREE OR TOWARDS ONE ANOTHER WHILE PUSHING TAXA TALAMPAYENSIS TO THE 0LEISTOCENE EG -EIOLANIA PLATYCEPS TO THE WITH FAST RATES TOWARDS THE BASE OF THE TREE )NDEED MOST TAXA WITH STEM LINEAGE OF TURTLE EG *OYCE  !NQUETIN  3TERLI CONTROVERSIAL PHYLOGENETIC RELATIONSHIPS SEE 'AUTHIER ET AL ET AL  AND THESE RESULTS HAVE BEEN UNCONTROVERSIAL IN REGARDS  FOR EXAMPLES WITHIN ,EPIDOSAURIA DISPLAY DEVIANT RATES OF TO THE PLACEMENT OF THESE TAXA WITHIN 4ESTUDINATA EG 'AFFNEY MOLECULAR EVOLUTION (UGALL ET AL   ET AL  *OYCE AND 3TERLI  *OYCE ET AL A EVEN 4HE VAST MAJORITY OF RECENT MOLECULAR ANALYSES OF TURTLE THOUGH NONE OF THESE STUDIES UTILIZED GLOBAL AMNIOTE MATRICES 4HE RELATIONSHIPS USED CONCATENATED DATA IE A SUPER MATRIX REASON FOR THIS LACK OF CONTROVERSY IS OF COURSE THE PRESENCE OF APPROACH USING SEQUENCE DATA FROM S TO S OF GENES  FULLY DEVELOPED TURTLE SHELLS IN ALL OF THESE TAXA A gKEY CHARACTERu 4HIS APPROACH IS UTILIZED REGULARLY AS IT ALLOWS INCORPORATION OF ASSUMED TO HAVE GREAT WEIGHT *OYCE AND 3TERLI   THE GREATEST AMOUNT OF DATA BUT STUDIES HAVE SHOWN THAT IT IS ,I ET AL  USED A GLOBAL MATRIX TO ASSIGN /DONTOCHELYS PRONE TO SYSTEMATIC ERROR AS WELL EG .ISHIHARA ET AL  SEMITESTACEA TO THE STEM LINEAGE OF TURTLE AND ,YSON ET AL  3ALICHOS AND 2OKAS   5SING ONE OF THE LARGEST DATA SETS UTILIZED THE SAME MATRIX TO ASSIGN %UNOTOSAURUS AFRICANUS TO THE ASSEMBLED TO DATE ,U ET AL  RECENTLY EVALUATED THE STEM LINEAGE AS WELL )NTERESTINGLY WHEREAS THE ANALYSIS OF ,I ET AL PHYLOGENETIC SIGNAL CONTAINED IN   ORTHOLOGOUS GENES  PLACES TURTLES ALONG THE STEM OF ,EPIDOSAURIA NOT !RCHOSAURIA

3 THE ANALYSIS OF ,YSON ET AL  HYPOTHESIZED THAT TURTLES ARE SHAPE OF THE DORSAL VERTEBRAE AND THE EXPANDED RIBS FOUND IN THIS OUTSIDE OF $IAPSIDA (OWEVER IS THE HYPOTHESIS THAT % AFRICANUS IS A TAXON WITH THOSE OF TURTLES BUT HESITATED FROM PLACING IT ALONG THE STEM TURTLE CONTINGENT UPON THE PLACEMENT OF THE ENTIRE CLADE OUTSIDE STEM LINEAGE OF TURTLES 7ATSON f LATER REINVESTIGATED % OF $IAPSIDA )S IT POSSIBLE THAT % AFRICANUS IS A STEM TURTLE BUT THAT AFRICANUS BASED ON ADDITIONAL MATERIAL AND EXPANDED THE LIST OF TURTLES ARE NEVERTHELESS SISTER TO ARCHOSAURS OR LEPIDOSAURS POSSIBLE SYNAPOMORPHIES TO INCLUDE THE PURPORTED PRESENCE OF  )N A RECENT STUDY ,EE  EXPLORED THIS QUESTION BY ELONGATE DORSAL VERTEBRAE ACTUALLY  SEE BELOW THAT ARE MUCH RETROkTTING TURTLES AND %UNOTOSAURUS AFRICANUS TO PREVIOUSLY LONGER THAN THE CERVICALS AND CAUDALS THE PURPORTED PRESENCE OF  PUBLISHED MORPHOLOGICAL DATA SETS WITH CONTRADICTORY OUTCOMES EXPANDED RIBS ACTUALLY  SEE BELOW AN ANTERIOR SHIFT OF THE IN REGARDS TO TURTLE ORIGINS )N ALL ANALYSIS % AFRICANUS WAS PLACED DORSAL RIBS RELATIVE TO THE DORSAL VERTEBRAE AND THE PURPORTED AS SISTER TO TURTLES AT THE BASE OF THE REPTILIAN TREE MUCH AS ALREADY PRESENCE OF DERMAL OSSIkCATIONS NOT CONkRMED TO BE PRESENT IN NOTED BY ,YSON ET AL  THEREBY GIVING ADDITIONAL CREDENCE TO LATER STUDIES ,YSON ET AL   -ANY YEARS LATER 0ARSONS AND THE NOTION THAT % AFRICANUS IS A TREE CHANGING TAXON SEE 7ILLIAMS f DISMISSED A RELATIONSHIP OF % AFRICANUS WITH SAMPLING ABOVE  )N THE NEXT ANALYSIS ,EE  FORCED TURTLES AS TURTLES BECAUSE THE RIBS OF MODERN TURTLES AS SEEN IN MARINE SISTER TO ARCHOSAURS AS SUPPORTED BY THE ABUNDANT MOLECULAR DATA TURTLES WITH REDUCED COSTALS ARE PURPORTEDLY NOT AS WIDE AS THOSE SEE ABOVE AND NOTICED THAT THIS ARRANGEMENT ONLY REQUIRES THE OF % AFRICANUS 4HIS ASSESSMENT WAS NOT ONLY INCORRECT SEE ADDITION OF FEW STEPS -ORE IMPORTANTLY HOWEVER EVEN THOUGH BELOW BUT ALSO HAD FAR REACHING EFFECTS AS MOST FOLLOWING % AFRICANUS WAS ALLOWED TO lOAT IN ALL ANALYSES ALSO THOSE WITH AUTHORS DISREGARDED THE %UNOTOSAURUS HYPOTHESIS COMPLETELY FORCED TOPOLOGIES IT CONSISTENTLY WAS RETRIEVED AS A STEM TURTLE 4HE DISCOVERY OF /DONTOCHELYS SEMITESTACEA ,I ET AL  4HE ANALYSIS OF ,EE  IS IMPORTANT FOR TWO REASONS )T kRST IMMEDIATELY