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Historical Biology An International Journal of Paleobiology Publication details, including instructions for authors and subscription information: http://www.informaworld.com/smpp/title~content=t713717695 Osteology and phylogeny of a new species of Araripesuchus (Crocodyliformes: Mesoeucrocodylia) from the Late Cretaceous of Madagascar Alan H. Turner a a Department of Geoscience, University of Iowa, Iowa City, IA, USA

Online Publication Date: 01 January 2006 To cite this Article: Turner, Alan H. (2006) 'Osteology and phylogeny of a new species of Araripesuchus (Crocodyliformes: Mesoeucrocodylia) from the Late Cretaceous of Madagascar', Historical Biology, 18:3, 255 - 369 To link to this article: DOI: 10.1080/08912960500516112 URL: http://dx.doi.org/10.1080/08912960500516112

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MACN Michigan Aires, China; Buenos of of Naturales, Republic Pakistan-University Produc¸ao Mineral, People’s of da Beijing, Paleoanthropology, Nacional Survey and Departmento Geological Paleontology do Vertebrate GSP-UM, Geologia Brazil; e Janeiro, Mineralogia de de Rio Divisao GDM-DNPM, Chicago; History, Natural Abbreviations: Institutional uem utn A nvriyo naaaio Madagascar Antananarivo, of University UA, Austin; Museum, u sdsigihdfo hmb h rsneo oratpmrhe.Adtoal,tenwfr ak ubro derived of number a lacks form new the Additionally, autapomorphies. four of in presence the present by features them from distinguished is but Keywords: n ersuis hsgopi eitda oecoeyrltdt escin hnt oouhas When notosuchians. to than neosuchians to related closely more as depicted crocodyliform Malagasy is the wegeneri group with clade This a peirosaurids. forms and and synapomorphies unambiguous five by diagnosed iatcltdps-rna lmns hslresml,culdwt Tsas losadtie ecito fits of description detailed a allows CT-scans, with with as characters well coupled derived as of remains, sample, number post-cranial a associated large shares and form skulls This new partial The three elements. morphology. animal, complete post-cranial nearly a disarticulated including individuals, five least orsodne .H unr iiino aenooy mrcnMsu fNtrlHsoy eta akWs t7t tet e York, New Street, 79th at West Park Central [email protected] History, E-mail: Natural of USA. Museum 10024, American Paleontology, NY of Division Turner, H. A. Correspondence: of species new A Abstract USA 52242, IA City, Iowa Hall, Trowbridge 121 Iowa, of University Geoscience, of Department TURNER H. ALAN Cretaceous Madagascar Late of of the species from new Mesoeucrocodylia) a (Crocodyliformes: of phylogeny and Osteology Biology Historical hlgntcrltosisof relationships Phylogenetic sicue nteaayi,i srcvrda ebro the of member a as recovered is it analysis, the in included is rcdlfre,osteology, Crocodyliformes, Araripesuchus 06 83:255–369 18(3): 2006; , .gomesii A. ´ Paleontologı y a and rsn nteMeaaoFraino aaacri ecie.Tetxni nw rmat from known is taxon The described. is Madagascar of Formation Maevarano the in present Araripesuchus .patagonicus A. MH mrcnMsu fNtrlHsoy e ok MH il uemof Museum Field FMNH, York; New History, Natural of Museum American AMNH, ´ el nvria ainldlCmhe Neuque Comahue, del Nacional Universidad la de a q 06Tyo Francis & Taylor 2006 Araripesuchus r vlae sn asmn nlssicuig2 eourcdla taxa. mesoeucrocodylian 29 including analysis parsimony a using evaluated are . aeCeaeu,Mdgsa,Meaaoformation Maevarano Madagascar, Cretaceous, Late , arn ta.20,CryRgr 02 Curry- 2000, 2002, O’Connor Curry-Rogers 2004). and Forster 2002, and Forster Sampson al. Rogers 1998, 2001, et 1996, 1998, Carrano al. and al. including et theropods, et (Forster dinosaurian saurischian, birds abelisaurid several described sauropods, entirely Buckley the armored and is point, Whatley this O’Connor component 2002, and To al. 1999, Forster 2004). et Brochu 2003, Carrano and 2000, 2000, Buckley al. 1998, et 1998, Buckley al. al. et et Gottfried 1998, Sampson Krause and Asher 1997, .wegeneri A. Araripesuchus ´ ainld ie,Nae,Rpbi fNgr MUC Niger; of Republic Niamey, Niger, de National e rrpscu gomesii Araripesuchus savldseisof species valid a as Araripesuchus .gomesii A. ld.Atog,ispsto ntecaeis clade the in position its Although, clade. ´ ,Agnia M,TxsMemorial Texas TMM, Argentina; n, þ .patagonicus A. and Araripesuchus rrpscu patagonicus Araripesuchus aaagscu insignis Mahajangasuchus . Araripesuchus . Araripesuchus is , Downloaded By: [Columbia University] At: 05:34 17 September 2007 97.Teeicuelretx,sc as and such 2002) taxa, al. large et champsa (Buckley include taxa These separate 1997). seven least at including hrceitc swl stertnino some of retention the taxa, not American as traits autapomorphic well of plesiomorphic number as a on characteristics based is conclusion 256 patagonicus ut fdrvdfaue rsn ntesmall- the in present as features such derived taxon forms of bodied show smaller that suite remains crocodyliform as a are beds well these from as 2003), clarki (Buckley Simosuchus form al. long-snouted a et 2001), 1999, Brochu Apesteguı ersnsanwseisof species it new as here a described is 1997, represents and al. 1999) et Brochu (Buckley and Buckley authors previous by referenced obsrgrigtemmesi of membership the regarding doubts Neosuchia. to group sister the is that clade a in it places aspects and clarifies of taxon taxonomic Malagasy the new its from of information Inclusion present unclear. remains status at phylogenetic unquestioned, is Mesoeucrocodylia 2003, Norell in and Pol membership 1999, 2000, its al. al. Although Pol et 2004). et Ortega 1999, 2000, Wu al. Brochu 1994, et of Buckley and Clark number 1991, Buckley Gasparini a al. 1997, et to, 1959, Gasparini sister (Price 1971, or groups within, mesoeucrocodylian placed been has h rcdlfr opnn seteeydiverse, extremely is component crocodyliform The diinly ubro uhr aeraised have authors of number a Additionally, ic h ao a rtdescribed, first was taxon the Since Araripesuchus iue1 G-NM432-R, DGM-DNPM 1. Figure .H Turner H. A. ´ BfeatadTqe 99 Rasmusson 1979, Taquet and (Buffetaut 2005). a and aaagscu insignis Mahajangasuchus Araripesuchus opooyadpyoeei analysis phylogenetic and morphology rrpscu gomesii Araripesuchus Bclye l 00.As recovered Also 2000). al. et (Buckley rrpscu buitreraensis Araripesuchus hsmtra a been has material This . xiie nteSouth the in exhibited rrpscu gomesii Araripesuchus Araripesuchus , Bclyand (Buckley Araripesuchus Araripesuchus Araripesuchus Pland (Pol Tremato- .This ieitrrtto fcailstrs ,Dra iw ,Vnrlview. Ventral B, view. Dorsal A, sutures. cranial of interpretation Line . ecie yBfeat(91,wt h South the with of (1981), species Buffetaut American The by al. 2002). of et Broin described de Ortega status Lapparent 1994, contaxonomic and Kellner Prasad 1990, 2000, al. et (Michard eh 91.Teseie speevdo w slabs two on preserved is 24450; specimen (AMNH The formation 1991). Hecht same the from recovered oto h uue ntesotadpeevdskull preserved and discerned. snout and be the can preserved on table well sutures is the of region is most palatal left The mandible the supra- side. to displaced complete lateral right but articulation, and nearly near in orbit A preserved and left fenestra. table the skull temporal to poorly the but posterior of complete, much occiput nearly lacking Price a skull by of preserved, described consists initially It was (1959). Brazil Santana of (Albian) Cretaceous Formation Early the from 1) Figure h ooyeof holotype The gomesii to Araripesuchus referred species taxon. the other of members the on thorough Araripesuchus a comments discusses it provide Rather, and taxa. to these of intended each of not redescription is section This Araripesuchus taxa. mesoeucrocodylian other by tested hypothesis including this not test will were study This however, phylogenetically. hypotheses, These tioned. nams opeeseeo of skeleton complete almost An Araripesuchus .wegeneri A. n eiw u nesadn fthese of understanding our reviews and ytmtc n eiwo membership of review and systematics .gomesii A. pce n ra apeof sample broad a and species ntepyoeei nlsswith analysis phylogenetic the in Araripesuchus rc 1959 Price rrpscu wegeneri Araripesuchus DMDP 432-R; (DGM-DNPM .gomesii A. a enques- been has a later was , Downloaded By: [Columbia University] At: 05:34 17 September 2007 h ooye(U V29 iue ,3 consists 3) 2, Figures 269; PV (MUC (2000). holotype al. The et Ortega by described Argentina Province, aeu Aba)RoLmyFraino Neuque of Formation Limay Cre- containing Rio Early block the (Albian) from single taceous specimens a individual four from least known at is taxon This rrpscu patagonicus Araripesuchus specimen this warranted. of is discussion and and detailed description a brief specimen thorough the a of however, completeness sides and the (1991), given Hecht photographed by both published detail. been was from description has and specimen prepared preservation AMNH The acid extraordinary aspects. been ventral revealing has and dorsal 24450 both showing iue3 UP 269, MUCPV 3. Figure iue2 UP 269, MUCPV 2. Figure rrpscu patagonicus Araripesuchus Ortega rrpscu patagonicus Araripesuchus tal et 2000 . eie ieitrrtto fcailstrs ,Dra iw ,Vnrlview. Ventral B, view. Dorsal A, sutures. cranial of interpretation line Revised . ´ kl ndra iw()advnrlve B.Scale (B). view ventral and (A) view dorsal in Skull . n h rfotlde o xadltrlyt any to a laterally as expand not prefrontal frontal. does the the to prefrontal parallel described runs The that 61) Ortega few band specimen. bony (2000: a “narrow, type of al. the in interpretation et present original features the skull with disagree 283) and hindlimb 270, as well elements. 268b, referred post-cranial as other 268, skull surface other additional 267, ventral an The comprise PV palate. the (MUC the and was specimens expose skull block to The the prepared scapulae. from obscures and removed armor preserves 269 forearms dorsal PV MUC the articulated series. vertebral and the of type details the is the in rostrum of the half missing of anterior end the anterior-most and The skull postcranium. complete nearly a of a h potnt oeaieti aeiland material this examine to opportunity the had I New Araripesuchus rmMadagascar from ¼ cm. 1 257 Downloaded By: [Columbia University] At: 05:34 17 September 2007 aea iw ,Sm mg ihln nepeainsprmoe pnit. upon superimposed interpretation line with image Same B, view. lateral in in patagonicus condition the bone, A. the nasal of is the edge prefrontal contacting anteromedial-most and prefrontal the lacrimal only the With of similar. surface dorsal the of this corroborating the of extent conclusion. contact full the other preserves prefrontal/lacrimal The 267) prefrontal/lacri- PV mesoeucrocodylians. the (MUC other a corresponds of skull in location (high- be in-filling) suture the mal feature to sediment to closely here linear slight more interpreted prominent by is A lighted orbit, crack. medial illus- the the narrow with lacrimal, of inline The nearly and border dorsally, interpretation. lying prefrontal 63: as (p. the trated authors’ skull between the the of suture border illustrates drawing anterior line 5) the The fig. of orbits”. level the the of at degree significant 267, MUCPV 4. Figure 258 hsseisof species This wegeneri Araripesuchus other in the is it that as interpretation subdermal and Araripesuchus my inset is is bar of It postorbital border bar. anteroventral low, postorbital a the holotype, the marks the in ridge the fact, distinct, In margin jugal. skulls, but lateral the is of the both body from jugal the of inset In the appears to unable bar observation. of was postorbital I the body this edge”. anterior main of its corroborate on the least insertion at in dermal, the bar 24450, postorbital AMNH Ortega of specimen in holotype note jugal 61) the (p. the “unlike authors The that of drawing. 5) body fig. the (2000: al.’s et with flush as shown rsrigrgtjgladmc ftepalate; the of much and rostrum jugal complete nearly right (a preserving 700 GDF Niger. holotype Gadoufaoua, The of Formation Elrhaz Cretaceous ie hsnwitrrtto Fgr ) h size the 4), (Figure interpretation new this Given h o-iepsobtlbrof bar postorbital rod-like The .gomesii A. .H Turner H. A. . taxa. Araripesuchus osntdfe ral rmthat from greatly differ not does rrpscu patagonicus Araripesuchus uftu 1981 Buffetaut rrpscu gomesii Araripesuchus skonfo h Early the from known is .patagonicus A. eainhpbtenperna,lcia,mxla n aa oe.A kl nright in Skull A, bones. nasal and maxilla, lacrimal, prefrontal, between Relationship . and is oiina h other the mediolateral same the as and roughly position in dentary, verticalization posterior the as of flattening 700) well anterior as (GDF the of dentary, holotype flattening dorsoventral same The the shows correct. is lamina”. dentary the completely posterior of surface not flat lateral the and of description vertical This “deep, a having fho it ailr ot.Tedfeec between difference patagonicus The tooth. A. maxillary sixth or fifth nepeaino oeo h etrs reae al. et of other dentary Ortega the features. from characterizes the 69) it of (2000: some distinguish on interpretation to reputed Araripesuchus features be of not of should species taxon a this that considered argued have (2000) al. et an as biogeographic but more description. of a thorough from lacking, benefit is therefore, is should, and it importance taxon mesoeucrocodylian, this African of published A study taxon. Taquet this detailed to and pertain may that Buffetaut and bones scutes, limb and dermal vertebrae, described isolated mention (1979) been yet not 18) lr 18)as oe httoweathered two for that exist noted braincases also (1986) Buffetaut by Clark description brief (1981). a given was 6) 5, Figures rsino h etmniua au n poor and ramus 700. GDF in mandibular ramus com- right left a the of dorsoventral the from preservation slight of stem to to pression due appears misinterpretation description of 6B). authors’ 5B, splenial (Figures The width The mandibular ventral the mandible. of half the wegeneri of A. surface ventral wegeneri A. the of compaction GDF dorsoventral of slight taxon. preservation Patagonian the poor and the 700 to due pronounced aigetra ae Pie15,Hct1991, Hecht 1959, (Price nares external facing ihr ta.(90,Klnr(94 n Ortega and (1994) Kellner (1990), al. et Michard te pce of species Other of splenial the that note 69) (2000: al. et Ortega oee,de o omaymr than more any form not does however, , om oeta afo h it fthe of width the of half than more forms pce.Idsge,hwvr ihtheir with however, disagree, I species. and Araripesuchus rrpscu wegeneri Araripesuchus Araripesuchus .wegeneri A. Araripesuchus aeanterolaterally have aaaon the taxa—around sprasover perhaps is iiganumber a citing , .wegeneri A. hthave that as Downloaded By: [Columbia University] At: 05:34 17 September 2007 .Brochu.) C. nGF70 h te etrssi odistinguish to said features wegeneri other the A. The in confirm 700. nares GDF to external on unable directed was wegeneri frontally A. I the of 2000). presence al. et Ortega 700, GDF 5. Figure rsneo v rmxlaytehin teeth premaxillary five of presence rcdlmrh n v rmxlaytehare teeth premaxillary among common five are and maxillary teeth posterior crocodylomorphs premaxillary the on Five denticles teeth. of presence the iue6 D 700, GDF 6. Figure teatro i ftersrmi eroded is rostrum the of tip anterior —the rmother from rrpscu wegeneri Araripesuchus Araripesuchus rrpscu wegeneri Araripesuchus kl ndra iw()advnrlve B.Scale (B). view ventral and (A) view dorsal in Skull . .wegeneri A. pce was species ieitrrtto fcailstrs ,Dra iw ,Vnrlview. Ventral B, view. Dorsal A, sutures. cranial of interpretation Line . and euhsbuitreraensis pesuchus .wegeneri distinguishing A. feature unique unambiguously only the rsn nteMalagasy the in present aeonwr eerdt “aff. to from referred teeth isolated were feature, this Cameroon on Based 7). (Figure hssuya ela h el described newly the as well as study this Bue ta.19) rsdadLpaetd Broin de Lapparent and Prasad 1990). al. et (Brunet hslae etcltdpseirmxlaytehas teeth maxillary posterior denticulated leaves This New Araripesuchus rmother from PladApesteguı and (Pol ¼ Araripesuchus m Poorpscuts of courtesy (Photographs cm. 1 rmMadagascar from Araripesuchus Araripesuchus ´ ecie in described 2005). a species Arari- 259 ” Downloaded By: [Columbia University] At: 05:34 17 September 2007 etto in dentition BclyadBoh 1999), Iberosuchus Brochu and (Buckley Eremosuchus W n us1996a), Sues and (Wu 1977), Archer and (Hecht pristicham mekosuckines one e.g. with taxa affinities Calvo phylogenetic another, crocodyliform close and show numerous not in Turner do that found Berg 1977, of is Archer 1956, and be 2005) and (Langston to Hecht value proven 1966, has phylogenetic serrations) limited with (laterally morphology compressed dentary tooth 2002). Larsson ziphodont left and Furthermore, Sues fragmentary see a though 1994, from (Buffetaut known currently is ifrn ot opooy ontseany be see character. should autapomorphic any character other not than significant a phylogenetically and do such more distinct considered for that I a reason grounds morphology, is compelling sufficient it tooth not Although, different other is taxon. such the to renaming as respect and with species, species, the aaauhswegeneri Hamadasuchus renamed provisionally similarity, 20)ntdtesmlrt ewe h et of teeth the between similarity wegeneri the A. noted (2002) 700, GDF 7. Figure et hwn etce ntohmri.(mg rmthe from (Image Broin.) margin. de Lapparent tooth de F. of on courtesy Serrette denticles photograph showing teeth 260 ti yve httepeec fziphodont of presence the that view my is It n hscnlso suspect. conclusion this find I .H Turner H. A. Atns1975). (Antunes n “aff. and .wegeneri A. Bfeat1989), (Buffetaut rrpscu wegeneri Araripesuchus Araripesuchus . ssml natpmrh of autapomorphy an simply is sns(agtn1956), (Langston psines Sebecus Itasuchus aaauhsrebouli Hamadasuchus n,bsdo this on based and, ” .wegeneri A. ls-po maxillary of Close-up . Mahajangasuchus Clet1946), (Colbert Pie1955), (Price Araripesuchus Platyognathus saff. as eainhp of Relationships nobtlfnsr mle hnteorbit. an the and than choana, smaller internal fenestra sized antorbital a moderate forming a maxilla slit, with premaxilla narrow synapomorphies the of constricted contact Other rostrum the a teeth at row. are relationship smaller this tooth supporting opposite maxillary enlarged in contact premaxilla– opposite tooth the maxilla to and edge points; posterior with these teeth row, at dentary tooth outward fenestra; of extending maxilla single middle or of the a in with dorsolaterally enlarged quadrate the teeth nares of a any, oriented; external if dorsally little, bar; of forms that internarial number premaxilla a a include: characters by derived These corroborated paraphyletic). apomorphies. unambiguous is sense, relationship traditional This the in Notosuchia, diin aprn 17)ntdsmlrte between Araripesuchus similarities noted (1971) Gasparini addition, ic t nta description, initial its Since mesoeucrocodylians nesodta lotalo hs hrcesare characters these now for plesiomorphic of and is crocodyliforms all it among widespread almost course, Of that armor). understood dermal and amphicoelous vertebrae, wrinkles, of irregular sculpture to specialized, tend bones and cranial number in denti- reduced symphysis, the tion in participate splenials mandibular external fenestra, large a supraoccipital roof, cranial in fenestra, participates supratemporal not supratemporal in palatine, may or a may participate frontal and orbits, fenestra, the than pterygoid antorbital smaller fenestra an by of divided, formed presence or nares single nares internal bar external postorbital subdermal, quadrate, slightly inclined high vertical a slightly in rather and a large skull are position, that orbits size, snout, short medium a with to (small acteristics oouhde oigissmlrte with terrestris similarities its noting Notosuchidae, em,wsntn itnto from distinction a it noting (this, fenestra was maxillo-palatal a seems, terminating of nares lack external the given an dorsally, were: Characters grouping this Uruguaysuchidae. for the in together n lr 98 placed 1988) Clark and analyses. cladistic hypertrophied on early based these and were of larger maxilla, suggestions None 1971). the maxilla, (Gasparini teeth in and maxillary teeth premaxilla of the number a having between fenestra), notch maxillo-palatal a possesses which guaysuchus obnto faprnl rmtv n derived first and (1959) described Price primitive When 1991). apparently (Hecht characteristics of combination a preserves it that given phylogenetically place to difficult eae onouhasthan neosuchians to related sbre”Ntsci oinclude to Notosuchia “suborder” al ldsi nlss(lr 96 94 Benton 1994, 1986, (Clark analyses cladistic Early olwn hs aprn 17)eetdthe erected (1971) Gasparini this, Following Araripesuchus hsntdsmlrt nldd1 char- 18 included similarity noted This . rrpscu gomesii Araripesuchus , Sphagesaurus and Araripesuchus and Uruguaysuchus Notosuchus Araripesuchus Araripesuchus and oother to eicue tin it included he , Notosuchus Cak1986). (Clark Araripesuchus Notosuchus nldn them including smr closely more as Notosuchus a proven has Notosuchus (making , Uru- .In , Downloaded By: [Columbia University] At: 05:34 17 September 2007 aae ieadtxnsmln.Bscly two Basically, for been sampling. have in proposed mesoeucrocodylians taxon differences basal within large and positions sometimes size the to dataset due likely are oantscincae(o n oel20,Pland Pol 2003, Norell and Apesteguı (Pol taxon clade sister notosuchian the as a or to 2003), notosuchian (Pol a Neosuchia with and clade polytomy unresolved an in 2001), polytomy unresolved an with in 2000), mesoeucrocodylians 2004, Gado basal and polytomy Turner (Larsson other unresolved and 2000, an Neosuchia in al. with 2005), Calvo et and Buckley Turner 1999, Brochu placed rcue l 02 unr20,Tre n Calvo and Turner 2004, Turner 2000, 2002, and al. al. et Buckley Ortega et 2000, Brochu 1988, al. et Clark Buckley 1999, Brochu and to (Benton historical sister placement—i.e. the Neosuchia systematic to its of correspond perceptions and found consistently Sebecosuchia h itrtxno h ld Peirosauridae clade the of taxon Mahajangasuchus sister the osnu mn hlgntcaaye.I h most the analyses, In recent analyses. phylogenetic of lack among a consensus given difficult proves question this Answering aa eourcdlas naaye hthv a have that for analyses position resolved In mesoeucrocodylians. basal eoeteArcntxnfrom taxon African the remove nlss oeaaye depict analyses Some analysis. 94 reae l 00,a h itrto sister the as 2000), al. (Clark Neosuchia et to Libycosuchus sister Ortega the as 1994, 1991), al. et parini rmatxn vnmr owieohrapcsof Is aspects other while so more ignored. are even species morphology a taxon, prefrontal/ exclude higher to a the used of from be of cannot indicative shape and contact less the nasal morphology, than or relationships dental more level no distinct is Broin a ziphodonty de Although, Lapparent and dental 2002). on (Prasad based or alone 2000) morphology al. et (Ortega phenetic been erprsdt s,“Is ask, to rephrased perhaps 2004). be should Turner monophyly 2003, of question Norell this and Therefore, Pol 2003, Pol 2000, relationship sister the about between doubt no been has There oouhasta escin W ta.1997, Apesteguı and al. Pol et 2003, Norell (Wu and neosuchians Pol than notosuchians osdrdaseisof species a considered Is and Turner 2000, al. et Buckley 2005). Calvo 2000, al. 1994, et (Clark Ortega notosuchians than neosuchians to closer nsbeun ok hlgntcaaye have analyses phylogenetic work, subsequent In Araripesuchus Araripesuchus Notosuchus Araripesuchus .gomesii A. ´ 05.Dfeecsbtenteeanalyses these between Differences 2005). a þ þ Araripesuchus Notosuchus n te oouhas(eeoe al. et (Sereno notosuchians other and Trematochampsa notosuchian? a monophyletic? Araripesuchus þ and Trematochampsa Araripesuchus .patagonicus A. stesse ao of taxon sister the as Araripesuchus W ta.19) as 1997), al. et (Wu ae nphylogenetic on based .wegeneri A. Araripesuchus a one among bounced has þ Araripesuchus w oain are locations two , Caririsuchus Bclyand (Buckley Otg tal. et (Ortega ”Efrsto Efforts ?” ´ 05 or 2005) a properly lsrto closer (Gas- have þ Notosuchus rcu19,Bclye l 00.Ti character, This in 2000). present anterior and also al. is Buckley and et however, 1994, Buckley medial Clark 1999, in the Brochu 12-1 along (character nasals border the contact Malawisuchus uhasrmiseuvcl u ubro char- of number a grouping. this but support acters equivocal, remains suchians 00.Ti etr speeti te croco- other in present is (e.g. feature dylomorphs al. et This Buckley 1999, 2000). Brochu and Buckley 1994, Clark l notespaeprlfnsrein fenestrae supratemporal the at not into or slightly all only extends frontal The 1986). Clark escin a encsayt eov hs issues. these resolve to non- necessary other be and may neosuchians notosuchians among sampling taxon 05 rsse ontscin W ta.19,Pol 1997, Apesteguı al. and Pol et (Wu 2003, Norell notosuchians and to sister or 2005) aeo h rw caatr171i uke tal. et Buckley addition, in 117-1 In (character 2000). crown the of base Malawisuchus Anatosuchus deo h otria.O h post-cranium, the On postorbital. the Araripesuchus on edge Araripesuchus 93 o 2005). Pol (Rusconi femur the 1933, of edge anterolateral the on flange a hsfauewith feature this h neirpr ny lr 18)as oe that noted and also orbit (1986) on Clark the only. sculpturing Baurusuchus between part with anterior narrow, bar the is the fossa that supratemporal feature the 96.Wt h icvr fteMalagasy the Gasparini of 1993, discovery Rage pesuchus the and With two (Buffetaut Buffetaut 1996). these Cretaceous 1982, between Early link 1981, the faunal during close as landmasses used a been for has and evidence America, South and Africa both biogeo- Araripesuchus the noted of first importance graphic (1979) Taquet and Buffetaut history Biogeographic 2000). al. et Buckley 1999, Brochu and (Buckley oacs h nenlcailmrhlg fthe of morphology cranial internal Malagasy the access To analysis tomographic Computed Methods biogeographic 2004). different it (Turner the scenarios of as test valuable a paleobiogeography provides Gondwanan Late of questions Cretaceous these to central it makes Madagascar That landmasses. pesuchus Gondwanan three in found Whether In Araripesuchus New xmndi hsstudy, this in examined ssae ihArc,SuhAeiaand America South Africa, with shared is Araripesuchus , Araripesuchus , and Comahuesuchus , Araripesuchus soeo e aawt pce rsn in present species with taxa few of one is Sebecus hr euiratrltrlyfacing anterolaterally peculiar a share and h he et r osrce tthe at constricted are teeth cheek the Notosuchus Mahajangasuchus Protosuchus Malawisuchus Araripesuchus Comahuesuchus , Simosuchus , Araripesuchus Libycosuchus .gomesii A. h klso h A8720 UA the of skulls the , and , scoeyale ihnoto- with allied closely is Theriosuchus .gomesii A. rmMadagascar from , and Hesperosuchus Uruguaysuchus oee,as shares also however, , , aelciasthat lacrimals have Araripesuchus Uruguaysuchus and Uruguaysuchus , caatr2- in 23-1 (character Like . , ´ 05.Further 2005). a .patagonicus A. Trematochampsa Notosuchus , .patagonicus A. Araripesuchus Oe 1879, (Owen Sarcosuchus ). l have all snow is Arari- Arari- share 261 and and , , , , Downloaded By: [Columbia University] At: 05:34 17 September 2007 262 n opesduigQikiesphoto-jpg QuickTime’s 198) size using 1, image (0, compressed Photoshop original Adobe and their in adjusted from levels with unreduced are data files All YawSpinHeadVoila.mov). PitchSpin- HeadVoila.mov, of (RollSpinHeadVoila.mov, movement renderings the the during transparent made (Roll- matrix gradually with “removed” and YawSpinHeadNomatrix.mov), digitally mov, PitchSpinHeadNomatrix. matrix matrix the SpinHeadNomatrix.mov, the with (RollSpinHeadMatrix.mov, with mov), images YawSpinHeadMatrix. generated the were PitchSpinHeadMatrix.mov, images. in CT digital models the for of present of each allowed set and for This A matrix density the high. bone of was “removal” between density each within contrast matrix much vary The not does sample. which of sandstone, density fine-grained matrix the make is the specimens of to the Most specimen axes. surrounding the orthogonal IA) three rotate the around that Fairfield, models Inc., left three-dimensional (VayTek, in are Voxblast which specimens the view. through lateral sagittal right left The view. from to dorsal proceeds passes in (SagFlipHeadMatrix.mov) and slice movie surface each dorsal with the ventrally in (HorFlipHead- on slice movie starts horizontal Matrix.mov) each The the with view. of anterior posteriorly tip movie proceeding the coronal rostrum, at The 210). begins 1, in (CorFlipHeadMatrix.mov) using leveled (15, were Photoshop images compressed the Adobe their for med- are values Grayscale (quality: from ium). and compression three unreduced photo-jpg size the QuickTime’s are image along All original generated axes. were slice-by- orthogonal QuickTime animations format. file slice 8-bit an to reduced asn sn h rgnl1024 original the using Maisano format TIFF 16-bit in files. slices data were individual interslice Images as mm. 0.078125 an archived also for at was thickness images interslice planes cutaway The mm. horizontal 0.078125 in of resliced and spacing was data sagittal slice reconstruc- mm), coronal 40.252967 the the The of and was 600. view scale 5000, tion of spacing reconstruction offset reconstruction field of the inter-slice (maximum field mm and 40 The thickness mm. mm, 0.08464 slice 0.08464 an two a offset, on no and scanned of with were wedge, orientation) samples mount empty The (angular view. per rotation views left samples per and 1000 acquired mA 0.165 were using and slices kV 120 Fifteen to unfiltered. set energies X-ray tAsi.Teutahg eouinsubsystem resolution ultra-high Texas The of 1024 University Austin. computed the X-ray in at at resolution scanned facility high (CT) were the tomography at 2297 2002 PR of June FMNH and (holotype) h ooa lcswr lomnpltdin manipulated also were slices coronal The aawr rcse yteato n Jessica and author the by processed were Data .H Turner H. A. £ 04CDcmr a mlydwith employed was camera CCD 1024 £ 04images 1024 sg eerfr oti ao’ itn geographic distant taxon’s known other this to voyage. relative long to location a refers for ironically here used Usage also walk”, a for out Tsangatsangana Etymology sp. nov. TSANGATSANGANA, ARARIPESUCHUS 1983) Whybrow and (Whetstone MESOEUCROCODYLIA 1930) (Hay CROCODYLIFORMES 1970) (Walker CROCODYLOMORPHA paleontology Systematic these suboptimal how of length were determine the are. lengths topologies to to that alternate compared The MPTs then tree. trees were the trees constraint those these the of only with compatible keeping option, lcmn of placement Notosuchus Sofr 02.Teigopcnitdo 29 of three of consisted taxa— outgroup protosuchian the ingroup and taxa The mesoeucrocodyliform 2002). (Swofford diinlhuitcsace eecnutdin conducted were in searches 2003). PAUP generated Maddison heuristic were and Additional Notosuchia (Maddison of v4.06 MacClade group sister the elctsadte ieto n reconnection PAUP and using bisection run branch-swapping tree (RA) (TBR) weighted addition and random equally 1000 replicates of with searches consisted heuristic analysis Parsimony analysis Phylogenetic http://research. this at throughout viewed found be be can amnh.org/users/turner. movies can generated CT analysis images work. CT the axis the of three coronal Examples by the created. a also along was Lastly, movie cutaway medium). dimensional (quality: compression hlgntcpsto of v2c position phylogenetic TreeRot using Decay calculated sequences. 1999). also TBR (Sorenson RA were replicates, 10 indices bootstrap boot- and 1000 nonparametric branch-swapping, with using Nodal examined trees. strapping, parsimonious was most number the support the not of but these topology length nor of tree the Ordering 111 affects and characters states. 