RENEWED INTEREST IN A POTENTIAL LINK OF % AFRICANUS DEMONSTRATES THAT TWO COMPETING MORPHOLOGICAL ANALYSES WITH WITH TURTLES #ARROLL  BECAUSE / SEMITESTACEA BRIDGES THE DIFFERENT TAXONOMIC FOCUSES IE DE"RAGA AND 2IEPPEL f ;WITH APPARENT MORPHOLOGICAL GAP BETWEEN % AFRICANUS AND TRUE MODIkCATION FROM 2IEPPEL AND 2EISZ f ,I ET AL  ,YSON TURTLES IE AMNIOTES WITH A FULL TURTLE SHELL  ! LINK HAS SINCE ET AL  AND ,EE = AND -ÌLLER AND 4SUJI  ;WITH BEEN FORMALLY PROPOSED ,YSON ET AL  AND BACKED UP BY A MODIkCATIONS FROM -ODESTO ET AL  4SUJI ET AL   SERIES OF ADDITIONAL STUDIES ,YSON AND *OYCE  ,EE  AND ,EE = CONVERGE ON THE PLACEMENT OF TURTLES OUTSIDE OF ,YSON ET AL A B   4HE LIST OF UNIQUE CHARACTERS CROWN $IAPSIDA BUT THAT THIS RESULT MAY NOT BE STATISTICALLY THAT UNITE % AFRICANUS WITH / SEMITESTACEA AND TURTLES HAS SIGNIkCANT RELATIVE TO OTHER SOLUTIONS 4HE MORPHOLOGICAL SIGNAL IS GROWN TO BE IMPRESSIVE &IG   &OR SIMPLICITY ) WILL DISCUSS THESE THEREFORE NOT ABLE TO RESOLVE THE PLACEMENT OF TURTLES WITHIN IN ANATOMICAL GROUPS EVEN THOUGH THEY COULD BE FURTHER AMNIOTES FOR THE MOMENT JUST LIKE THE MOLECULAR DATA PERHAPS ANATOMIZED BECAUSE OF A SAMPLING BIAS OR PERHAPS BECAUSE THE THREE MAJOR REPTILIAN LINEAGES DIVERGED FROM ONE ANOTHER VERY RAPIDLY DURING $ORSAL 6ERTEBRAL #OUNT%LONGATION OF $ORSAL 6ERTEBRAE 2ELATIVE TO THE ,ATE 0ALEOZOIC THE SAME TWO BIASES EFFECTING MOLECULAR DATA /THER 6ERTEBRAE 4HE ANALYSIS OF ,EE  SECONDLY REVEALS THAT THE INTERPRE !S ALREADY NOTED BY 3EELEY f AND 7ATSON f %UNOTOSAURUS http://doc.rero.ch TATION OF % AFRICANUS AS AN INTERMEDIATE STEM TURTLE IS HIGHLY AFRICANUS CAN BE UNITED WITH TURTLES BASED ON DORSAL VERTEBRAL ROBUST RELATIVELY IMMUNE TO PERTURBANCES AND THAT THIS PLACE COUNT AND ELONGATION OF THE DORSAL VERTEBRAE RELATIVE TO MENT IS INDEPENDENT FROM THE PLACEMENT OF TURTLES WITHIN SURROUNDING PARTS OF THE BODY 4HE NUMBER OF VERTEBRAE VARIES !MNIOTA 3O EVEN THOUGH MORPHOLOGY CANNOT RESOLVE THE GREATLY AMONG VERTEBRATES BUT SUCH AN EXTREME REDUCTION IS PLACEMENT OF TURTLES WITHIN AMNIOTES WITH GREAT CONkDENCE UNIQUE TO THESE TAXA -ÌLLER ET AL   -ORE IMPORTANTLY EITHER THE SIGNAL REFERRING % AFRICANUS TO THE STEM LINEAGE OF WHEREAS THE REDUCTION IN VERTEBRAL NUMBER IN MOST TAXA IS TURTLES IS STRONG AND CANNOT BE lIPPANTLY BRUSHED ASIDE 4HIS CORRELATED WITH A SHORTENING OF THE DORSAL COLUMN % AFRICANUS CONTRASTS THE PLACEMENT OF SAUROPTERYGIANS ALONG THE STEM / SEMITESTACEA AND TURTLES SHOW A UNIQUE ELONGATION OF THESE LINEAGE OF TURTLES WHICH HAS ONLY BEEN SUPPORTED BY A SINGLE VERTEBRAE RELATIVE TO THE SURROUNDING REGIONS 4HIS TREND IS ONLY ANALYSIS DE"RAGA AND 2IEPPEL f AND SOME OF THE FOLLOWING REVERSED IN SOME MODERN TURTLES AFTER THE ACQUISITION OF EXTREME STUDIES UTILIZING THIS MATRIX BUT IS EASILY PERTURBED THROUGH THE NECK MOBILITY 7ERNEBURG ET AL   ADDITION OF NEW TAXA ANDOR CHARACTERS 2IEPPEL AND 2EISZ f -ODERN TURTLES UNIVERSALLY HAVE  DORSAL RIBS OF WHICH THE ,YSON ET AL  ,EE   !LL STUDIES THAT UTILIZE kRST AND TENTH ARE GREATLY SHORTENED AND THE SECOND TO NINTH FORM SAUROPTERYGIANS AS MODEL ORGANISMS FOR THE ORIGIN OF TURTLES COSTALS IE RIBS THAT INTERACT WITH THE DERMIS TO FORM A COMPOSITE THEREFORE LACK A SOLID PHYLOGENETIC FOOTING WITH METAPLASTIC BONE MORE DETAILS BELOW  4HE MOST PRIMITIVE TURTLES WITH A WELL PRESERVED SHELL SUCH AS 0ROGANOCHELYS 4(% %5./4/3!5253 (90/4(%3)3 QUENSTEDTI OR 0ROTEROCHERSIS ROBUSTA ALSO REVEAL THE PRESENCE )N THE TYPE DESCRIPTION OF %UNOTOSAURUS AFRICANUS 3EELEY f OF  DORSAL RIBS BUT THE kRST IS GREATLY ELONGATED LIKE THE LAST ALREADY COMPARED THE NUMBER OF DORSAL VERTEBRAE THE ELONGATE CERVICAL WHEREAS THE REMAINING NINE FORM COSTALS 'AFFNEY f

4 http://doc.rero.ch

&IGURE  ! SIMPLIkED CLADOGRAM SUMMARIZING THE HIERARCHICAL DISTRIBUTION OF CHARACTERS AND TAXA DISCUSSED IN THE TEXT