79 or states 77, presence/absence character 67, of include 49, sets 45, nested unordered. 37, potentially as represent 15, 9, treated 1, characters Characters multistate all weighted, equally with were that characters morphological Orthosuchus estvt nlsswscnutdrgrigthe regarding conducted was analysis Sensitivity * sn h efretplgclconstraints” topological “enforce the using h aae nlzdcmrss129 comprises analyzed dataset The . n oouha re osriigthe constraining Trees Notosuchia. and Araripesuchus aaayepeso enn “just meaning expression Malagasy , Protosuchus Araripesuchus ihnNtsci,o as or Notosuchia, within , Araripesuchus Hemiprotosuchus ihrsetto respect with * v4.0b10 species. and Downloaded By: [Columbia University] At: 05:34 17 September 2007 30 MHP 37;tba FN R2300, PR PR (FMNH FMNH tibiae 2300, 2337); PR PR FMNH FMNH 2330, 8734, UA 2331); (UA 8776, PR FMNH (UA femora 2330, PR pubes FMNH 8778, 2330); UA 8777, PR 2330); (FMNH PR FMNH 2328); ischium 8768, PR UA (FMNH PR 2303, manual PR ilia FMNH FMNH 2328); 2301, 2328); PR 2312, (FMNH PR unguals PR manual FMNH metacarpals (FMNH 8759, 2339); 2312, phalanges UA PR PR 8736, FMNH (UA (FMNH 2324, ulnaria PR 2339); FMNH 2324, PR PR FMNH 2310, FMNH (UA PR FMNH radialia FMNH PR 8737, 2324); UA 2329, PR 2327, 8736, FMNH (FMNH PR PR pisiform 8758, FMNH 2339); 2326, PR (UA FMNH PR ulnae 2327, FMNH 2324, 2339); PR PR FMNH FMNH FMNH 2324, 2325–2327, PR 2321, (UA PR (FMNH PR radii humeri FMNH 2340); 2302, 2313); PR FMNH PR PR FMNH FMNH (UA 8721, 8741, coracoids UA 2334); PR 8740, FMNH 8779, 2313, (UA 8755); PR scapulae (UA FMNH 2333); chevrons PR 8733, 2334); (FMNH UA PR osteoderms 8732, FMNH (UA 8755, ribs 8729– UA dorsal 2309, (UA PR 2330); FMNH PR caudals 2308, FMNH PR 2330); FMNH 8735, PR UA 8731, FMNH FMNH 8774, 8767, UA 2301, (UA UA sacrals PR 8722, 8769–8773, 2304–2306); (UA UA PR dorsals FMNH 8742, PR UA FMNH 2323); 8739, PR 8723–8727, UA FMNH 8728, 2307, (UA postorbital 8764); PR cervicals 2322); FMNH (UA 8757, 2336); (UA PR nasals palpebrals 2315); FMNH PR 2332); (FMNH 8765, PR (UA FMNH 8752, FMNH frontals (UA 2317, jugals UA 2321); PR 8761, UA PR 8760, FMNH UA 8762, 8756, PR PR (UA (FMNH (FMNH maxillae premaxillae 2334); splenial 8754); (UA 2318); ribs 8751, angular PR UA 2316); dorsal FMNH 8750, (UA 8763, vertebrae, partial dentaries UA caudal 2299); angular, PR and right (FMNH dorsal splenial, right maxilla, ilium, dentary, right angular, right and skull, surangular nearly Partial left 2298); palpebral, left radius, PR left of (FMNH hindlimbs portions right right sacrum, and obscured column, vertebral partial complete radius ulna, left jugal, right and mandible, Partial right 2297); damaged PR skull, (FMNH left three astragalus humerus, right and the and calcaneum of shaft right the pubis ribs, of dorsal portion partial coracoid, damaged three head, ribs, femur, dorsal left lacking damaged of right femur head of proximal right end and proximal ilium ribs, left scapulae, with humerus, right vertebrae and presacral left twelve right partial and first left the including skull, hyoid, complete nearly large A Paratypes posterior lacking fenestrae. skull supratemporal the complete beyond portion nearly 8720, UA Holotype rcs rjcigfo h oslpr fthe of part dorsal the attenuating. that and from retroarticular mandible a premaxilla and projecting a bar, internarial lacrimal, process the the of little contact forms not nasals do tooth, that dentary 10th hypertrophied a possessing ifr rmalother all from Differs Diagnosis 2000; al. et 8). Rogers 1998, Figure of al. Hartman et and village Rogers (Krause 1998, Madagascar the northwestern near in Berivotra Late Basin, Mahajanga Cretaceous), (Campanian?—Maastrichtian, occurs Maevarano locality the of This Formation Member 93-33. Anembalemba the MAD within locality, student, from single 1993 graduate 15, a August Antananarivo on Rabarison, of Augustin Joseph University discovered a was material by referred all and holotype The locality and Horizon unidentified 2314). 2314); PR FMNH 8749, PR 8747–UA FMNH (UA (UA unguals unguals: 8749, (UA pedal phalanges 8747–UA unidentified 2337); 2330); FMNH PR PR FMNH 2319, 8736, FMNH PR 2330, FMNH 2311, PR PR 8744–UA FMNH UA FMNH 8743, 2337, 8746, (UA PR phalanges pedal FMNH 2338); 2330, PR PR PR FMNH 8780, FMNH UA PR 2319, 8736, (FMNH (UA astragalus PR metatarsals 2335); FMNH 2330); FMNH PR 2300, FMNH 2335, PR 2330, FMNH PR PR 8781, (UA FMNH fibulae 2337); 2330, PR FMNH etro a niae h td ra ,Mpo Madagascar, of Map B, Basin area. Basin. Mahajanga study Mahajanga the of of location indicates showing Map map of A, near Box center area). location. (grayed Formation Maevarano of Geographic exposures showing 8. Figure New Araripesuchus Araripesuchus rmMadagascar from pce by species 263 Downloaded By: [Columbia University] At: 05:34 17 September 2007 rna ev n aoi oaia h etquadrate left the The detailing foramina. carotid preserved, and well nerve is cranial is occiput table skull The the of present. most and frontal The 18A). (Figure posterior-most the and the of bones, consisting roofing pterygoid. cranium dermal of partial some the elements, a chondrocranial lateral is and occipital it FMNH 2299, Like material. PR least skull the recovered is the 8766 of UA the complete fissure. of metotic view a and provide capsules individual to otic out this prepared The in partially was preserved chondrocranium. and well the a is of magnum provides elements foramen left of likewise, the view of This, wall clear the medial The fenestra. the C). to to 18B, up supratemporal damaged caudal (Figure is orbits side cranium the right of the border of posterior but two-thirds preserved, roughly skull. the is of surface ramus ventral the to mandibular of compacted portion anterior right The present this. to the is anterior thereby is frontal bone no bones The chondrocranium but the dermatocranium. dermal of palatal lateral portions and the to of access providing sides, much both On lacks half. posterior it the of consists skull The and palate a dorsal the provide of cranial braincase. aspects external 2299) as internal well on PR as information structures FMNH of source 2298, secondary of PR morphology and FMNH cranial images internal detailed Araripesuchus the provide on data information These 2297. PR bone the ambiguous. of sutures some portions and damaged rendered compaction preservation moderate the during because though, this poorer than of is fragmentary condition skull overall preserved a The jaw. lower preserves and well skull 2297 entire less PR FMNH but holotype. complete more skulls. preserved the of jugal. smallest the left the the and is to of It adjacent portion preserved displaced small not surangular very a left is for the except posterior- skull missing, is The the jaw lower material. of are the portion 28), of most 18, isolated remainder is the Figures holotype The from 2299; condition. good PR FMNH in 8775, preserved FMNH UA 8766, 2298, 14–17), (UA PR Figures skulls bone partial 2297; nearly four PR the and other (FMNH of One skull pattern. much complete suture details isolated cranial is and of and 9–13) texture preserved number Figures well 8720; a (UA very as holotype well The remains. as blocks associated aeilfor Material preservation and form General mandible and Skull Description: 264 A87 opie uho h otro skull posterior the of much comprises 8775 UA preserving skull, partial a is also 2299 PR FMNH individual. complete nearly a is 2298 PR FMNH FMNH and 8720 UA for obtained were data CT is and five the of largest the is 2297 PR FMNH .H Turner H. A. h ata kls(A86,U 8775, UA 8766, (UA skulls partial The . .tsangatsangana A. srcvrdi three in recovered is ietdgov.Ti per iia oaforamen a to similar in appears present This groove. ventrally a directed with associated is teeth foramen This premaxillary 20). fifth (Figure and fourth the in between corner that gomesii to A. premaxilla/maxilla similar the foramen, to corresponding depression process. as maxilla. posterolateral just the extensive beginning the to face more prior contact with substantial becomes a contact by no contact evidenced narrow and to this rounded little a Dorsally, is show with margin nasals smooth posterior the hypertrophied The while constriction. a nasal away tooth, accommodate curves the dentary maxilla to from The B). the ventrolaterally fifth maxilla faces 9A, of (Figure and the fenestra anteriorly process first divides narial posterolateral the premaxilla external The and The anterolaterally. largest smallest. the the the with being premaxilla the in fourth present teeth five are There Premaxilla the of portion mandibular the and exposed. of quadrate pterygoid aspect the have ventral that skulls of few the the of end one is it Also, posterior choana. internal the of preserves 8775 portions UA is complete. nearly ear and preserved inner are and largely and braincase predominatelyabsent,theleftandrightlaterosphenoids present the is Although, basisphenoid complete. the of much and qiaeta h oae in duct. foramen topographically lacrimal the not the as are of equivalent however, foramina, end two anterior The the as suggested ewe h rmxlaadtemaxilla. any the in of determined in specimens be the cannot the participated nasals. of bar premaxillae the the the of by extent, formation and the what formed to largely or is premaxilla If, bar internarial the The between rmxlai ocv.Tedra oto fthe of portion dorsal The concave. is premaxilla neosuchians. is derived feature some in the other This paralleled of 21). with position (Figure the shared tooth roughly its maxillary and at fourth tooth position festooned maxillary dorsal third at enlarged slightly most position the of ventral is level most the margin its rectangular/trapezoidal reaching alveolar dorsoventrally, a The forms with snout. combined nasals, the This, only horizontal medially. with oriented portion of vertically portion dorsal-most nearly facial is The maxilla 2321. FMNH PR the 8720 specimen FMNH and UA in 2297 in preserved PR preserved maxillae is partial maxilla with complete nearly A Maxilla lose ntepseirmri sasmall a is margin posterior the on seen Also h neirmri ftejnto ihthe with junction the of margin anterior The hr safrmni h posteroventral the in foramen a is There . .patagonicus A. Araripesuchus .tsangatsangana A. hc reae l (2000) al. et Ortega which , .patagonicus A. .patagonicus A. pce n is and species . resides and Downloaded By: [Columbia University] At: 05:34 17 September 2007 robtlbr hc ssrca u a small a has but the surficial forms is portion which exposed bar, only posterior-most the preorbital is under The even It exposure maxilla. fossa. dorsal no antorbital posterodorsal is laterally—there the and posterior of view. the lateral surface forms in L-shape lacrimal inverted The an has lacrimal The Lacrimal bears surface foramina. small dorsal anteriorly, grooves and, The longitudinal shallow form. of composed in sculpturing linear rather the in as but with sutures zigzag not diagonally are and sutures and These prefrontal. frontal posteriorly the slightly with transversely widens narial It external the fenestra. of margin Anteriorly, posterodorsal narrow. the transversely forms it and long is nasal The Nasal the of The margin posterior fenestra. shape. the tooth, antorbital to in third prior just spatulate ends the row become tooth After teeth the largest. with maxillary the conical the are being the teeth tooth above three third least first just at The held teeth. maxilla foramina eight The 21). a large (Figure and margin moderately texture alveolar sculpted The of mildly jugal. a row the nearly bears of the surface process lateral anterior of is the fenestra. half maxilla by the overlapped anterior antorbital Posteriorly, fenestra. the the antorbital circular forms in maxilla participation The the from aaa rcse ftemxlaaesml and simple the are of maxilla surface the 22B). ventral (Figure of unsculpted the processes nasal contrast, the palatal of location In the to conchae. correspond may they that hsdfesfo h odto in condition lacrimal. the thereby the from contacting suture, differs from overlapping This nasals the broad the and a lacrimal separating the as in contacts ventral maxilla prefrontal the is of ascending the maxilla process Dorsally, 21). the (Figure dorsal to of opposed ventral portion the anterior-most than farther margin. anteriorly extends maxilla FN R22,Fgr 2) lr (1986) Clark 22A). in ridges Figure similar very fragment 2321, maxilla described are one PR on ridges preserved (FMNH These only are intersect- depressions. and of thin and series the 1970). ridges complex however, a ing tooth, (Parsons bear maxillary processes third crocodylians palatal the living to Posterior in found nasals the taxon. this to in due lacrimal prefrontal maxilla the the with The contact with wedge-shaped. contact a and lacks size equal roughly patagonicus A. nall In h ail ak h nenlsrt n sinuses and struts internal the lacks maxilla The Araripesuchus .gomesii A. hr h armladperna r of are prefrontal and lacrimal the where ifr rmti condition—the this from differs pce h aasaeexcluded are nasals the species Protosuchus .gomesii A. suggesting , .gomesii A. and omn h neoeta deo h orbit. by the overlapped is Unlike of jugal. lacrimal and the maxilla robust edge of the margin is anteroventral ventral The lacrimal posteriorly the expands the forming pillar of the Ventrally, pillar 23). narrows throughout. (Figure maxilla descending and the under The lacrimal lacrimal at the medially of the and end anteriorly the opens in duct The dorsally The anteriorly. bone. runs lacrimal the the of duct size the along one-third roughly is posteriorly and maxilla bar. anteriorly prefrontal length, the entire its contact under along extensive is prefrontal an lacrimal the forms The with region. and The overlap and past throughout, 23). this narrow (Figure in just fenestra thin fenestra is the maxilla maxilla antorbital of the margin anterior the under the past unexposed of just continues forward The midpoint extends fenestra. lacrimal the antorbital the the of remainder into extending lamina ltl eaae h armlfo otc ihthe with that contact com- from maxilla fenestra, lacrimal nasal. antorbital a the separates the with pletely over and coupled dorsally maxilla process, the extends of This regions nasal. posterior process the anterior short divides a the has that comprises, It as orbit. well the and as of wall area anterior broad pre-orbital dorsal is the solely, almost prefrontal the Anteriorly, Prefrontal palpebral. the for face contact in lacrimal the of edge lateral osntdfe uhfrom much differ in not does contact the which (2000), al. of et naso-prefrontal-lacrimal Ortega both to Contrary prefrontal, nasal. the and contact lacrimal wedge-like a sacako ig Otg ta.20:fi.5). fig. in 2000: short and al. prefrontal not et the is (Ortega between ridge nasal contact or the prefrontal crack Therefore, and lacrimal a the the as between mistaking while contact attributed lacrimal true the the was to process—they bone prefrontal the of posterior the remainder as the prefrontal. identified the only they of extent structure the due The of is exists misinterpretation difference a large to a that observation al.’s et in contact shorter lacrimo-nasal is however, similar taxa, are two 4). bones the three (Figure between the authors) of by proportions noted relative that The unlike is present is aa sabgos(iue4.Ol nclose a on apparent. does become and 268 elements Only two MUCPV the lacrimal of 4). between rostrum contact the (Figure the the of between ambiguous examination 269) contact is (MUCPV a nasal holotype of the the given presence fact, of In long. preservation very suture lacrimo-nasal the h armldc svr ag.Ispseiropening posterior Its large. very is duct lacrimal The hsi unlike is This New .patagonicus A. Araripesuchus .gomesii A. .patagonicus A. AN 45) hc has which 24450), (AMNH .gomesii A. hnin than rmMadagascar from .tsangatsangana A. .gomesii A. .gomesii A. wa difference (what .patagonicus A. h posterior the , ete is neither , .Ortega ak a lacks 265 Downloaded By: [Columbia University] At: 05:34 17 September 2007 266 .H Turner H. A. Downloaded By: [Columbia University] At: 05:34 17 September 2007 New Araripesuchus rmMadagascar from