5 kG  *OYCE  *OYCE ET AL A  4HE CERVICAL LIKE NATURE APPOSITIONAL PERICHONDRAL AND METAPLASTIC PORTIONS OF THE RIB OF THE kRST THORACIC VERTEBRA AND RIB OF BASAL TURTLES IS STRIKING ) ALSO CALL ANY RIB WITH PERICHONDRAL OUTGROWTHS A PROTO COSTAL 'AFFNEY f kG  AND IT IS POSSIBLE THAT THIS VERTEBRA IS A AND A RIB WITH PERICHONDRAL AND METAPLASTIC OUTGROWTHS A COSTAL CERVICAL THAT WAS gCAPTUREDu BY THE SHELL /NCE THIS gDORSALIZEDu AND SIMILARLY TERM ANY STILL HYPOTHETICAL DORSAL VERTEBRA WITH VERTEBRA IS REMOVED FROM CONSIDERATION THE BASAL CONDITION OF PERICHONDRAL OUTGROWTHS A PROTO NEURAL AND A DORSAL VERTEBRA TURTLES OVERLAPS WITH THAT FOUND IN /DONTOCHELYS SEMITESTACEA ,I WITH PERICHONDRAL AND METAPLASTIC OUTGROWTHS A NEURAL ET AL  AND % AFRICANUS ,YSON ET AL A IN REGARDS TO THE 0ARSONS AND 7ILLIAMS f ORIGINALLY OBSERVED THAT THE NUMBER OF DORSAL VERTEBRAE AND RIBS 4HE ONLY APPARENT DISPARITY ENDOCHONDRAL RIBS OF MODERN TURTLES ARE SIGNIkCANTLY NARROWER BETWEEN THESE GROUPS PERTAINS TO THE NUMBER OF CERVICAL VERTEBRAE THAN THE OVERLYING METAPLASTIC BONE WHICH IS MOST APPARENT IN ,I ET AL  ,YSON ET AL A AND IT IS NECESSARY TO POSTULATE MANY JUVENILE AND VARIOUS AQUATIC TURTLES WHERE THE DISTAL THE ADDITION OF TWO CERVICAL VERTEBRAE FROM % AFRICANUS TO / PORTIONS OF THE RIBS ARE NOT COVERED BY METAPLASTIC BONE 4HESE SEMITESTACEA AND THE ADDITION OF YET ANOTHER CERVICAL VERTEBRA AND AUTHORS HOWEVER ERRED BY PRESUMING THAT THE BROADENED THE CAPTURE OF THE MOST POSTERIOR CERVICAL BY THE SHELL FROM / ELEMENTS OF % AFRICANUS CONSIST ONLY OF THE RIB AND LACK SEMITESTACEA TO 4ESTUDINATA &IG   ) SPECULATE THAT THE PERICHONDRAL OR METAPLASTIC OUTGROWTHS 4HIS ASSUMPTION DOES ACQUISITION OF ADDITIONAL CERVICAL VERTEBRAE IS LIKELY LINKED TO NOT APPEAR TO BE FALSE AT kRST LOOK BECAUSE THE EXPANDED RIBS OF THE ORIGIN OF THE TURTLE SHELL WHICH DEMANDED THE ACQUISITION OF A MOST AMNIOTES CONSIST OF ENDOCHONDRAL BONE ONLY (OWEVER MORE ELONGATE AND lEXIBLE NECK TO COMPENSATE FOR THE EXTREME ,YSON ET AL A RECENTLY REVEALED THAT THE EXPANDED RIBS OF % STIFFENING OF THE TRUCK 7ERNEBURG ET AL   4HE LOSS OF THE AFRICANUS ACTUALLY CONSIST OF A VERTICALLY ORIENTED ENDOCHONDRAL MOST POSTERIOR COSTAL AT THE BASE OF CROWN 4ESTUDINES IS ALREADY RIB AND HORIZONTALLY EXPANDED PERICHONDRAL OUTGROWTHS OF THE WELL DOCUMENTED BY NUMEROUS PHYLOGENETIC STUDIES EG RIBS ,YSON ET AL A kGS   A CHARACTER PREVIOUSLY THOUGHT 'AFFNEY ET AL f *OYCE   TO BE UNIQUE TO TURTLES 3CHEYER ET AL   !LTHOUGH ,I ET AL  STATE THAT / SEMITESTACEA LACKS COSTALS IT IS APPARENT FROM $ORSAL 2IBS %XPANDED BY 0ERICHONDRAL /UTGROWTHS AND -ETAPLASTIC THE IMAGES ACCOMPANYING THE TYPE DESCRIPTION ,I ET AL  "ONE kGS   THAT THE COSTALS ARE 4 SHAPED IN CROSS SECTION ,YSON 3EELEY f AND 7ATSON f ALREADY HIGHLIGHTED THE PRESENCE OF ET AL   (OWEVER ONLY HISTOLOGICAL ANALYSIS WILL REVEAL EXPANDED RIBS AS A POTENTIAL DIAGNOSTIC SYNAPOMORPHY BETWEEN WHETHER THIS TAXON POSSESSED PROTO COSTALS OR TRUE COSTALS )T IS % AFRICANUS AND TURTLES BUT THIS CHARACTER WAS LATER DISMISSED BY SIMILARLY UNCLEAR IF THE MIDLINE OF / SEMITESTACEA POSSESSED 0ARSONS AND 7ILLIAMS f BECAUSE THE RIBS OF MODERN TURTLES PROTO NEURALS OR TRUE NEURALS /NLY TESTUDINATES ARE THEREFORE APPEAR TO BE LESS EXPANDED THAT THOSE OF % AFRICANUS 4HE LATTER KNOWN TO HAVE TRUE COSTALS AND TRUE NEURALS AS DEkNED HEREIN CONCLUSION HOWEVER WAS MISGUIDED BECAUSE 0ARSONS AND &IG   7ILLIAMS f FAILED TO DISTINGUISH BETWEEN THE RIB PORTION AND 4HE COSTALS OF % AFRICANUS DIFFER PRIMARILY FROM THOSE OF CROWN THE METAPLASTIC OUTGROWTHS OF THE RIBS IN BOTH TAXA TURTLES BY HAVING VERTICALLY ORIENTED RIBS AND BY LACKING SUTURAL !LTHOUGH PALEONTOLOGICAL DATA NOW CLEARLY REJECT THE TRADI CONTACTS ,YSON ET AL  kGS   BUT THIS MORPHOLOGICAL GAP http://doc.rero.ch TIONAL HYPOTHESIS THAT THE COSTALS OF TURTLES ARE THE RESULT OF AN IS BRIDGED BY /DONTOCHELYS SEMITESTACEA WHICH LACKS SUTURAL EVOLUTIONARY FUSION OF THE RIBS AND OVERLYING OSTEODERMS ,I ET AL CONTACTS BETWEEN THE COSTALS ,I ET AL  kG  AND NUMEROUS  THESE BONES ARE NEVERTHELESS COMPOSITES FORMED THROUGH STEM TURTLES THAT POSSESS VERTICALLY ORIENTED RIBS AS WELL 'AFFNEY THE EXPANSION OF THE RIB INTO THE SKIN THROUGH PERICHONDRAL AND f kG  *OYCE ET AL  kG  *OYCE ET AL A kG   4HE METAPLASTIC OUTGROWTHS EG 3CHEYER ET AL   %MBRYOLOGICAL ONLY MAJOR TRANSITIONS WITHIN THIS CHARACTER COMPLEX THEREFORE STUDIES REVEAL THAT THE RIBS GROW INTO THE DERMIS DURING ONTOGENY OCCUR WITHIN THE KNOWN TURTLE STEM LINEAGE &IG   INITIALLY THROUGH AN EXPANSION OF THE RIBfS PERICHONDRAL COLLAR 4HE REALIZATION THAT % AFRICANUS POSSESSES EXPANDED RIBS THAT AND THEN THROUGH THE METAPLASTIC OSSIkCATION OF THE OVERLYING ARE STRUCTURALLY EQUIVALENT WITH THOSE OF TURTLES IS PARTICULARLY (IRASAWA ET AL  kG  OR SURROUNDING 3CHEYER ET AL  SIGNIkCANT BECAUSE IT CORROBORATES PREDICTIONS MADE BY kG  DERMAL TISSUE !CCORDINGLY TO SOME DEkNITIONS THE EMBRYOLOGICAL STUDIES AS TO THE ORIGIN OF THESE STRUCTURES ,YSON PERICHONDRAL AND METAPLASTIC PORTIONS OF THE COSTAL SHOULD BE ET AL A  CONSIDERED TO BE gAPPOSITIONAL BONEu 0ATTERSON f OR g:UWACHSKNOCHENu 3TARCK f AND THE ENTIRE COSTAL COULD 4HE 4URTLE "REATHING !PPARATUS THEREFORE BE REGARDED TO BE ENDOSKELETAL IN NATURE 2IEPPEL AND %XTANT AMNIOTES EXHIBIT A BROAD RANGE OF BREATHING MECHANISMS 2EISZ f  ) kND THIS POINT TO BE SEMANTIC AT BEST AND MISLEADING BUT ALL GROUPS SHOW SOME TYPE OF COSTAL VENTILATION IE BREATHING AT WORST BECAUSE THE CATEGORIZATION OF THE ENTIRE COSTALS AS PURELY THROUGH THE EXPANSION AND CONTRACTION OF THE RIBCAGE  4URTLES ARE ENDOSKELETAL UNNECESSARILY DOWNPLAYS THE ROLE OF THE DERMIS IN THE NOTABLE EXCEPTION TO THE RULE BECAUSE THE FORMATION OF THE THE FORMATION OF THESE BONES ) HEREIN DISTINGUISH CLEARLY BETWEEN SHELL PRECLUDES MOVEMENT BETWEEN THE RIBS )NSTEAD TURTLES RELY THE ORIGINAL ENDOCHONDRAL PORTIONS OF THE RIB VERSUS THE ON A UNIQUE SYSTEM FORMED BY TWO PAIRS OF ANTAGONISTIC

6 ABDOMINAL MUSCLES THE - TRANSVERSUS AND - OBLIQUUS SCAPULA IS NOT LOCATED WITHIN THE RIB CAGE AS OFTEN ERRONEOUSLY ABDOMINIS 'AUNT AND 'ANS f "RAINERD AND /WERKOWICZ REPORTED BUT RATHER ANTERIOR TO THE RIB CAGE ,YSON AND *OYCE   0ALEONTOLOGISTS HAVE PUZZLED OVER THE ORIGIN OF THIS   4HIS ARRANGEMENT IS HARDLY UNIQUE HOWEVER AS IT BROADLY BREATHING MECHANISM 2IEPPEL AND 2EISZ f BUT FEW INSIGHTS OCCURS AMONG BASAL AMNIOTES AND THEREFORE DEMANDS NO SPECIAL HAVE BEEN PRODUCED ON HOW THIS BREATHING SYSTEM MIGHT HAVE EXPLANATION ,YSON AND *OYCE   'IVEN THE ABSENCE OF NECK ORIGINATED AND TAIL ARMOR IN / SEMITESTACEA THEIR PRESENCE IN 0ROGANOCHELYS ! RECENT STUDY BY ,YSON ET AL  DEMONSTRATED THAT THE QUENSTEDTI AND #HINLECHELYS TENERTESTA IS NOW BEST INTERPRETED AS PRESENCE OF INTERCOSTAL MUSCLES CAN BE TRACED IN OSTEOLOGICAL A DERIVED SYNAPOMORPHY OF THESE TWO TAXA ONLY NOT A BASAL SPECIMENS WITH CONkDENCE BY TRACING THE kBERS MUSCLES LEAVE SYMPLESIOMORPHY OF 4ESTUDINATA CONTRA *OYCE ET AL   BEHIND IN BONE AT THEIR INSERTION POINTS USING HISTOLOGICAL &INALLY RECENT INVESTIGATIONS HAVE SHOWN THAT THE HOOKED SECTIONS 4HIS DISCOVERY IS IMPORTANT AS IT ALLOWS THE RECON ELEMENT IN THE PES OF TURTLES IS A COMPOSITE STRUCTURE FORMED BY STRUCTION OF THE BREATHING APPARATUS OF EXTINCT ORGANISMS WITH THE kFTH DISTAL TARSAL AND THE kFTH METATARSAL BUT THE HOMOLOGY OF CONkDENCE ,YSON ET AL  ALSO SHOWED THAT TURTLES ARE THE STRUCTURE WITH SIMILAR HOOKED ELEMENTS IN OTHER REPTILES REMAINS ONLY EXTANT AMNIOTES KNOWN TO HAVE FULLY LOST THEIR INTERCOSTAL UNCLEAR FOR THE MOMENT *OYCE ET AL B  MUSCULATURE AT POST HATCHING STAGE BUT ALSO THAT ALL GROUPS OF EXTANT AMNIOTES WITH BROADENED RIBS EG ANT EATERS LORIS TREE 4(% /2)'). /& 4524,%3 /##522%$ /. ,!.