Figure 9. UA 8720, Araripesuchus tsangatsangana. Skull in dorsal view (A, photograph; B, line interpretation of cranial sutures). Scale ¼ 1 cm. (Photograph by C. Leonard.) 267 Downloaded By: [Columbia University] At: 05:34 17 September 2007 268 .H Turner H. A. Downloaded By: [Columbia University] At: 05:34 17 September 2007 New Araripesuchus

Figure 10. UA 8720, Araripesuchus tsangatsangana. Skull in right lateral view (A, photograph; B, line interpretation of cranial sutures). Scale ¼ 1 cm. (Photograph by C. Leonard.) Madagascar from 269 Downloaded By: [Columbia University] At: 05:34 17 September 2007 270 .H Turner H. A. iue1.U 8720, UA 11. Figure rrpscu tsangatsangana Araripesuchus kl nvnrlve.Scale view. ventral in Skull . ¼ m Poorp yC Leonard.) C. by (Photograph cm. 1 Downloaded By: [Columbia University] At: 05:34 17 September 2007 iue1.U 8720, UA 13. Figure iue1.U 8720, UA 12. Figure rrpscu tsangatsangana Araripesuchus rrpscu tsangatsangana Araripesuchus kl nlf aea iw Scale view. lateral left in Skull . kl natro iw Scale view. anterior in Skull . New Araripesuchus ¼ ¼ m Poorp yC Leonard.) C. by (Photograph cm. 1 m Poorp yC Leonard.) C. by (Photograph cm. 1 rmMadagascar from 271 Downloaded By: [Columbia University] At: 05:34 17 September 2007 272 .H Turner H. A. Downloaded By: [Columbia University] At: 05:34 17 September 2007 New Araripesuchus rmMadagascar from

Figure 14. FMNH PR 2297, Araripesuchus tsangatsangana. Skull in dorsal view (A, photograph; B, line interpretation of cranial sutures). Scale ¼ 1 cm. (Photograph by C. Leonard.) 273 Downloaded By: [Columbia University] At: 05:34 17 September 2007 274 .H Turner H. A. Downloaded By: [Columbia University] At: 05:34 17 September 2007 New Araripesuchus rmMadagascar from

Figure 15. FMNH PR 2297, Araripesuchus tsangatsangana. Skull in right lateral view (A, photograph; B, line interpretation of cranial sutures). Scale ¼ 1 cm. (Photograph by C. Leonard.) 275 Downloaded By: [Columbia University] At: 05:34 17 September 2007 276 iue1.FN R2297, PR FMNH 16. Figure .H Turner H. A. rrpscu tsangatsangana Araripesuchus kl nvnrlve.Scale view. ventral in Skull . ¼ m Poorp yC Leonard.) C. by (Photograph cm. 1 Downloaded By: [Columbia University] At: 05:34 17 September 2007 savnrlflrn tiscnatwt h palatine the with contact its at prefrontal well flaring as the ventral medially, flares palpebral a that that process as orbital clear right dorsal the a is has the within displaced pillar it lodged by Nevertheless, and preserved cavity. length is broken which its is bone, along pillar the the midway and prefrontal of medially distorted lack left right The slightly The preservation. is of pillar 26). artifact prefrontal (Figure an be medial and pillar could the opening the narrower on of opening much the process are lacking pillars slanted, caudally prefrontal the and 99,atog tsol entdta hp and alligatorids. shape within that variable In noted is pneumaticity be of should extent it although 1999), odto sgnrlysmlrt htfudin found that to similar generally (pre- mississippiensis Alligator hollow is completely condition pillar recess process prefrontal frontal posterior the the with and pneumatic very becomes prefrontal. directed 24). the (Figure medially prefrontal/frontal of the between surface a suture pseudo-tongue-in-groove a ventral forms against This the Ventrally, flange rests. from abuts prefrontal directed process flange the ventrolaterally of this most a which has upon frontal cross- in The triangular and of thick that margin is process it dorsal section. where smooth. posterior the orbit, is along long the midway surface a nearly dorsal has grooves. extends the prefrontal longitudinal ridge The fine the dorsal with the to ridge, Lateral sculpted this to is Medial surface bone. the of midline sangana einn utpirt h ri,teprefrontal the orbit, the to prior just Beginning complex. is suture frontal/prefrontal The h oslsraeo h rfotlin prefrontal the of surface dorsal The .tsangatsangana A. iue1.FN R2297, PR FMNH 17. Figure a ogtdnlrdernigna the near running ridge longitudinal a has sensu imr19;Fgr 5.This 25). Figure 1995; Witmer h rfotlrcs ssmaller is recess prefrontal the , and liao mefferdi Alligator rrpscu tsangatsangana Araripesuchus .tsangat- A. (Brochu kl nlf aea iw Scale view. lateral left in Skull . n trgi.Ti ssmlrt h odto found condition the to similar in is This pterygoid. and h aitlpseirybtenteatro otends most meets anterior It the between is prefrontals. posteriorly parietal bone the the The by nasals. anterolaterally the a bordered of under in tips continuing most nasals frontal is anterior the the the the with between surface meets suture transverse narrow frontal dorsal nearly not the The Anteriorly, does orbits. and undivided. flat is transversely frontal The Frontal bar postorbital the of base siphonal small caudal 27A). a (Figure the is there at Also, ventral foramen (longitudi- pits). and ornamentation and grooves alveolar lateral nal an The have bar cross-section. surfaces post-orbital the triangular to This anterior a jugal 27B). the (Figure of most ventromedially gives longitudinal as rotates surface a jugal ventral posteriorly. the the bears becomes slanted posteriorly, surface also that, ridge is lateral and inward the its parietal obliquely Anteriorly, the directed is in is meet jugal the point It to the of rod-like. of a and surface projection slender to lateral dorsal The the tapers quadratojugal. with and contact slender overlapping infratemporal is or in posterior, process, and participate Its jugal, fenestra. not maxilla the antorbital does the preserved, crocodylomorphs, the poorly other is overlaps like dorso- it is lateral Although and jugal distinct lacrimal. broad the a Anteriorly, shows ventrally 27). and (Figure triradiate bowing is jugal The Jugal .gomesii A. New Araripesuchus AN 24450). (AMNH ¼ m Poorp yC Leonard.) C. by (Photograph cm. 1 rmMadagascar from 277 Downloaded By: [Columbia University] At: 05:34 17 September 2007 ueoscvte Fgr 9.Atrol,teei a with is marked there Anteriorly, is 29). (Figure frontal cavities the numerous Internally, postorbital. 28). (Figure fossa supratemporal fene the into supratemporal entering the of of Skulls 18. Figure uue nFN R29.Scale 2299. PR cranial of FMNH interpretation Line in C, 2299. PR sutures FMNH B, 8775. UA A, .Leonard.) C. 278 h rna a eysotcnatwt the with contact short very a has frontal The .H Turner H. A. rrpscu tsangatsangana Araripesuchus ta—h rna just frontal strae—the ¼ m Poorpsby (Photographs cm. 1 npseirview. posterior in h ideoeigit ubro hmes the chambers, three of number Posteriorly, a with frontal. into posteriorly opening run the middle to the continue of and open length for chambers ridge dorsal the the underneath half runs that recess large .Leonard.) C. h uuewt h ail.Scale maxilla. the with suture and premaxilla the of the detail showing contact, maxilla premaxilla/ left .Leonard.) C. iue1.Rcntuto ftesulof skull the Nash.) of Betti Reconstruction tsangatsangana 19. Figure iue2.FN R2321, PR FMNH 21. Figure 8720, UA 20. Figure iwo sltdmxla hwn etoe eta agnand margin ventral Scale festooned margin. alveolar showing the maxilla, above foramina isolated of view nlf aea iw Oiia lutainb Luci by illustration (Original view. lateral left in rrpscu tsangatsangana Araripesuchus rrpscu tsangatsangana Araripesuchus ¼ c .(Photograph1cm. by ¼ m Poorp by (Photograph cm. 1 Araripesuchus ls-pof Close-up . Lateral . Downloaded By: [Columbia University] At: 05:34 17 September 2007 ,Pltlve,soigsot ufc.Scale surface. smooth showing view, Palatal B, Fgr 0.Te ln aeoetal rdal and gradually lateroventrally slant They The in 30). frontal. as extensive (Figure the as of not are those parietalis cranii like crista recesses Also, lacks fenestra. bone temporo-orbital supratemporal this the the within is to postorbital for and restricted parietal the foramen between Contact the artery. fenestrae supratemporal borders the parietal of and The wall medial fenestrae. the constricted the forms supratemporal strongly meets the not is parietal between parietal the The Posterolaterally, squamosals. postparietal present. no but is the median indistinct parietal is of the supraoccipital small between the edge and suture a dorsal The crest. the with supraoccipital to concave is corresponding edge slightly projection posterior to The posterior 28). straight the (Figure the surface forms onto continue occipital not does and but table, skull undivided the of border is parietal The Parietal the of portion laterosphenoids posteromedial cristae. the The contact 29B). and the ascend (Figure for channel tracts ridged median a olfactory sharply border These well-developed cranii. suture, cristae with prefrontals The half. the continuous anterior which are the with along processes, bone lateral the runs of ridge midline small the a recesses postorbital– Ventrally, the contact. large underlying in and frontal–parietal orbit ending the to and posterior continuing two lateral 2321, PR FMNH 22. Figure rrpscu tsangatsangana Araripesuchus ¼ m Poorp yC Leonard.) C. by (Photograph cm. 1 Alligator ,Dra iwo h aaa aia hwn nenlrde n depressions. and ridges internal showing lamina, palatal the of view Dorsal A, . otcsavr ml oto fteprea ihnthe within it parietal the fossa. and of supratemporal contact, portion small parasagittal very a the a contacts meets and along postorbital table The medially skull jugal. frontal the the of from bar. side inset lateral the postorbital the is rod-like forming bar postorbital the portion The “V”-shaped of anteroventral a portion in the dorsal-most quadratojugal and the the suture broadly—the of ascen- of narrow process quadrate the ding portion contacting the portion posterior posteroventral contacting the with bifur- lamina, the and ventrally thin of descends cates A lamina supratemporal corner squamosal. broad the the anterolateral anteroposteriorly overlaps broadly the of and forms fenestra length It the fenestra. along midway 1991). al. et (Gasparini 1041) Notosuchus in seen 24450), is (AMNH This which rests. on palpebral shelf posterior lower 28). the a corner (Figure possesses corner indentation anterolateral indented concave This the slight The a with convex possessing surface. gently border overall dorsal an bearing anterolateral narrow flat, is a surface dorsal has postorbital The Postorbital supra- the within 314). CorFlip: high (2: fenestra temporal laterosphenoid the contact otrol,tepsobtletnsnearly extends postorbital the Posteriorly, Malawisuchus , New pr.os;MC N13,1040, 1037, RN MACN obs.; (pers. Araripesuchus .patagonicus A. Gmn 97 n peirosaurids and 1997) (Gomani rmMadagascar from Otg ta.2000) al. et (Ortega .gomesii A. 279 , Downloaded By: [Columbia University] At: 05:34 17 September 2007 eto hog h neirbre fobt,soigcontact showing orbits, of border frontal. and anterior prefrontals between the through 8720, UA section 24. Figure 8720, UA 23. Figure armladlcia eesmorphology. recess lacrimal and lacrimal 280 .H Turner H. A. rrpscu tsangatsangana Araripesuchus rrpscu tsangatsangana Araripesuchus ooa CT Coronal . ooa Tsie hog h otro oto ftersrm hwn internal showing rostrum, the of portion posterior the through slices CT Coronal . Fgr 1.Asmlrcniini on in found is matrix condition obscuring similar “V” and the A gomesii A. of fracturing 31). extent to (Figure dorsal due postorbital the the unclear with with “V”-shaped, suture The meet is to bar. PR anteriorly postorbital FMNH extends the in quadratojugal only The preserved 2299. is quadratojugal The Quadratojugal eiodl fteqart smlrto (similar quadrate is lateral the gonicus quadratojugal the in of The participate not hemicondyle margin. does and ventral dorsally obscures visible the however, of the jugal, of infratemporal most overlapping exposure. margin the The posteroventral lateral the at fenestra. forms narrow and moderate is fenestra has quadratojugal The and laterally onro h kl al.Tedra ufc slopes surface dorsal The table. skull posterolateral the the forms of that bone corner triradiate a is This Squamosal smooth. is quadratojugal the of otrol,teqartjgloelp h quadrate the overlaps quadratojugal the Posteriorly, u otatn htof that contrasting but , . .gomesii A. .Tesurface The ). .pata- A. Downloaded By: [Columbia University] At: 05:34 17 September 2007 mississippiensis qaoa opie n-hr ftesultable skull The the process. of paroccipital one-third the comprises forming squamosal otic and the overhanging downwards recess the bends and is long of process very posterolateral the attachment the and of concave, is portion the squamosal occipital The for musculature. ear groove external developed well the of wall lateral foramen. the artery comprises temporo-orbital and suture broad the a parietal supratemporal in the the contacts reach squamosal to the in lateral Medially, not fenestra. seen broad is that does squamosal to The but (similar bar postorbital postorbital the anteriorly under extends squamosal The laterally. gently very ( 8720 UA in recesses prefrontal of comparison and Detail 25. Figure h aea ufc ftesumslbrsafairly a bares squamosal the of surface lateral The C )a eni ooa lcstruhec skull. each through slices coronal in seen as D) —C, Notosuchus ). o eeoe etal tti otc si sin is the it contacts as quadrate contact the this Medially, at is basioccipital, crocodylians. ventrally quadrate the developed The not laterosphenoid. contact and to most basisphenoid, quadrate in horizontally the as Internally, extends inclined mesoeucrocodylians. dorsolaterally basal is quadrate The the Quadrate of long a length is the there runs 32). and (Figure process that chamber paroccipital chamber single hollow, a cylindrical completely by is lobe formed posterolateral The width. rrpscu tsangatsangana Araripesuchus New Araripesuchus A )adTMm78 ( m-7487 TMM and B) —A, rmMadagascar from Alligator 281 Downloaded By: [Columbia University] At: 05:34 17 September 2007 mississippiensis .Leonard.) C. iue2.U 8752, UA 27. Figure in is it no as however, present depression, is shallow a ridge foramina, has two otic quadrate the the surrounding the recess to otic siphonium. anterior the preotic small Within Just a bears incisure. quadrate the otic incisure otoccipital large surface a and dorsal The bears the canal. quadrate contact, cranioquadrate the the this forming to overlies Dorsal squamosal 34). (Figure ( otoccipital 8720 UA in recess prefrontal left through section CT Sagittal 26. Figure er ag raglrdpeso omdfo the from formed depression triangular large and quadrate slants a the reduced bears of surface surface very posterior articular The is medioventrally. The hemicondyle narrow. B). aspect medial lateromedially in 18A, spherical the nearly (Figure and is hemicondyle ventrally lateral The completely almost 33). 31, (Figures 24450) AMNH 282 otrol,temniua odlsaedirected are condyles mandibular the Posteriorly, .H Turner H. A. —B). rrpscu tsangatsangana Araripesuchus .gomesii A. Pie1959; (Price ,Mda iwo h ua.B aea iwo ua.Scale jugal. of view Lateral B, jugal. the of view Medial A, . uhwdrta al(iue1) h eta odris border ventral The 18). (Figure is tall it than view wider posterior much In dorsally. exposed not is bone This Supraoccipital ventral the 34). to (Figure only surface contributes quadrate the therefore rmteqart speet(unlike present the is from quadrate lamina the formed ascending No ae¨rum,from and squamosal. the of foramen large lobe posterior is the solel process of ridge bounded paroccipital the border is condylar to lateral which medial dorsally the ridge This condyle marks medial process. medial distinct the the paroccipital a of from terminus and running the process, quadrate, the paroccipital of surface dorsal rrpscu tsangatsangana Araripesuchus A n M -47( m-7487 TMM and —A) yteqart.The quadrate. the by y ¼ m Poorp by (Photograph cm. 1 Alligator Alligator and ) Downloaded By: [Columbia University] At: 05:34 17 September 2007 antb bevdin observed be cannot ti esdvlpdin developed less is It bv h oae anm hsrdei very is ridge This magnum. in foramen pronounced quadrates the The divided the between are above articulates. extending general, crest transverse in proatlas a occiput by the the and which exoccipitals, posterior small on a form process the they above where meet bones magnum, two foramen The 28). 