$ SHREWS EXHIBIT REDUCED AMOUNTS OF INTERCOSTAL MUSCULATURE 4HE PURPORTED ECOLOGY OF THE ORIGIN OF TURTLES HAS HISTORICALLY BEEN *ENKINS f  )T THEREFORE SEEMS TO BE UNIVERSALLY TRUE IN AMNIOTES USED TO SUPPORT VARIOUS PHYLOGENETIC SCENARIOS AS A SISTER GROUP THAT THE RIB CAGE IS MOST COMMONLY STIFFENED THROUGH AN RELATIONSHIP WITH VARIOUS TERRESTRIAL GROUPS OF VERTEBRATES IMPLIES A EXPANSION OF THE RIBS WHICH IN RETURN RESULTS IN A REDUCTION OF TERRESTRIAL ORIGIN OF THE GROUP WHEREAS A CLOSE RELATIONSHIP WITH INTERCOSTAL BREATHING AND THE NECESSITY TO EXPLORE ALTERNATIVE AQUATIC GROUPS IMPLIES AN AQUATIC ORIGIN *OYCE AND 'AUTHIER MODES OF BREATHING )N TURTLES THIS CONUNDRUM WAS SOLVED   &IVE LINES OF EVIDENCE HAVE BEEN DEVELOPED IN THE LAST THROUGH THE SPECIALIZATION OF THE ABDOMINAL MUSCLES !S ,YSON DECADE TO SUPPORT THE TERRESTRIAL NATURE OF THE EARLIEST TURTLES IE ET AL  NOTE THESE MUSCLES SYMPLESIOMORPHICALLY ATTACH TO TAXA AT THE BASE OF THE CLADE 4ESTUDINATA  &IRST BASAL TURTLES THE VISCERAL SIDE OF ALL DORSAL RIBS IN OTHER AMNIOTES BUT ARE UNIVERSALLY HAVE A REDUCED PHALANGEAL COUNT EG 'AFFNEY f CONSTRICTED TO THE ANTERIOR AND POSTERIOR DORSAL RIBS IN TURTLES BUT f 3TERLI ET AL  ANDOR SHORT HANDS CHARACTERS THAT NOT THE MIDDLE DORSAL RIBS )NCIDENTALLY THE POINTS OF INSERTION OF CORRELATED WITH TERRESTRIAL HABITAT PREFERENCES TODAY *OYCE AND THE ABDOMINAL MUSCLES CAN ALSO BE TRACED WITH CONkDENCE IN 'AUTHIER   3ECOND MANY BASAL TURTLES KNOWN FROM COMPLETE HISTOLOGICAL SECTIONS OF THE RIBS AND IT IS THEREFORE POSSIBLE TO SKELETONS ARE KNOWN TO HAVE OSTEODERMS EG 'AFFNEY f *OYCE RECONSTRUCT THE ORIGIN OF THE TURTLE BREATHING APPARATUS IN THE ET AL  ANOTHER FEATURE ONLY FOUND AMONG TERRESTRIAL TURTLES FOSSIL RECORD TODAY *OYCE AND 'AUTHIER   4HIRD AT LEAST TWO LINEAGES OF )N A kNAL STEP ,YSON ET AL  ANALYZED THE EXPANDED RIBS OF BASAL TURTLES HAVE TAIL CLUBS EG 'AFFNEY f f WHICH ARE %UNOTOSAURUS AFRICANUS AND ARRIVED AT THE SURPRISING CONCLUSION DEFENSIVE FEATURES THAT CANNOT BE UTILIZED UNDER WATER *OYCE AND http://doc.rero.ch THAT THIS TAXON  LACKS INTERCOSTAL MUSCLES AND  THAT THE INSERTION 'AUTHIER   &OURTH THE SHELL BONE HISTOLOGY OF BASAL TURTLES SITE FOR THE ABDOMINAL MUSCLES ARE RESTRICTED TO THE ANTERIOR AND SHOWS CHARACTERISTICS OTHERWISE TYPICALLY FOUND AMONG MODERN POSTERIOR DORSAL RIBS AND ARE LACKING IN THE MIDDLE DORSAL RIBS !LL TERRESTRIAL TURTLES 3CHEYER AND 3ANDER   !ND kNALLY DESPITE A OSTEOLOGICAL CORRELATES FOR THE UNIQUE TURTLE BREATHING SYSTEM ARE GLOBAL PREPONDERANCE OF MARINE AND FRESHWATER AQUATIC FAUNAS THEREFORE TO BE FOUND IN % AFRICANUS AS WELL 4HIS HIGHLY SURPRISING FROM THE 4RIASSIC BASAL TURTLES ARE UNIVERSALLY FOUND IN TERRESTRIAL RESULT PROVIDES UNUSUALLY STRONG SUPPORT FOR THE PHYLOGENETIC SEDIMENTS AND ASSOCIATED WITH TERRESTRIAL FAUNAS PARTICULARLY RELATIONSHIPS OF % AFRICANUS WITH TURTLES &IG   SAUROPODOMORPH DINOSAURS *OYCE AND 'AUTHIER  *OYCE 4URTLES ARE LINKED TO %UNOTOSAURUS AFRICANUS BY A NUMBER OF ET AL  A  !QUATIC TURTLE ABOUND IN FRESHWATER AQUATIC ADDITIONAL CHARACTERS BUT THESE LACK THE SAME WEIGHT AS THOSE SEDIMENTS GLOBALLY FOLLOWING THE -IDDLE *URASSIC EG 7INGS ET AL OUTLINED ABOVE BECAUSE THEY EITHER ARE NOT EXCLUSIVE TO THESE   4HIS COINCIDES WITH THE ORIGIN OF THE CROWN GROUP $ANILOV TAXA OR REPRESENT REPTILIAN SYMPLESIOMORPHIES OR ARE BASED ON AND 0ARHAM  *OYCE  !NQUETIN  WHICH IS DIFkCULT HOMOLOGY ASSESSMENTS 4HESE INCLUDE AMONG OTHERS THE UNIVERSALLY OPTIMIZED IN PHYLOGENETIC ANALYSES TO HAVE ORIGINATED PRESENCE OF PAIRED GASTRALIA LACKING LATERAL AND MEDIAL ELEMENTS FROM AN AQUATIC ANCESTOR 4HE DERIVED TURTLE STEM LINEAGE IS PRESENT AMONG NUMEROUS PARAREPTILES ,YSON ET AL A THE THEREFORE CLEARLY TERRESTRIAL AND A CLOSE RELATIONSHIP WITH MARINE SYMPLESIOMORPHIC PRESENCE OF CLEITHRA ,YSON ET AL A B NOT SAUROPTERYGIANS CANNOT BE CORROBORATED USING ECOLOGICAL SENSU *OYCE ET AL  CRANIAL TUBERCLES ,YSON ET AL  ALSO ARGUMENTS PRESENT IN A BROAD SAMPLE OF AMNIOTES AND A WIDE TRUNK ,YSON 4HIS CONSENSUS WAS SOMEWHAT CHALLENGED BY /DONTOCHELYS ET AL  ALSO PRESENT IN A BROAD SAMPLE OF AMNIOTES  ! RECENT SEMITESTACEA ,I ET AL  NOTE THAT ALL KNOWN SPECIMENS OF REEVALUATION OF THE SHOULD GIRDLE OF TURTLES REVEALS THAT THE / SEMITESTACEA ORIGINATE FROM SEDIMENTS THAT WERE DEPOSITED IN A