18, (see Figures surface below occipital the of most form exoccipitals The Exoccipital 35). (Figure canal transverse a middle through the regions connecting ear antrum mastoid large a possess retn PladApesteguı and (Pol Argentina extend in not as does much supraoccipital as well- the anteriorly a CT in and that seen be midline can imagery It the present. is is of crest surface nuchal side developed occipital either The on dorsally). depressed faces surface this oslsraeo h rcs ae osltrlya in as (in dorsolaterally eusuchians faces process most the of surface dorsal Poorp yC Leonard.) C. by (Photograph h urocptlof supraoccipital The notasstronglytriangularinshapeasinmosteusuchians. 2299, PR FMNH 28. Figure iwo kl al hwn nepeaino uue.Scale sutures. of interpretation showing table skull of view eue si h other the in fenestrae, as post-temporal reduced the processes to postoccipital Dorso- Medial “V”-shape. are fenestrae small present. right post-temporal the a downturn of forming rudiments sharp laterally, midline, very the a at with angularity broad .tsangatsangana A. Notosuchus .gomesii A. rrpscu buitreraensis Araripesuchus .tsangatsangana A. rrpscu tsangatsangana Araripesuchus rtue hs r very are These protrude. Araripesuchus Alligator ´ 2005). a AN 25)and 22450) (AMNH and and huhi does it though , Comahuesuchus .patagonicus A. pce.The species. a more a has Dorsal . ¼ from cm. 1 . etal oiscnatwt h aipeod This basisphenoid. the extends with and contact (laterally its squamosal) canal to and cranioquadrate ventrally quadrate the by of enclosed wall medial iue2.U 8720, UA 29. Figure rni(cf.C otro eto hwn ueosrcse within recesses frontal. midway numerous the showing section section cristae Posterior distinct C, CT and (ccrf). recesses cranii B, frontal three (mfr). the showing the orbits recess showing through frontal section Anterior medium A, prominent recesses. frontal showing sections ag etoaea rcs speetadfrsthe forms and present is process ventrolateral large A New Araripesuchus rrpscu tsangatsangana Araripesuchus rmMadagascar from ooa CT Coronal . 283 Downloaded By: [Columbia University] At: 05:34 17 September 2007 ihpsobtladfiuet omaprino h mandibular the of portion a form to Scale failure condyle. and contact postorbital ”V”-shaped with showing yellow), in (highlighted quadratojugal iue3.FN R2298, PR FMNH 31. Figure unlike is It 33A). Figure allows see which but preserved (also poorly fissure, is nerves, distinguishable metotic X and (cochlear IX 34). the the of (Figure passage tympani of crocodylians remnant in bulla does The the it as of prominence) poster- portion the Anteromedially, forms cavity. opisthotic) oventral to cerebral (fused the exoccipital the of walls lateral basioccipital the of base the of 34). level (Figure the at ( is 2297 PR contact FMNH between morphology parietalis cranii crista and parietal of Comparison 30. Figure M -47( m-7487 TMM 284 h xciiasfr h otro at fthe of parts posterior the form exoccipitals The .H Turner H. A. ¼ liao mississippiensis Alligator m Poorp yC Leonard.) C. by (Photograph cm. 1 rrpscu tsangatsangana Araripesuchus —B). Right . yolsa XI oaiaaepeet Halfway present. are foramina (XII) hypoglossal aspect. has posterodorsal fissure metotic more the a that in species modern of adults ooa Tscinsoigrcs ntepseoaea oeof lobe posterolateral the in 2297, recess squamosal. showing the PR section FMNH CT Coronal 32. Figure tlatoe n ehp eod medial second, a perhaps and one, least At rrpscu tsangatsangana Araripesuchus rrpscu tsangatsangana Araripesuchus A and —A) . Downloaded By: [Columbia University] At: 05:34 17 September 2007 tde o aetesih eta xaainse in small seen excavation central the slight the of ventrally. have developed not all slightly does is It almost that condyle forms occipital basioccipital The not Basioccipital to is however, artery tract, 2297. The PR this carotid FMNH cavity. in of ear preserved middle internal portion the dorsal-most into the foramen. dorsomedially carotid pass allows posterior smaller this to slightly This ventromedial a just is and there present is large a vagi quadrate foramen the and condyle occipital the between 2297, PR FMNH 33. Figure utcinsnss n ,CrnlC etostruhtepseirrgtqart fbanae n ,Sgta Tscin through sections CT Sagittal D, and B braincase. of quadrate right midline. posterior the the of through right sections CT braincase Coronal the C, and A sinuses. Eustachian rrpscu tsangatsangana Araripesuchus Tiaeyo aiu rics opooyicuigoi asl and capsule otic including morphology braincase various of imagery CT . oaiaaentepsdo MHP 27 u are Eustachian but 2297, PR lateral FMNH present on The exposed not are crest. are foramina median crests the parasagittal notch alongside small small Two Eustachian a median small opening. forms the to basioccipital entrance the the roofs that of tip ventral rnvrai rfnu (Fu Maha- profundus the occipito- M. for transversalis and attachment is basioccipitovertebralis as crest jangasuchus medial serves well-developed which a tubera present, but Basal present, the 36C). not for (Figure are channel tube of posterior Eustachian portion the median anterior holds the and basioccipital condyle antrum the large occipital a The have 18). does (Figure eusuchians most New Araripesuchus rmMadagascar from ¨ bigr17) The 1876). rbringer 285 Downloaded By: [Columbia University] At: 05:34 17 September 2007 aa itecnb ado h nenlmrhlg of morphology internal poor CT the is of the there said which be of for can skulls little Because data, two specimens. the in all preservation in preserved of B” “ridge to corresponding (1973). two Iordansky the of ventral the akdb w aaaitlrde like is ridges and surface parasagittal 22450) two external (AMNH by The a furrow. marked by marked transverse is pterygoid lateroventrally. The narrow the with pterygoid exoccipital. contact the ventral The the and the laterally medially, with basioccipital quadrate suture the contacts broad basisphenoid basisphenoid a compression the shares antero-posterior Posterodorsally, little (verticalization). very exposure, posteroventral exhibiting large a has basisphenoid The Basisphenoid cavity. cerebral the of floor the posterior the of forms part basioccipital 368– The CorFlip: 35). (2: Figure data 385; CT in present clearly and large otic right the of detail showing 2299, Scale occiput, capsule. PR the of FMNH view Ventrolateral 34. Figure 286 aipeodrsrmi rsn u poorly but present is rostrum basisphenoid A .H Turner H. A. ¼ m Poorp yC Leonard.) C. by (Photograph cm. 1 .patagonicus A. rrpscu tsangatsangana Araripesuchus MCV269), (MUCPV .gomesii A. . hscnatcnb envr lal in clearly very seen data be can CT contact quadrate. shows and This show quadrate. poorly laterosphenoid the very between not or however, contact the does prootic 345–347), CorFlip: therefore the (2: is and with latero- portion preserved contact the posterior-most well where the not exposed, specimens is both sphenoid In the of wall braincase. anterolateral the forms laterosphenoid The Laterosphenoid of located wall and opisthotic. the preserved dorsal is within of canal portion semicircular the lateral anterior the The forms foramen. perilymphantic and the of the the fissure, of border border with dorsal metotic posteroventral the delineates suture the bulla, tympanic the forms exact dist opisthotic its not The but is 2297 exoccipital PR FMNH lateral the preserved. poorly although very is canal, canal semicircular semicircular of half lateral anterior the the and canal 383–386; of semicircular portion anterior CorFlip: ventral the the (2: contains prootic prootic tympani The the 35). bulla of Figure VIII the margin the posterior to the of ventral on branch seen CT a been and for can 2299 incision PR the and FMNH however, both VIII scans, CT In the visible. the for nerves nor openings VII specimen anteroventral any the are in data 383–393, Neither CorFlip: 35. 2: Figure scan and CT prominence); 2299, (cochlear PR tympani FMNH anterior see bulla the the forms it of Internally, portion lateral braincase. the the on visible of is side prootic the whether unclear is It opisthotic and Prootic basioccipital. the the dividing the from seen to is opisthotic fissure Dorsal metotic a 36). the sinus into 33, rhomboid (Figures open sinus tube posterior rhomboid laterally, Eustachian and large median anterior the Moving the of side, channels preserved. left carotid the anterior on not this the especially and of are defined extent well foramina anterior not The is channel 33). be anterior to (Figure appears the tube preserved Eustachian to median corresponding the for structure PR channel the FMNH a in but surrounding basisphenoid and the 2297, bone within area poor between is the matrix Contrast of rostrum. or basisphenoid basisphenoid the h iln u ontcnatec te,smlrto similar other, each contact gomesii not A. do but midline the a with sutures within the this, basisphenoid. to rests the ventral contacts and, it fossa process ventral There shallow capitate anterodorsally. The postorbital frontal the parietal. with and sutures laterosphenoid the Dorsally, h pshtci eeal elpeevdin well-preserved generally is opisthotic The h neirtp fteltrshniscneg at converge laterosphenoids the of tips anterior The oce o h xto rna evsII nerves cranial of exit the for Notches . nusal nC data. CT in inguishable .gomesii A. . Downloaded By: [Columbia University] At: 05:34 17 September 2007 n I r ifiutt icr,btapa obe to appear but in as discern, edge, anterior to the difficult along present are III and 2297, PR FMNH 35. Figure utcintubes. Eustachian ogune obig oooost htse in character seen that the and in to present connectivity is homologous crocodylians bridge of no based principles congruence, then area 37) the the Figure in (goV, on groove laterosphenoid the of by wall indicated al. lateral traversing et indeed, the (Clark is, across branch the bulbi ophthalmic levator the and If M. 1993). branch in the ophthalmic supplying present branch nerve’s of not path trigeminal the fact, to corresponds the ovale in foramen the to is, dorsal bridge tsangatsangana a A. that suggest (Iordansky laterosphenoid the a bridge of the and 1973). body between main nerve mandibular the the its trigeminal and group, of the crown offshoot the of small eusuchian In branch 1992). early ophthalmic Norell and the CorFlip: (Clark in (2: imagery present CT to in evident corresponding be 336–352). structure may a this fractured, although Fgr 7,C.Altrshni rdemybe may bridge laterosphenoid A in present ovale) C). (foramen 37A, opening trigeminal (Figure the foramen of the edge from anteriorly ovale pass (supplying bulbi) The levator branch data. M. anterior CT the or the fossil forms the laterosphenoid IV from nerve discerned cranial be for cannot foramen A crocodylians. modern h al fteltrshni htaeexposed are that laterosphenoid the of walls The not is but crocodylians in present is bridge This .tsangatsangana A. Fgr 7.Asalwgroove shallow A 37). (Figure rrpscu tsangatsangana Araripesuchus FN R29)and, 2298) PR (FMNH .tsangatsangana A. Hylaeochampsa .gomesii A. . ooa Tscino kl hog rics hwn einadlateral and median showing braincase through skull of section CT Coronal . and nFN R29 Fgr 9) Tscans CT 39A). (Figure preserved 2297 partially PR only FMNH are in ectopterygoids The Ectopterygoid lateral 25B). the passage. (Figure form nasal the and scans 2297 of tall CT are walls relatively PR the are palatines FMNH in laminae seen and These the clearly 8720 of are UA and both laminae in dorsal None- preserved fenestra. the suborbital the of theless, of portion border a forming anterior laterally the extends palatines the of o xedrsrlyvr a nteetx.N palatal No taxa. these present. in is far fenestra very rostrally extend not of that Comahuesuchus from different but Notosuchus the anterior peirosaurids to the similar in is beyond This condition rostrally fenestra. cm 2 suborbital the or extend of 1 border They maxillae the margins. between with narrow lateral somewhat are can subparallel palatines palatines the The and of inferred. outline maxillae be general the a of shape, their processes with palatal the no the reveals bears of further and data presence CT narrow features. is distinct distinctive It A other present. 38). is (Figure septum choana choanal anterior internal the the and of palatines border the the of in preserves portion present 2297 caudal-most PR not FMNH specimens. or the preserved of most poorly are palatines The Palatine Like .gomesii A. New MC N13,14,14)or 1041) 1040, 1037, RN (MACN Araripesuchus MC 63)tepltnsdo palatines P6131)—the (MACN and .patagonicus A. rmMadagascar from ml portion small a , 287 Downloaded By: [Columbia University] At: 05:34 17 September 2007 osse eywl rsre oslsurface dorsal preserved well very 40). (Figure a possesses gomesii otecon a enin seen (as of choana absence posterior depression or the midline a presence to or the line suture so sagittal a ventral preserved surface the of ventral is exposure flat surface good broad No anteriorly. a faces has that plate pterygoid The Pterygoid bone 296). The 2:CorFlip: the contact (see to maxilla. bar enough postorbital posterior the the extend not and to of appears jugal portion the ectopterygoid any contact posterior-most lacks The to the and anterolaterally process. of extends extent, posterior surface significant ventral significant any pterygoidal the portion. the to ventral overlap of pterygoid its not edge in lateral does recess the flange, small forms a bone is moderately This revealed scans Also short, the C). a in 39B, (Figure reveal ectopterygoid robust 250–284) CorFlip: (2: 2297, PR FMNH 36. Figure aea utcintbs otro aoi oae,adbsocptlantrum. basioccipital and foramen, carotid posterior tubes, Eustachian lateral 288 .H Turner H. A. antb eemnd MHP 2299 PR FMNH determined. be cannot ) rrpscu tsangatsangana Araripesuchus .patagonicus A. and oiotlC etos(,B )o kl hog rics hwn einand median showing braincase through skull of C) B, (A, sections CT Horizontal . A. haa.Ti neirpoescp h posterior the internal caps the process of anterior enclosure This dorsal choanae. the and posterior boundary the form These of troughs. pair open a The lateroventrally forms midline. extension anterior the this of along surface forward ventral extends it as narrows the pterygoid. with the continuous of process dorsally, (palatal) facing anterior and surface) transverse side posteriorly flat facing rostral ridge a long the the This of than side (rather face. caudal the the ridge with posterior a on forming and process, continues basis- anterior narrow the distinct transversely distinction is a posteriorly, process with and dorsal The quadrate, foramen phenoid. latero- the the the of border contacts ovale), ventral that the process (forming dorsal sphenoid a has 2298) V29 oee,ti oslsraewsnot was surface dorsal this MUC however, and exposed. 