7 SMALL MARINE BASIN SURROUNDED BY LAND FROM THREE SIDES AND HAD POSED A SIMILAR PROBLEM FOR OVER A CENTURY BECAUSE THE STEM KNOWN TO CONTAIN TERRESTRIAL lORAS AND FAUNAS !LL KNOWN LINEAGE OF BIRDS WAS PRIMARILY kLLED BY THE ,ATE *URASSIC TAXON SPECIMENS THEREFORE COULD HAVE EASILY WASHED IN FROM THE !RCHAEOPTERYX LITHOGRAPHICA 4HIS TAXON DISPLAYS ENOUGH NEARBY COASTS ,I ET AL  FURTHERMORE NOTED THAT THE CHARACTERS TO CLEARLY LINK IT WITH CROWN BIRDS EG FEATHERS SHORTENED HANDS OF / SEMITESTACEA ARE EQUIVALENT IN LENGTH TO WINGS FUSED CLAVICLES REDUCED kNGERS BUT ALSO NUMEROUS MODERN TURTLES LIVING IN gSTAGNANT OR SMALL BODIES OF WATER u BUT SYMPLESIOMORPHIES THAT HIGHLIGHT ITS REPTILIAN AFkLIATIONS EG FAILED TO EMPHASIZE THAT THIS CATEGORY OF TURTLES IMPLIES SWAMPY TEETH LONG TAIL GASTRALIA (UXLEY f  ! LACK OF ADDITIONAL FOSSILS FRESHWATER HABITATS NOT COASTAL MARINE WATERS *OYCE AND HOWEVER PROVIDED ENOUGH ROOM FOR AN ACRIMONIOUS DEBATE 'AUTHIER   !LTHOUGH ,I ET AL  AT NO POINT INDICATE REGARDING THE PRECISE PLACEMENT OF BIRDS WITHIN !RCHOSAURIA EG MARINE HABITAT PREFERENCE FOR THIS TAXON THE IMAGE OF / /STROM f -ARTIN ET AL f #HATTERJEE f  4HIS LACK OF SEMITESTACEA AS A MARINE PROTO TURTLE HAS NEVERTHELESS BEEN TRANSITIONAL FOSSILS FURTHERMORE CEMENTED THE IDEA THAT MOST BURNED INTO THE COLLECTIVE MIND OF THE SCIENTIkC COMMUNITY AVIAN CHARACTERS ORIGINATED IN PARALLEL WITH ONE ANOTHER IN DIRECT THROUGH COUNTLESS RECONSTRUCTIONS OF THIS TAXON PORTRAYING / RELATIONSHIP WITH lIGHT EG &EDUCCIA f  SEMITESTACEA AS AN AGILE SWIMMER OF OPEN MARINE WATERS EG /VER THE COURSE OF THE LAST  YEARS AN ASTOUNDING SAMPLE OF "ENTON ET AL   (OWEVER THIS HABITAT PREFERENCE IS HIGHLY INTERMEDIATE FORMS HAS IDENTIkED BIRDS AS HIGHLY DERIVED UNLIKELY FOR THIS TAXON BECAUSE NEITHER ITS STIFFENED TORSO NOR ITS THEROPOD DINOSAURS BEYOND ALL REASONABLE DOUBT :HOU   REDUCED LIMBS AND TAIL WOULD HAVE ALLOWED EFFECTIVE PROPULSION IN !LTHOUGH THE DEBATE CONTINUES REGARDING THE PRECISE INTERRELA MARINE SETTINGS TIONSHIPS OF VARIOUS THEROPOD GROUPS EG 8U ET AL  /NE OBSERVATION ALLOWS ME TO EVEN QUESTION THE FRESHWATER 'ODEFROIT ET AL  THE PRIMARY STEPS WITHIN THIS TRANSITIONAL AQUATIC HABITAT PREFERENCES FOR / SEMITESTACEA ) AGREE WITH ,I SERIES ARE NOW CLEAR /NE OF THE MOST SURPRISING REALIZATIONS OF THIS ET AL  THAT THIS TAXON PLOTS WITH VARIOUS gSWAMPYu TAXA IN RESEARCH PROGRAM HAS BEEN THAT THE MAJORITY OF CHARACTERS THE TERNARY DIAGRAM OF *OYCE AND 'AUTHIER  BUT THIS HISTORICALLY ASSOCIATED WITH BIRDS AND lIGHT EG VARIOUS TYPES OF DIAGRAM IS BASED ON MODERN TURTLES WITH A PHALANGEAL COUNT OF FEATHERS HOLLOW BONES FURCULUM AVIAN BREATHING ORIGINATED      OR LESS 4HE PRIMARY DIFFERENCE BETWEEN TERRESTRIAL ONE BY ONE OVER THE COURSE OF TENS OF MILLIONS OF YEARS DEEP WITHIN TURTLES AND AQUATIC TURTLES IS THE RELATIVE LENGTH OF THE PROXIMAL THE THEROPOD CLADE EG 8U ET AL  3CHACHNER ET AL  PHALANGES BECAUSE TURTLES THAT WALK ON DRY LAND NEED SHORT &OTH ET AL  AND WERE NOT DEVELOPED IN CONCERT WITH lIGHT PHALANGES TO ALLOW ROLLOVER WHEREAS AQUATIC TURTLES REQUIRE 4HIS CONCLUSION AT THE SAME TIME PROVIDES THE MOST CONVINCING ELONGATE PHALANGES FOR THE FORMATION OF EFFECTIVE lIPPERS OR SUPPORT FOR THE THEROPOD ORIGIN OF BIRDS BECAUSE BIRDS ARE LINKED PADDLES *OYCE AND 'AUTHIER   4HE UNGUALS ARE TYPICALLY TO THEROPODS BY A LONG LIST OF SYNAPOMORPHIES THAT WERE ELONGATE IN BOTH GROUPS TO SUPPORT THE CLAWS 5NLIKE ALL PREVIOUSLY THOUGH TO BE UNIQUE TO BIRDS TESTUDINATES / SEMITESTACEA STILL POSSESS A PRIMITIVE PHALANGEAL )N THE OLDER LITERATURE TURTLES WERE VARIOUSLY ALLIED WITH COUNT OF      ,I ET AL  BUT THE PROXIMAL PHALANGES %UNOTOSAURUS AFRICANUS 7ATSON f PLESIOSAURS -OODIE f ARE NOTABLY ROBUST AND SHORTENED AS IN EXTANT TERRESTRIAL TURTLES PLACODONTS *AECKEL f gCOTYLOSAURSu #OPE f TEMNOSPOND http://doc.rero.ch 4HIS TAXON IS THEREFORE PUSHED INTO THE SEMI AQUATIC HABITAT ZONE YLS 6ALL¼N f PAREIASAURS 'REGORY f DIADECTIDS /LSON IN THE TERNARY DIAGRAM OF *OYCE AND 'AUTHIER  BY ITS f AND CAPTORHINOMORPHS #ARROLL f BUT MOST OF THESE PLESIOMORPHIC PHALANGEAL COUNT NOT BECAUSE IT HAS THE LONG HYPOTHESES WERE SUPPORT BY ISOLATED CHARACTERS AND LACKED A PHALANGES CHARACTERISTIC OF AQUATIC TURTLES GLOBAL PERSPECTIVE %ARLY CLADISTIC HYPOTHESES ON TURTLE ORIGINS )N CONCLUSION /DONTOCHELYS SEMITESTACEA WAS LIKELY A FULLY PLACED TURTLES AS SISTER TO CAPTORHINIDS 'AFFNEY AND -C+ENNA TERRESTRIAL STEM TURTLE AND AT MOST AN INHABITANT OF SWAMPY f 'AFFNEY AND -EYLAN f 'AUTHIER ET AL fA BUT LARGER FRESHWATER ENVIRONMENTS 4HIS CONCLUSION DOES NOT HELP RESOLVE COMPUTER ASSISTED ANALYSES HYPOTHESIZED A SISTER GROUP RELATION THE ORIGIN OF TURTLES IN ANY DETAIL AS MOST GROUPS OF AMNIOTES HAVE SHIP TO gANAPSIDu PROCOLOPHONIDS 2EISZ AND ,AURIN f ,AURIN A STRONG TERRESTRIAL SIGNAL IN THE 0ERMIAN AND 4RIASSIC INCLUDING AND 2EISZ f OR PAREIASAURS ,EE f f gPARAPSIDu ARCHOSAURS AND %UNOTOSAURUS AFRICANUS (OWEVER THE POTENTIALLY SAUROPTERYGIANS 2IEPPEL AND DE"RAGGA f DE"RAGA AND FRESHWATER AQUATIC HABITAT PREFERENCES OF / SEMITESTACEA SHOULD 2IEPPEL f OR gPARAPSIDDIAPSIDu LEPIDOSAURS -ÌLLER   NOT TACITLY BE USED AS EVIDENCE IN FAVOR OF RELATIONSHIPS WITH )NTERESTINGLY ALL MODERN ANALYSIS EXCLUDED % AFRICANUS A PRIORI MARINE SAUROPTERYGIANS EVEN THOUGH THIS TAXON HAD CONSISTENTLY BEEN LISTED AS SISTER TO TURTLES IN STANDARD PALEONTOLOGICAL TEXTBOOKS AND CLASSIkCATIONS $)3#533)/. !.$ #/.#,53)/.3 THROUGHOUT MUCH OF THE CENTURY EG (UENE f 2OMER f 4HE ORIGIN OF TURTLES HAS PUZZLED PALEONTOLOGISTS FOR MORE THAN #ARROLL f   YEARS BECAUSE THE FOSSIL RECORD HAD FAILED TO PROVIDE /NE REASON WHY NONE OF THESE HYPOTHESES FULLY CONVINCED THE RECOGNIZABLE INTERMEDIATE FORMS BEYOND THE ,ATE 4RIASSIC SYSTEMATIC COMMUNITY IS BECAUSE TURTLES REMAINED HIGHLY 0ROGANOCHELYS QUENSTEDTI 'AFFNEY f  4HE ORIGIN OF BIRDS APOMORPHIC #ARROLL  AND BECAUSE NONE OF THE CHARACTERS

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11 /SBORN (&  4HE REPTILIAN SUBCLASSES $IAPSIDA AND 3YNAPSIDA AND 3EELEY (  /N A NEW REPTILE FROM 7ELTE 6REDEN "EAUFORT 7EST THE EARLY HISTORY OF THE $IAPTOSAURIA -EM !M -US .AT (IST m %UNOTOSAURUS AFRICANUS 3EELEY  1 * 'EOL 3OC m  3HEDLOCK !- "OTKA #7 :HAO 3 ET AL  0HYLOGENOMICS OF /STROM *(  !RCHAEOPTERYX AND THE ORIGIN OF BIRDS "IOL * ,INN 3OC NONAVIAN REPTILES AND THE STRUCTURE OF THE ANCESTRAL AMNIOTE m GENOME 0ROC .AT !CAD 3CI 53! m 0ARSONS 43 7ILLIAMS %%  4WO *URASSIC TURTLE SKULLS A 3HEN 88 ,IANG $ 7EN *: ET AL  -ULTIPLE GENOME ALIGNMENTS MORPHOLOGICAL STUDY "ULL -US #OMP :OOL m FACILITATE DEVELOPMENT OF .0#, MARKERS A CASE STUDY OF TETRAPOD 0ATTERSON #  #ARTILAGE BONES DERMAL BONES AND MEMBRANE PHYLOGENY FOCUSING ON THE POSITION OF TURTLES -OL "IOL %VOL BONES OR THE EXOSKELETON VERSUS THE ENDOSKELETON )N !NDREWS 3- m -ILES 23 7ALKER !$ EDITORS 0ROBLEMS IN VERTEBRATE EVOLUTION 3TARCK $  %MBRYOLOGIE 3TUTTGART 'EORG 4HIEME ,ONDON !CADEMIC 0RESS P m 3TERLI * DE LA &UENTE -3 'UILLERMO 32 2OUGIER 7  !NATOMY AND 0HILIPPE ( :HOU 9 "RINKMANN ( 2ODRIGUE . $ELSUC &  RELATIONSHIPS OF 0ALAEOCHERSIS TALAMPAYENSIS A ,ATE 4RIASSIC TURTLE (ETEROTACHY AND LONG BRANCH ATTRACTION IN PHYLOGENETICS "-# FROM !RGENTINA 0ALAEONTOGR ! m %VOL "IOL  3TERLI * 0OL $ ,AURIN -  )NCORPORATING PHYLOGENETIC UNCERTAINTY 0HILIPPE ( "RINKMANN ( ,AVROV $6 ET AL  2ESOLVING DIFkCULT ON PHYLOGENY BASED PALAEONTOLOGICAL DATING AND THE TIMING OF PHYLOGENETIC QUESTIONS WHY MORE SEQUENCES ARE NOT ENOUGH 0,O3 TURTLE DIVERSIkCATION #LADISTICS m "IOL E 3TRIMMER + VON (AESELER !  