268 269, PV MUC 64). both PV 2000: of al. examination et my (Ortega In concave” than rather “flat as h oslsraeof surface dorsal The hsatro rcs sbodpseiryand posteriorly broad is process anterior This h trgi nthe in pterygoid The .tsangatsangana A. .patagonicus A. a described was FN PR (FMNH Downloaded By: [Columbia University] At: 05:34 17 September 2007 rs ntedroaea odr.Tepeyodof pterygoid the The between resting ventrally, borders. more distinct sits the species dorsolateral a modern has the of and on palatines process the crest on anterior high sits the pterygoid Unlike crocodylians, palatines. the of modern processes dorsal the of portion Left Scale C, 2298. process. PR capitate FMNH possible of of showing details laterosphenoid 8775, and UA nerve of ophthalmic view for lateral groove Left B, ophthalmic nerve. in with and associate possibly artery features and as well laterosphenoid parietal, as ovale, basisphenoid, foramen laterosphenoid, of between relationships Details tsangatsangana 37. Figure .Leonard.) C. ,Rgtltrlve fFN R29,showing 2299, PR FMNH of view lateral Right A, . ¼ m Poorpsby (Photographs cm. 1 Araripesuchus aa hs r ogadnro n etwti the within rest and narrow CT and in long identifiable are but These broken data. badly are vomers The slight Vomer or this preservation if of the however, palatines. result on the unclear, of the palatines disarticulation is is the It separation of palate. ends secondary posterior-most the ol etrb hrceie saspu.In septum. a as the characterized divides gomesii be “spine” A. completely the CorFlip: better respect, it (1: this would data In and CT passage. 25) in nasopharyngeal fig. spine seen shaped be 317–442; 2000). can “T” al. “spine” et a (Ortega This choanae internal possesses the divides pterygoid that the that rna ilr u ul slcigo h dorsal the on lacking is bulla in a as surface. but them pillar, capping frontal not tsangatsangana processes A. palatal dorsal r rsre ste omteatro odro h internal the of border anterior the form they palatines the choana. as of portions preserved caudal surrounding are the the Only with removed. palate digitally the matrix of reconstruction CT dimensional iue3.FN R2297, PR FMNH 38. Figure thsbe oe in noted been has It New and Araripesuchus .patagonicus A. h trgi otcstepre- the contacts pterygoid The . .gomesii A. rrpscu tsangatsangana Araripesuchus rmMadagascar from h etmseparates septum the , and .patagonicus A. Three- . 289 Downloaded By: [Columbia University] At: 05:34 17 September 2007 iiihspseiry steavoipseirto posterior alveoli morphology the This as slight posteriorly, margin. with diminishes mandibular spatulate anteriorly a forms anterior directed this first together, the mildly Taken procumbency. and alveoli, teeth remaining 1–11 medio- nine the alveoli to from broad in lower margin situated alveolar moderately element the with and characteristic laterally, flat very dorsoventrally a tsangatsangana is A. dentary The Dentary in as palatines. palate the the on and clarki exposed maxillae not are the They of processes palatal 2297, PR FMNH 39. Figure h urudn arxdgtlyrmvd h lmnsaeicmlt ihteetn fterpeevto ihihe ngen nC, In green. in highlighted preservation their of with extent ectopterygoids the right with and left incomplete the are of elements reconstruction CT The three-dimensional removed. a scale (C) digitally and matrix (B); surrounding skull reconstructed the CT through slice coronal a 290 ¼ .H Turner H. A. Bclye l 2000). al. et (Buckley m Poorp yC Leonard.) C. by (Photograph cm. 1 h rttido h etr is dentary the of third first The . rrpscu tsangatsangana Araripesuchus Simosuchus opooyo cotrgi,a hw napoorp fdra ufc (A); surface dorsal of photograph a in shown as ectopterygoid, of Morphology . hl h otro hr srltvl mohand in smooth seen relatively is third pits, unornamented. small relatively posterior bears the surface ventral while the The after alveolus. dorsally ninth slopes tenth gradually the dentary maxillary at The third tooth. indentation enlarged the subtle accommodate to a alveolus is B). there 41A, ceasing (Figure and Laterally, narrow tooth row dentary a to tooth penultimate by the the Posterior below to replaced parallel alveoli. are running foramina the groove the the to 15, along ventral alveolus less row dentary a or form lateral foramina more neurovascular small directed are alveolus vertically. tenth the hsgnrldna opooyi iia othat to similar is morphology dental general This of series A alveoli. sixteen has dentary the total, In .patagonicus A. , .gomesii A. and .wegeneri. A. Downloaded By: [Columbia University] At: 05:34 17 September 2007 a ypyelrgo.D ooa eto iwytruhobt,soigtevria etr ossigltl rn etoeillmn.In dorsoventrally lamina. the ventromedial showing no or 8720, little UA possessing of dentary rostrum vertical scale anterior the showing through B, orbits, section and through Coronal midway A C, section dentary. Coronal D, the region. of symphyseal outline flat complex the showing 2318, PR iue4.Dnaymrhlg in morphology Dentary 41. Figure 2298, PR FMNH 40. Figure trgi n aeopeod Scale the laterosphenoid. between is contact process and true anterior the The pterygoid displacing pterygoid. thereby of dorsally surface deflected dorsal of view lateral .Leonard.) C. ¼ m Poorpsb .Leonard.) C. by (Photographs cm. 1 rrpscu tsangatsangana Araripesuchus rrpscu tsangatsangana Araripesuchus ¼ c.(Photograph1cm. by Left . ,Dra iwo etr MHP 38 ,Rgtltrlve fFMNH of view lateral Right B, 2318. PR FMNH dentary of view Dorsal A, . ) ovrey h otro hr ftednayis dentary all the of in third 41B, thin posterior (Figure the lamina deflected Conversely, little dorsally C). a bears of way and the in flattened dorsoventrally completely oigo o n oel(04,ti aemor- same this (2004), both Norell in seen and is phology Pol the of Contra extend fenestra. coding not mandibular does external but the angular beneath the overlaps Ventrally, dentary 42A). the the (Figure of fenestra margin mandibular upper lamina external the forming ventro-medial laterally, surangular no to D). little 41A, (Figure with oriented nmda iw h ekla roetpr anteriorly tapers groove Meckelian the view, medial In ipasayhn htcudb ecie as in dentary described like the be third, of anterior could third morphology. The that middle the “convex”. the anything of only oversimplification displays reality, an perhaps of In is of This bit deflected. dentary ventrally a and the dorsal described is that 64) patagonicus (2000: A. al. et Ortega .wegeneri A. h otro agni hnadoelp the overlaps and thin is margin posterior The New Araripesuchus Araripesuchus and shvn cne”ltrlprofile lateral “convex” a having as .tsangatsangana A. .patagonicus A. aaadaevertically are and taxa rmMadagascar from and ,isnearly .gomesii A. .gomesii A. 291 . , Downloaded By: [Columbia University] At: 05:34 17 September 2007 ypyelrgo yn a n ra.Teei no the in is as with symphysis There terrestris the view broad. of and would dorsal constriction flat abrupt rami in lying When “V”-shape dentary region symphyseal 43A). a right formed and (Figure the have left of mandible the length the articulated, the dentary of of The symphysis two-thirds alveolus. ninth roughly the the comprises to into back posteriorly further canal conti- 55–173). Meckelian CorFlip: the (1: the marking dentary groove of the the nuation at of present demarcating terminus is foramen alveolus, anterior small eighth A contact. the splenial below and the surangular point the between the of contact view the to Medial transiliens, C, Cartilago lamina. the triangular for projecting Scale lamina dorsally overlapping coronoid. small triangular dentary possible the the the a showing showing showing 2299, and 2299, 2298, PR PR angular, FMNH PR FMNH of FMNH angular of in and surangular dentary mandible the left posterior of of view view Dorsal Lateral A, B, morphology. angular. mandible the Posterior 42. Figure hsi hn lnaebn htbgn tthe is splenial at The begins fossa. to Meckelian that the posteriorly of bone most expanding elongate enclose symphysis, thin, a mandibular is This Splenial taxon. former the in present symphysis suchus 292 h ogai ftesmhssetnsantero- extends symphysis the of axis long The .H Turner H. A. AN 45)lcstede trough-like deep the lacks 24450) (AMNH MC-N13;pr.os) and obs.), pers. 1037; (MACN-RN ¼ m Poorpsb .Leonard.) C. by (Photographs cm. 1 Notosuchus Araripe- elas well V-hpdpoesbtentednais This other in dentaries. found the that like between is “V”-shape process “V”-shaped a rtbrnelctda h uuebtentetwo in the between elements suture the splenial at located protuberance mandible. the of the surface symphysis, ventral of the one-third the approximately forms to and and narrows splenial Posterior N31) GDF603). MACN (MNN 1041), P6131; 1040, (MACN in RN1037, seen symphysis (MACN splenial U-shaped the ftesmhsa egh(iue4A.This is symphysis 43A). the in at (Figure found splenial that like to length dentary of symphyseal proportion the strongly one-third of roughly a constituting PR forming and takes suture FMNH symphysis splenial interlocking mandibular of The the 2298. data in PR FMNH CT part be as also 2297, well can as PR but 2297, FMNH 2316, in PR FMNH seen in preserved best and etal,teslnastprt on forming point a to taper splenials the Ventrally, nmda iw hr sasalpseirydirected posteriorly small a is there view, medial In .wegeneri A. Malawisuchus GF700). (GDF .gomesii A. Gmn 97,btdfesfrom differs but 1997), (Gomani .tsangatsangana A. Pie15,Hct1991) Hecht 1959, (Price Araripesuchus Comahuesuchus Anatosuchus Notosuchus .Alarge as Downloaded By: [Columbia University] At: 05:34 17 September 2007 iwo nioae ih peil hwn ag foramen large showing splenial, right isolated Scale oralis. an intermandibularis of view kl ihtesronigmti iial eoe.BadC ooa Tscin hog nu,soigteslna/etr otc and contact splenial/dentary the showing snout, through sections CT symphysis. Coronal the C, at and splenials B removed. of digitally interdigitation matrix surrounding the with skull iue4.FN R2316, PR FMNH 44. Figure in is it as gomesii present A. is oralis intermandibularis foramen 8720, UA 43. Figure rmd2d6 hssih ovxt sunique is alveoli convexity dental slight and This mandible to the d12–d16. the of from forms This wall shape. convex medial slight a on which lamina taking splenial, the thins, This of alveolus. remainder the tenth with the continuous of is the level overlaps the and at millimeters dentary few a anteriorly projects aebe rsre,tog e iue4)cno be cannot 42) to Figure determined. not see not appears though preserved, (which type been is morphology coronoid have its the the splenial so with specimens, contact convexity. the in the and of any of any observable in lack margin preserved is that posterior-most it splenials The that have extent specimen) the (to osly h peilpsessati aiathat lamina thin a possesses splenial the Dorsally, .tsangatsangana A. Fgr 44). (Figure rrpscu tsangatsangana Araripesuchus — .gomesii A. ¼ rrpscu tsangatsangana Araripesuchus m Poorp yC Leonard.) C. by (Photograph cm. 1 and .patagonicus A. ,Vnrlve fmniua ypyi natredmninlC eosrcino the of reconstruction CT three-dimensional a in symphysis mandibular of view Ventral A, . Lateral . eaoyl Fgr 5,B.I vrl hp,the the shape, overall to In of corresponding B). ceratohyals 45A, likely (Figure slender, ceratohyals and long h otro afcre wyfo h anterior the 35 from a away roughly cross-section. curves at in half portion cylindrical posterior and straight The is bone the gomesii A. niecooyin,btsmlrt te basal other to similar but crocodylians, fenestra. the mandibular Unlike of the to process anterior and posterior above dentary the contacts surangular The Surangular of 45C). ceratohyal (Figure the that dorsoventrally shows Araripesuchus and imagery CT compressed expanded. becomes element richardsoni vignaudi paau r nw rmafwcrocodylomorphs— hyoid few the a of e.g. from Portions known only mention. are (1991) apparatus cursory Hecht a but them hyoids, gave the preserve 24450) opsi orydescribed. poorly crocodylo- is non-crocodylian in morphs apparatus hyoid The Hyobranchials h lmnspeevdin preserved elements The eprscu agilis Hesperosuchus New Fe ta.2002). al. et (Frey h rsre neirhl ftebd of body the of half anterior preserved The . CletadMo 1951), Mook and (Colbert Araripesuchus shlo hogotislength its throughout hollow is .tsangatsangana A. Cake l 2000), al. et (Clark 8 nl.Ti oto fthe of portion This angle. rmMadagascar from .tsangatsangana A. .gomesii A. r iia to similar are Protosuchus Geosaurus (AMNH 293 are Downloaded By: [Columbia University] At: 05:34 17 September 2007 rjcini iia ota enin seen m-7487), that (TMM to similar bordered is This is C). projection 42B, it medial (Figure and groove “U”-shaped small smooth a The by is laterally a projection dorsally. the is of projecting surface there lamina fenestra, triangular mandibular external above pars surangular the the of externus surface upper mandibulae the On adductor medialis. the of insertion of sculpted. surface slightly lateral is The bone surangular. ventral- the dorsal the the thin of is overlaps A portion however, most line. angular angular, jaw the the the of lamina with with the parallel contact and of The straight wall surface. Unlike lateral articular 47B). the (Figure gomesii fossa forming glenoid descends arch mandibular an it bearing portion as posterior-most the only with posterior fenestra. and mandibular external dorsal the the of borders of half and concave approximately is forms surangular the of margin The 42A). anteroventral (Figure dentary the by overlaps) to overlapped opposed (as is surangular the mesoeucrocodylians, 2297, PR FMNH 45. Figure arxdgtlyrmvd ,Crnlscinsoigtehlo neiro h yis nA scale A, surrounding In the with hyoids. skull the the of of interior reconstruction CT hollow three-dimensional the a showing in section hyoids Coronal of C, view Ventrolateral removed. B, digitally bones. hyoid matrix right and left of view and 294 ls iiaiyo hp n oain h tutrsin the structures the Given location, and shape 1973). of similarity (Iordansky close externus adductor the of mandibular superficialis pars the of attachment from formed the is groove the and transiliens, Cartilago the nmda iw hr sasalwfsafrthe for fossa shallow a is there view, medial In straight is surangular the of margin dorsal The nteetx,tepoeto evsa h otc for contact the as serves projection the taxa, these In aescu trigonatus Paleosuchus .H Turner H. A. h uaglrde o omaypr fthe of part any form not does surangular the , amncrocodilus Caiman rrpscu tsangatsangana Araripesuchus FN 81980). (FMNH .mississippiensis A. TMm-7365) (TMM etadrgthod,wt nenldti rmC mgr.A biu ventral Oblique A, imagery. CT from detail internal with hyoids, right and Left . A. oe agno h nua sanro ig that ridge narrow a is angular the of the from and margin Continuing 2298). lower larger PR FMNH FMNH becoming (see and individuals 2297 pits PR larger in the spaced with closely more pits, small with the which This through passes. fenestra. groove cartilage deep mandibular Meckelian a external comprises the forms surface process of anterior features. floor long distinctive the the of no surface bears dorsal and the The of smooth surface is medial The angular known 46). is (Figure angular 8754 UA isolated from an 42A and (Figure respectively) angulars C, specimens; and right most complete both 2298 nearly in PR preserve FMNH preserved and 2297 that PR poorly FMNH however, projection is bone posterior process. retroarticular This the thin of surface lateral the external a the overlies posterior of the surface and curved ventral-most surangular, of the dorsally overlaps border that a portion ventral fenestra, the process mandibular anterior forms acute long, a that of consists angular The Angular in present not 24450). is (AMNH lamina triangular the for and determined be cannot of structure presence this The mentioned. crocodylians modern the tsangatsangana A. h aea ufc fteaglri ornamented is angular the of surface lateral The ieysre h aefnto sin as function same the served likely ¼ m Poorp yC Leonard.) C. by (Photograph cm. 1 .patagonicus A. .gomesii A. Downloaded By: [Columbia University] At: 05:34 17 September 2007 sntnal spoone si h omrtaxa. former the in as pronounced as nearly not is eusuchian ,Coeu oslve fgeodfsaadrtoriua rcs.I ,scale removed. C, digitally In matrix process. surrounding retroarticular the with and skull fossa the B, glenoid of skull. of reconstruction to CT view articular three-dimensional of dorsal a relationship Close-up in overall showing C, articular removed, right digitally of matrix view surrounding posterior the Oblique with skull the of reconstruction CT dimensional iue4.FN R2297, PR FMNH 47. Figure in found This that ridge. to distinct a similar of is form the ridge in not is division other other this the the in In present species. is portion dorsally