1UARTET 0UZZLING A QUARTET 0LATZ *% #ONLON *-  wAND TURN BACK AGAIN .ATURE  MAXIMUM LIKELIHOOD METHOD FOR RECONSTRUCTING TREE TOPOLOGIES 0OLLOCK $$ %ISEN *! $OGGETT .! ET AL  ! CASE FOR EVOLUTIONARY -OL "IOL %VOL m GENOMICS AND THE COMPREHENSIVE EXAMINATION OF SEQUENCE 4HOMSON 2# 0LACHETZKI $# -AHLER $, -OORE "2  ! CRITICAL BIODIVERSITY -OL "IOL %VOL m APPRAISAL OF THE USE OF MICRO2.! DATA IN PHYLOGENETICS 0ROC .ATL 2ANNALA " (UELSENBECK *0 9ANG : ET AL  4AXON SAMPLING AND !CAD 3CI %m% THE ACCURACY OF LARGE PHYLOGENIES 3YST "IOL m 4SUJI ,! -ÌLLER *  !SSEMBLING THE HISTORY OF THE 0ARAREPTILIA 2EISZ 22 ,AURIN -  /WENETTA AND THE ORIGIN OF TURTLES .ATURE PHYLOGENY DIVERSIkCATION AND A NEW DEkNITION OF THE CLADE &OSSIL m 2ECORD m 2EST *3 !ST *# !USTIN ## ET AL  -OLECULAR SYSTEMATICS OF 4SUJI ,! -ÌLLER * 2EISZ 22  -ICROLETER MCKINZIEORUM GEN ET SP PRIMARY REPTILIAN LINEAGES AND THE TUATARA MITOCHONDRIAL GENOME NOV FROM THE ,OWER 0ERMIAN OF /KLAHOMA THE BASALMOST -OL 0HYLOGENET %VOL m PARAREPTILE FROM ,AURASIA * 3YST 0ALEONTOL m 2IEPPEL / A 4URTLES AS DIAPSID REPTILES :OOL 3CR m 4SUJI ,! -ÌLLER * 2EISZ 22  !NATOMY OF %MEROLETER LEVIS AND THE 2IEPPEL / B 0ARAPLACODUS AND THE PHYLOGENY OF THE 0LACODONTIA PHYLOGENY OF THE NYCTEROLETER PARAREPTILES * 6ERT 0ALEONTOL m 2EPTILIA 3AUROPTERYGIA  :OOL * ,INN 3OC m  2IEPPEL /  4URTLES AS HOPEFUL MONSTERS "IO%SSAYS m 4ZIKA !# (ELAERS 2 3CHRAMM ' ET AL  2EPTILIAN TRANSCRIPTOME 2IEPPEL / DE"RAGGA -  4URTLES AS DIAPSID REPTILES .ATURE V A GLIMPSE IN THE BRAIN TRANSCRIPTOME OF kVE DIVERGENT http://doc.rero.ch m 3AUROPSIDA LINEAGES AND THE PHYLOGENETIC POSITION OF TURTLES 2IEPPEL / 2EISZ 22  4HE ORIGIN AND EVOLUTION OF TURTLES !NNU %VO$EVO  2EV %COL 3YST m 6ALLEN %  "EITRAGE ZUR +ENNTNIS DER /NTOGENIE UND DER 2OMER !3  /STEOLOGY OF THE REPTILES #HICAGO 5NIVERSITY OF VERGLEICHENDEN !NATOMIE DES 3CHILDKROTENPANZERS !CT :OOL m #HICAGO 0RESS  3ALICHOS , 2OKAS !  )NFERRING ANCIENT DIVERGENCES REQUIRES 7ANG : 0ASCUAL !NAYA * :ADISSA ! ET AL  4HE DRAFT GENOMES OF GENES WITH STRONG PHYLOGENETIC SIGNALS .ATURE m SOFT SHELL TURTLE AND GREEN SEA TURTLE YIELD INSIGHTS INTO THE 3CHACHNER %2 ,YSON 42 $ODSON 0  %VOLUTION OF THE RESPIRATORY DEVELOPMENT AND EVOLUTION OF THE TURTLE SPECIkC BODY PLAN .AT SYSTEM IN NONAVIAN THEROPODS EVIDENCE FROM RIB AND VERTEBRAL 'ENET m MORPHOLOGY !NAT 2EC m 7ATSON $-3  %UNOTOSAURUS AFRICANUS 3EELEY AND THE ANCESTRY 3HAFFER (" -INX 0 7ARREN $% ET AL  4HE WESTERN PAINTED TURTLE OF THE #HELONIA 0ROC :OOL 3OC ,OND m GENOME A MODEL FOR THE EVOLUTION OF EXTREME PHYSIOLOGICAL 7ERNEBURG ) (INZ *+ 'UMPENBERGER - ET AL  -ODELING NECK ADAPTATIONS IN A SLOWLY EVOLVING LINEAGE 'ENOME "IOL 2 MOBILITY IN FOSSIL TURTLES * %XP "IOL " -OL $EV %VOL $/)JEZ 3CHEYER 4- 3ANDER 0-  3HELL BONE HISTOLOGY INDICATES B  TERRESTRIAL PALAEOECOLOGY OF BASAL TURTLES 0ROC "IOL 3OC " 7ESTPHAL &  4HE DERMAL ARMOUR OF SOME 4RIASSIC PLACODONT m REPTILES )N "ELLAIRS ! #OX #" EDITORS -ORPHOLOGY AND BIOLOGY OF 3CHEYER 4- "RÌLLMANN " 3¶NCHEZ 6ILLAGRA -2  4HE ONTOGENY REPTILES ,ONDON !CADEMIC 0RESS P m OF THE SHELL IN SIDE NECKED TURTLES WITH EMPHASIS ON THE 7ILLISTON 37  4HE PHYLOGENY AND CLASSIkCATION OF REPTILES HOMOLOGIES OF COSTAL AND NEURAL BONES * -ORPHOL m #ONTRIB 7ALKER -US m

12 7INGS / 2ABI - 3CHNEIDER *7 ET AL  !N ENORMOUS *URASSIC :ANGERL 2  4HE HOMOLOGY OF THE SHELL ELEMENTS IN TURTLES * TURTLE BONE BED FROM THE 4URPAN "ASIN OF 8INJIANG #HINA -ORPH m .ATURWISSENSCHAFTEN m :ARDOYA 2 -EYER !  #OMPLETE MITOCHONDRIAL GENOME SUGGESTS 8U 8 :HOU :( 0RUM 2  "RANCHED INTEGUMENTAL STRUCTURES IN DIAPSID AFkNITIES OF TURTLES 0ROC .ATL !CAD 3CI m 3INORNITHOSAURUS AND THE ORIGIN OF FEATHERS .ATURE m :HOU :  4HE ORIGIN AND EARLY EVOLUTION OF BIRDS DISCOVERIES 8U 8 9OU ( (AN &  !N !RCHAEOPTERYX LIKE THEROPOD FROM #HINA DISPUTES AND PERSPECTIVES FROM FOSSIL EVIDENCE .ATURWISSENSCHAF AND THE ORIGIN OF !VIALAE .ATURE m TEN m http://doc.rero.ch

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