the curved and process retroarticular the individual overlapping largest 42A). the Figure 2299; in PR (FMNH is projecting pronounced ridge This portion. posterior particularly curved dorsally the the from dividing lamina dorsally, deflects iue4.U 8754, UA 46. Figure h iiinbtentepseirprojection posterior the between division The Mekosuchus rrpscu tsangatsangana Araripesuchus Araripesuchus u in but , rrpscu tsangatsangana Araripesuchus .tsangatsangana A. Simosuchus pce,however, species, Araripesuchus aea iwo nioae etaglr Scale angular. left isolated an of view Lateral . n the and opooyo h riua hglgtdi elw.A oslve fthree- of view Dorsal A, yellow). in (highlighted articular the of Morphology . it Fgr 7.Tegeodfsai ogl vlrwith ovular roughly is fossa glenoid The 47). (Figure seen. the be with cannot surface angular contact and lateral surangular its its matrix, and in rock surface by ventral articular obscured partially left and in the is articulation element the is that Given 2298. PR preserved FMNH specimen articular only The Articular for area surface large a attachment. provides pterygoideus perhaps ridge This h oslsraeo h riua scomplex is articular the of surface dorsal The ¼ m Poorp yC Leonard.) C. by (Photograph cm. 1 New Araripesuchus ¼ m Poorp yC Leonard.) C. by (Photograph cm. 1 rmMadagascar from 295 Downloaded By: [Columbia University] At: 05:34 17 September 2007 otrol rjcigpoesoiiaigfo h oslpr ftemnil.B, mandible. the of part dorsal the from originating process projecting posteriorly rjcigpoesoiiaigfo h eta ato h adbe Scale mandible. the of part ventral the from originating process projecting aiapeeton present retroarticular lamina the closely of more process resembles It slender. process. retroarticular the to of fossa base glenoid anterior the the around contact extending and fossa any the wall lateral of the form forming instead not fossa, glenoid the does of part surangular The in eusuchians. present wall most glenoid The posterior quadrate. the the lacks fossa on complementing glenoid hemicondyle the medial 45 the to a of corresponds slant approximately slant at This ventrally angle. slants The the fossa hemicondyle. twice articulating medial respective nearly their appear of fossae area two the Additionally, of 1037). articular RN the the (MACN as prominent on as present nor defined ridge sharply an as This not quadrate. is by the ridge of divided hemicondyle a side are either to on corresponding fossae fossae two the with These ridge anteroposterior fossae. broad two of variation Systematic 48. Figure 296 h eratclrpoesi ogadrelatively and long is process retroarticular The area. and size in equal roughly are fossae two The .H Turner H. A. .gomesii A. n ak h xesv medial extensive the lacks and .patagonicus A. Araripesuchus oouhsterrestris Notosuchus .Likeother eratclrpoess A, processes. retroarticular 8 riuainof articulation still specimen. 48). the but on (Figure preserved position, not extension ae¨rum is dorsal foramen posterior A more direct the a retaining from originates Instead, it posteriorly. extending position, ventral a from uv oslya h rcse fmr derived more of processes the other Unlike do. as neosuchians dorsally curve Araripesuchus hrce ecie yCake l n Gomani and only al. et in that Clark present as generally, by is al. described et “noto- Ortega character that by used appears was not it suchian” movement, Second, longitudinal enlargement. allowing refer simply (1997) articular Gomani an and (1989) trait is al. the Two et First, Clark made. (1997). which be Gomani to to need and clarification (1989) of al. points notosuchians”, et this to Clark to “exclusive citing refer being They previously “enlarged”. as as trait area refer articular they text the in to and extension, medial and posterior a riua rcs of process articular ¼ reae l 20)fiue h cranio-mandibular the figured (2000) al. et Ortega cm. 1 rrpscu patagonicus Araripesuchus rrpscu tsangatsangana Araripesuchus h rcs ae oslybtfisto fails but dorsally faces process the , .gomesii A. .tsangatsangana A. Notosuchus and MCV29 a posteriorly a has 269) (MUCPV .patagonicus A. Araripesuchus FN R29)hsa has 2297) PR (FMNH , Malawisuchus osntoriginate not does h retro- the , shaving as and Downloaded By: [Columbia University] At: 05:34 17 September 2007 rrpscu wegeneri Araripesuchus lni os s“nagd in “enlarged” is fossa glenoid clade. Chimaerasuchus of Variation 49. Figure et—ohdnayadpremaxillary/maxillary—are and dentary teeth—both lhuhkonfo other from known Although Teeth notosuchian of group restricted a crocodyliforms. a remain to of appear articulation characteristic jaw long a and Propalinal motion motion. propalinal of longitudi- indicative or grooves surface nal articular glenoid the of extension tsangatsangana A. udaeatclt r nedlre hnthe that it than appear However, the hemicondyles. not larger of the does hemicondyles of indeed facets the are articulating which articulate on quadrate fossae two the ti ehp rubeta h riua ufc fthe of surface articular the that arguable perhaps is It n a o piie o larger a for optimized not was and GF70 ihfiepeailr et.C, teeth. premaxillary five with 700) (GDF Araripesuchus oss infiatposterior significant a possess .gomesii A. Araripesuchus Araripesuchus rmxlaytohnme.A, number. tooth premaxillary , .patagonicus A. pce,the species, ie that given or rrpscu gomesii Araripesuchus rrpscu tsangatsangana Araripesuchus yia ftehtrdn etto on nmost in found dentition heterodont the of typical ailr et,ada neemndnme of number twelve undetermined or an eleven teeth. and dentary 49B), teeth, (Figure maxillary premaxillary five ubro etr teeth; dentary of undetermined an number and maxillary, 11–13 premaxillary, aueo uho the disarticulated of the much Given of detail. nature dental the the obscur- cases of articulated, most most preserved ing in are mandible because and is skull This described. poorly ftehi h dentary; the in number teeth undetermined an of and Comparatively, teeth, and maxillary dentary. eleven teeth, the maxillary in gomesii A. eleven total teeth least eighteen at 49A), (Figure h Tiaigo h w kls uhmr fthe described. of be can more taxon much the skulls, for two morphology the dental of imaging CT the ngnrl h et of teeth the general, In .tsangatsangana A. New a orpeailr et Fgr 49C), (Figure teeth premaxillary four has GMDP 3-)wt orpeailr teeth. premaxillary four with 432-R) (GDM-DNPM Araripesuchus U 70 ihfiepeailr et.B, teeth. premaxillary five with 8720) (UA .tsangatsangana A. a v rmxlayteeth premaxillary five has .patagonicus A. rmMadagascar from .tsangatsangana A. .wegeneri A. aeiland material a four has 297 has are Downloaded By: [Columbia University] At: 05:34 17 September 2007 te aai o iia ota of that to similar not is (e.g. taxa other teeth baurusuchids dentary certain pachecoi enlarged are there with and in hypertrophied of tooth row last dental The the mesoeucrocodylians. basal among pit a 160–190). in CorFlip: (1: rest m4 would and tooth m3 maxillary between third this the occluded, of When extent tooth. the to hypertrophied as is unique very profile it is tooth procumbent dentary tenth weakly The 41B). and (Figure a outward giving splay teeth slightly, nine forward first These size decrease again. then in and size increase in d1–9, of largest teeth the is These which d5, until premaxilla. the of those bear morphology). not do are spatulate margins denticles. their and distinct and (a striated posteriorly are blunt teeth moving These more size and in shorter maxillary remaining decrease The teeth 2002). and Broin (Prasad ziphodont de not Lapparent are the therefore on and denticles distinct carinae, any in bear not ridges do shallow but enamel with the medio- marked is compressed surface weakly conical, The are laterally. very teeth and hypertro- three recurved, a first slight These by the tooth. followed of third are size phied and the (roughly teeth) first small The premaxillary are pm3. teeth of maxillary size two the size—roughly in fifth the decreased and largest size the being in fourth increase the with teeth posteriorly premaxillary The to point. sub- taper acute and an are recurved slightly teeth cross-section, in circular premaxillary The crocodylomorphs. ti otsmlrt htse in seen that In to 1986). Clark similar most richardsoni is It R29 n MHP 28 h ta is atlas The 2298. PR FMNH FMNH in and preserved is 2297 region cervical PR the of Much preservation and form General skeleton axial Postcranial teeth carinae. Description: the known on denticles GDF distinct their have of 7) to teeth (Figure 700 maxillary regards posterior with of morphology—the species imaging ques- other CT this elucidate or Only will tion. discoveries specimens known of fossil currently species future other the Only whether uncertain Araripesuchus is but It (Wermuth five, uncommon 1953). not and is formula four tooth in between of variance number varies The teeth species. premaxillary the among similarity general 298 ail.Termiigtehi h etr are and dentary premaxilla posteriorly. the size in in the decreasing teeth spatulate, between and remaining short notch series, The dental a the maxilla. in in anterior more resting is tooth trophied yetoh fdnaytehi o widespread not is teeth dentary of Hypertrophy like shaped are dentary the of teeth nine first The oprsnwt other with Comparison .H Turner H. A. .I ihrcs,tehprrpyse nthe in seen hypertrophy the case, either In ). .wegeneri A. AN 04 obr n ok1951, Mook and Colbert 3024; (AMNH aeahprrpiddnaytooth. dentary hypertrophied a have .richardsoni P. Comahuesuchus eitssbtnilyfo the from substantially deviates Araripesuchus oee,tehyper- the however, , MC 63)is P6131) (MACN .tsangatsangana A. Baurusuchus Protosuchus hw a shows . eercvrdfo h lc otiigteother the containing 2307) block tsangatsangana PR the Araripesuchus FMNH from and recovered 8728 were isolated (UA Two vertebrae spines. neural cervical are and lack cervicals but These c2–c5 complete exposed. entire however, mostly cervicals series, only the the this are In c8? 2298 to girdle. posterior PR pelvic terminating the preserved, FMNH is the series In of vertebral partially some right detail. are illuminates The imagery c1–c5 obscured CT matrix. and rock. damaged by by is obscured hidden c7 are of side c3 lateral of and surface axis dorsal the the of Much c5. through articulation littered area bone. an of reveal fragments images broken CT with but not out, was column prepared vertebral the axis and between elements region the atlantal The the to to process. odontoid as sutured the unclear as was centrum it pleurocentrum making the column, whether vertebral remain- the the of from two der dislocated These what preserved pleurocentrum. to were atlantal adjacent elements the lying be is to which appears 2297, PR intercentrum FMNH the in by only represented unambiguously opee armwsas eoee FN PR (FMNH mostly recovered isolated, ilia. An also exposed. left was the is sacrum and of that complete, surface right all dorsal are both the sacrum of to portions attached Unfortunately, 2298 PR respective are in their others 2298. with PR of rib FMNH association number in dorsal vertebrae close a in one and preserved d12 preserves with 2297 articulation incomplete. PR or FMNH disarticulated generally are preserved and described preserved the well all generally intact. fully are containing These block specimens. larger likely the are from and spines, series. dorsal neural the in trans- and anterior their these individual processes retaining exposed, preserved, larger verse fully better not slightly are While PR vertebrae a 2299). FMNH PR with to (FMNH block belonging same six the are 2298 processes There in transverse unexposed. present are their dorsals spines of neural much their lack and a these that of block. given the All higher within unexposed be remain the to may vertebrae likely few of is number orientation This 2298. the images skull. the to to CT respect due with and column help vertebral obscured the little to is provide articulated the morphology not of much is articulated, dorsal are c8 ribs is of the c7 Because rib both, vertebra. In the 2297 2298. and PR PR FMNH damaged FMNH in preserved in not disarticulated and axis and were atlas which the of ribs, exception the with vertebrae their nFN R29,tevrereaepeevdin preserved are vertebrae the 2297, PR FMNH In h armi opee ti rsre nFMNH in preserved is It complete. is sacrum The those and missing are ribs dorsal the of Most recovered were dorsals isolated nine Additionally, PR FMNH in vertebrae dorsal 14 least at are There to attached were ribs cervical the specimens both In specimens. Downloaded By: [Columbia University] At: 05:34 17 September 2007 BclyadBoh 99 and 1999) 2000), of Brochu and that al. (Buckley to et similar (Clark is 2297) oee,te ontcnatec te dorsally. other each contact of not intercentrum The do they however, iue5.FN R2297, PR FMNH 50. Figure in Like 2000). al. et of and reception (Ortega the Crocodylia condyle for from occipital shaped” known the “ring are are arches arches neural neural PR FMNH patagonicus atlantal A. or 2297 the PR FMNH 2298, either in and 50) not 190–215 the Figure a Though HorFlip: 2: crocodylians. from (see although in present only unambiguously known taxa, known well is is crocodylomorph structure atlas of the of handful arch neural The complex Atlas–axis blocks. the of prepared any in and block. in large intact the these from fully recovered of were but vertebrae one caudal isolated to only three addition In and these, vertebra. two respective 2298 its Only with PR articulation from series. FMNH come the with preserved of likely are arches middle haemal and identified unambiguously the small in The rather somewhere 2298. are PR FMNH vertebrae with association in preserved remains but right sacrum The rib. this the preserved. with from is disarticulated preserved rib also and sacral is damaged rear ilium is right vertebra the sacral only anterior The 2301). necnrm(re)adpsil erlac natredmninlC eosrcino h kl ihtesronigmti digitally matrix surrounding the with skull the of structures. two reconstruction same CT the three-dimensional showing a section in Coronal arch B, neural removed. possible and (green) intercentrum obnsceryietfibea atai r present are gastralia as identifiable clearly bones No are caudal as identifiable vertebrae nine Only MCP 6) nMCP 6,the 269, PV MUC In 269). PV (MUC oouhsterrestris Notosuchus .tsangatsangana A. rrpscu tsangatsangana Araripesuchus aaagscu insignis Mahajangasuchus eprscu agilis Hesperosuchus rcdlsporosus Crocodylus Pl2005), (Pol FN PR (FMNH Tiaeyo osbealsnua rh(elw.A neoeta iwo atlas of view Anteroventral A, (yellow). arch neural atlas possible of imagery CT . MHP 28lcsisnua pn edrn it rendering spine edges. neural central its the lacks reach 2298 PR not on FMNH do and depressions anteroposteriorly. medially, run These depressions constricted and shallow is side centrum. anteriorly either the centrum of axial pronounced length The the more across slightly is diminishes ridge centrum. the of This surface ventral median the A along extends 51–53). ridge (Figures wide than longer is axis the processes. and any dorsal lacking smooth, The are progress- margins blade-like. medial then becoming a and rib expands revealing the capitulum ively of visible the shaft fully behind The is constricts facet. rib contact depressed left The slightly exposed. the is head rib of left the complete, single capitulum of nearly half anterior is a the rib only atlantal but possess right The 51). and (Figure compressed, ventrally facets. capitular large moderately of arch, remnants the with 2297 only neural PR FMNH in it damaged and that is odontoid suggests The 52) was. pleurocentrum (Figure 2298 axis PR FMNH although the to fused taxon. latter the in seen process opesdlike compressed etal epa in as dorso- curves deep not which is ventrally visible, postzygapophysis is The ventromedially. postzygapophysis Only right damaged. or the exposed poorly are neurapophyses hlo ocvt eaaigteln capitular long the separating concavity facets. shallow with long than a broader element 15529)—an (FMNH ncnrs otealsitretu,tecnrmof centrum the intercentrum, atlas the to contrast In dorso- anteriorly, slender are ribs atlantal The was process odontoid the whether unclear is It In .patagonicus A. New Araripesuchus Alligator MCP 6) h atlantal the 269), PV (MUC Notosuchus tlcstesallateral small the lacks It . rmMadagascar from Pl20) but 2005), (Pol 299 Downloaded By: [Columbia University] At: 05:34 17 September 2007 .Leonard.) C. om oss ra,atrpseiryexpanded anteroposteriorly crocodyli- spine. broad, neural Most axial a shape. possess its forms determine to impossible 2297, PR FMNH 51. Figure ae h w tutrso h centrum. the the sepa- both on area structures on two double- little the axis Very is rates the diapophysis. rib and contacts axial parapophysis The directly fossa. and circular headed shallow there diapophysis a the to is nearly posterior than Just are shape. PR they larger in cross-section ovular FMNH are in of and diapophyses parapophyses The diapophysis the exposed. right are the 2297 and FMNH of 2298 diapophysis the left PR to The dorsal ridge. just ventral centrum, median the of edges anteromedial eni hs aa h pn,hwvr sntas not of is spine however, axial spine, the terrestris The as restricted taxa. anteriorly those in seen perhaps 5;Sglp 5–7;Fgr 3.Teeimages These 53). Figure that suggest 154–174; 327– CorFlip: SagFlip: (2: imagery 354; CT in viewable is 2297 are PR that 2005). poster- (Pol spines shape shifted in axial triangular breadth, and with iorly, anteroposterior form in reduced general this from 300 otyaohss h pn a o sbodas broad anterior as (cf. substantial extension not the lacking was crocodylians, modern spine the of The axial terminus the the the postzygapophyses. to that posteriorly developed than and was spines, spine anteroposteriorly neural cervical broader succeeding slightly was h xa aaohssaesaladlctdo the on located and small are parapophyses axial The tlataprino h xa erlsieo FMNH of spine neural axial the of portion a least At .H Turner H. A. Chimaerasuchus Pl20)(iue 52–54). (Figures 2005) (Pol .tsangatsangana A. liao mississippiensis Alligator oouhsterrestris Notosuchus W n us19b deviate 1996b) Sues and (Wu rrpscu tsangatsangana Araripesuchus a erlsiethat spine neural a had M m-7487) TMM Pl20)and 2005) (Pol Notosuchus eta iwo neircria etba eis Scale series. vertebral cervical anterior of view Ventral . etclyoine Fgr 2.I hsrset tis of it prezygapophysis respect, the nearly chus this to In is similar 52). again and (Figure oriented anteriorly vertically projects prezygapophysis oFi:1510 iue 1 3 u preserves but 53) 51, 356–383; Figures CorFlip: partially (2: 135–180; is 2297 (c3) HorFlip: PR cervical FMNH third in the damaged of arch neural The vertebrae cervical Postaxial length. is entire its rib along the concave of strongly bent surface centrum. though dorsal breadth, The axial in their slightly. uniform dorsomedially is the contact rib the on are of both shaft The surfaces heads and two articular together respective The closely tuberculum. spaced circular is ovular the rib more right and than larger the is but capitulum The 2297 exposed. PR best FMNH in ribs preserved both of are capitula The 51). (Figure elements headed of rsn nteai of axis the on present orpeevto,i sipsil odtriei a if determine to ( to Due lamina impossible exposure. is suprapostzygapophyseal lateral it preservation, partial the poor giving a medioventrally surface slightly articular ventrally, slants face instead not does but surface laterally articular curving The posteriorly, slightly. project the and of spine base neural the from originate They prezygapophyses. h xa isin ribs axial The h rzgppye r eysal iia othat to similar small, very are prezygapophyses The h otyaohssaemc ogrta the than longer much are postzygapophyses The Chimaerasuchus W n us1996b). Sues and (Wu u nietoeof those unlike but .tsangatsangana A. .tsangatsangana A. sensu ¼ m Poorp by (Photograph cm. 1 Notosuchus . o 05 is 2005) Pol Chimaerasu- r double- are The . Downloaded By: [Columbia University] At: 05:34 17 September 2007 evclrb.B eta iwo neircria etba.Scale vertebrae. cervical anterior of view Ventral B, ribs. cervical h erlsiei rae neootrol hnthose than anteroposteriorly broader is spine 52). neural (Figure The c3 on spine neural the lacks arch, neural and the pedicel anterior the preserving although of 2298, PR prezygapophyses. FMNH much and spine neural the 2298, PR FMNH 52. Figure rrpscu tsangatsangana Araripesuchus evclvrerlmrhlg.A etltrlve,soignua rhsand arches neural showing view, lateral Left A, morphology. vertebral Cervical . ¼ m Poorpsb .Leonard.) C. by (Photographs cm. 1 otrol vrtepszgppye,btntt the to not but postzygapophyses, the over anterior posteriorly slight rests spine neural The a cervicals. remaining the in present is alsonot is There thespinethat of portion dorsal cervicals. the to curvature remaining the of New Araripesuchus rmMadagascar from 301 Downloaded By: [Columbia University] At: 05:34 17 September 2007 xetse in seen extent 2297, PR FMNH 53. Figure Notosuchus crocodylians. modern the in closely seen more condition resembling 1999) Brochu and Buckley posterior the shows (C) section spine. medial-most the of the development the bisect the of and extent exactly contacts anterior the just not reveals spine, (B) (A) do section middle section the Sections lateral-most spine, neural spine. The spine. neural and the shape vertebra the of showing vertebra, position axis the and through sections CT sagittal of 302 nohrsalbde eourcdlassc as such mesoeucrocodylians small-bodied other In .H Turner H. A. and Chimaerasuchus aaagscu insignis Mahajangasuchus rrpscu tsangatsangana Araripesuchus h pn scentered is spine the , U 8654; (UA Series . ietdvnrlywt h riua aesexpanded. facets articular the with ventrally column. directed the and cervicals anterior through the on posteriorly short are parapophyses The moves posterior one as the larger on size in increases the cervicals. ridge on seen This that distinct axis. like a anteroposteriorly is ventrally running present ridge is What any vertebrae. corners. on well-developed the processes of anteroventral hypapophyseal the distinct no the are to There at due located linear parapophyses appears this ventrally contact viewed when although slightly circular, is non- surface to articular among anterior amphiplatyan The condition mesoeucrocodylians. eusuchian are prevalent 56) the 52, amphicoelous, 51, (Figures most seen crocodyliforms. trait other profile—the in rectangular commonly a Sues has unlike taxa and which view, these lateral Wu of in 2005, triangular spines are neural (Pol the pedicels Additionally, 1996b). neural the over eosrt h otro oiino h erlsieoe the over spine neural the c6 of and position c5 postzygapophysis. Vertebrae posterior vertebrae. the fifth, cervical the demonstrate eighth showing and column seventh, vertebral and sixth, skull the of reconstruction iue5.FN R2297, PR FMNH 54. Figure aaohssaeoua nltrlve n become and view lateral in ovular are Parapophyses vertebrae cervical eight the of centra the All rrpscu tsangatsangana Araripesuchus .tsangatsangana A. .CT Downloaded By: [Columbia University] At: 05:34 17 September 2007 rzgppye.Tepszgppye fc3 of postzygapophyses The prezygapophyses. slightly 2297). is PR (FMNH there curvature although dorsal more cervicals, anterior the from significantly vary that processes prezygapophyseal have otro evcl fcooyin Pl20) The 2005). of (Pol cervicals the posterior crocodylians in more of seen cervicals that closely posterior more resembles morphology Chimaerasuchus oee,i te o-rcdla mesoeucroco- crocodylians. as non-crocodylian most such in other dylians, condition seen in This that However, medial. resembles articulation directed the closely with partially slightly surface slanted strictly not but are horizontal facets The anterodorsally. c4 directed and c3 are of processes prezygapophyseal The circular. tuberculum. for rib facets the flat with articulation bear diapophyses in postaxial All seen trunk. orientation the a the assume approaching to begin projection diapophyses the lateral that c7 until remain not is It diapophyses vertebrae, orientation. directed ventrally succeeding less The or more the a maintaining arch. in flattened neural dorsoventrally the from and seen as are such insignis depressions taxa These crocodyliform centrum. other the in of wall the are in depressions deep and are there parapophysis, the centrum and to ventral dorsal the Both ventrally. than of laterally more midline directed dorsoventral the nearer to originate parapophyses the posteriorly, Moving Leonard.) C. by (Photograph morphology. 2297, rib PR FMNH cervical 55. Figure h otyaohssaea ag sthe as large as are postzygapophyses The h aeso h rzgppye r roughly are prezygapophyses the of facets The ventrally project diapophyses flattened the c3, On oouhsterrestris Notosuchus U 64 uke n rcu19:Pae1) Plate 1999: Brochu and Buckley 8654; (UA W n us19b h cervical the 1996b) Sues and (Wu oouhsterrestris Notosuchus Pl2005). (Pol rrpscu tsangatsangana Araripesuchus .tsangatsangana A. Mahajangasuchus Pl20)and 2005) (Pol onot do ealo evclnua pns hwn urzgppyellmnea elas well as laminae suprazygapophyseal showing spines, neural cervical of Detail . opooyi lopeetin present also is morphology This 55). PR (Figure laminae FMNH suprapostzygapophyseal with on continuous the is based rollspinhead), (2: that, data ridge CT and low 2297 a spines the by of marked and surface are lateral tall The lateral 55). in moderately 54, broaden arches. (Figures to are view begin spines neural the spines c7 by the the but narrow, over of majority centered the The back, are c4 From spines arches. neural neural tall very have which osvnrladatrpseirlnts therefore lengths, from to differing anteroposterior proportion in and equal roughly dorsoventral are They among similar. very common be 2005). (Pol to mesoeucrocodylians in basal appears small-bodied seen processes those physeal the as pronounced terrestris bears but as postzygapophyses are laminae not the axis. These short of the laminae. suprapostzygapophyseal surface of those dorsal are posterior-projecting The resemble long that 2298) have postzygapophyses cervicals PR remaining (FMNH aiafrteatcmn fteitrpnu ligament interspinous (Fu the of attachment the for median lamina small best a the bears spine 2297 the PR surface, this FMNH On in exposed. c6 of spine neural the 2298,with PR FMNH and 2297 PR FMNH both in exposed 2000). for al. determined et (Ortega be cannot but 24450) erae nsz ntepseircervicals posterior the on lamina size be this cannot in crocodylians, vertebrae modern cervical decreases In other the determined. on lamina h erlace ftecria etba r all are vertebrae cervical the of arches neural The h otro ufc ftenua pnsi poorly is spines neural the of surface posterior The ¨ bigr17) h rsneadetn fthis of extent and presence The 1876). rbringer rsneo uhlmneo postzyapo- on laminae such of Presence . New Araripesuchus oouhsterrestris Notosuchus rmMadagascar from .gomesii A. or Chimaerasuchus .patagonicus A. Notosuchus (AMNH 303 , Downloaded By: [Columbia University] At: 05:34 17 September 2007 neirpoeso ahrbi vrapdb the depression. by “U”-shaped dorsal overlapped the the anterior in is the resting with crocodyliforms, rib process rib preceding the each other of process of posterior in process As anterior tuberculum the capitulum. between passing and entire and rib the the running of is length depression shaft broad “U”-shaped the dorsoventrally of shallow surface a a dorsal and the On that point process. process acute posterior anterior pro- an an to capitular of tapers consists and shaft tubercular The cesses. the to perpendicular separated gap. and small size a in by equal roughly tuberculum are The capitulum double-headed. and are taxon ribs new postaxial the All in 52A). 51, 1921, (Figures Whybrow conservation of that (Mook and ribs cervical Whetstone the group and 1951, 1983) Mook the and is within rib Colbert cervical conserved crocodyliform highly the of morphology The ribs Cervical 2005). (Pol posteriorly prominent remain rjcigfo h oslsraeo h posterior the of lamina surface directed dorsal the posterodorsally from thin projecting a of presence Mo 91,wiein while 1921), (Mook 2307, PR FMNH 56. Figure C,addra D iw Scale view. (D) dorsal and (C), 304 nitrsigfauein feature interesting An being by characterized is rib the of shaft The .H Turner H. A. oouhsterrestris Notosuchus rrpscu tsangatsangana Araripesuchus ¼ m Poorpsb .Leonard.) C. by (Photographs cm. 1 .tsangatsangana A. .tsangatsangana A. hs laminae these sltdprilatro evclvrer natro A,pseir() etlateral left (B), posterior (A), anterior in vertebra cervical anterior partial Isolated . reflect sthe is iia etr speeto h evclrb of ribs cervical the on present is insignis Mahajangasuchus feature structure. of similar barb-like surface distinct lateral A more a the forms from This rib. elevated larger the slightly In as 96). well pronounced, 52A, more as becomes (Figures projection shaft the rib specimens the of process otro evclrb n h neirdra ribs the dorsal between anterior In the form last 1921). and in the (Mook ribs intermediate which cervical is convention in posterior rib the crocodylians follows modern cervical vertebrae division with dorsal This used and 57). cervical (Figure between transition h etao h neirdra etba are is vertebrae surface dorsal articular centra. anterior anterior cervical the The the of asymmetrical. like centra amphiplatyan amphicoelous The are weakly 58–63) (Figures to centra dorsal All vertebrae Dorsal the to shaft. extends rib more that the a of lamina has midpoint and anterior ribs thin dorsal curved anterior prominent less and of shorter shafts is the rib than c8 to the perpendicular of not shaft is The ribs, them. dorsal in ribs like shaft, cervical the like but spaced and closely are tubercular processes The capitular intermediacy. this demonstrates h ih i ftec speevdadmrsthe marks and preserved is c8 the of rib right The .tsangatsangana A. U 8654). (UA h i fc8 of rib the , Downloaded By: [Columbia University] At: 05:34 17 September 2007 aia Poorp yC Leonard.) C. by (Photograph lamina. Poorp yC Leonard.) C. by (Photograph iue5.Psto fprppye nvrered-3in d1-d3 vertebrae on parapophyses of Position 58. Figure on parapophyses the dorsal of four presence first continued the the least at on present poor vertebrae. is to asymmetry, it the due of but extent unclear the is exposure or It preservation posteriorly. posterior reflecting the expanding and edges than dorsolateral the plane) with surface horizontal articular the (in broader 2297, PR FMNH 57. Figure 28,soigprppye nnua rhadcnrmbd.B ih aea iwo 2add FN R29) Scale 2299). PR (FMNH d3 and d2 of view lateral Right B, body. centrum and arch neural on parapophyses showing 2298), hsuiu opooyapast erltdto related be to appears morphology unique This rrpscu tsangatsangana Araripesuchus 8add is 8soigtastoa auewt eldvlpdatro thin anterior well-developed with nature transitional showing c8 ribs. d1 and c8 . rrpscu tsangatsangana Araripesuchus h neirbreso h etaaentepne as expanded not are centra the of borders however, anterior crocodylians, the In crocodylians. within found aaohsso 1d a epiiiefor primitive be in may unknown (its d1–d3 crocodyliforms on Parapophyses of to dorsals (FMNH appears anterior insignis condition the d1–d3 in similar of present A be centra 58). Figure the 2298; of PR surface lateral the BclyadBoh 99 i.7 lt 1). plate 7, Fig. 1999: Brochu and (Buckley New ,Lf etoaea iwo 1add FN PR (FMNH d2 and d1 of view ventrolateral Left A, . Araripesuchus rmMadagascar from Protosuchus Mahajangasuchus ,btis but ), ¼ cm. 1 305 Downloaded By: [Columbia University] At: 05:34 17 September 2007 n 3o nadditional an d2, of d1, d3 as interpreted and are isolated 2298 block PR the FMNH three on containing 58B) Figure observation, 2299; PR this unsurprising (FMNH vertebrae upon in (Mook Based This Crocodylia in 1921). 2298. found is condition PR this that FMNH given on a d1–d3 lack of centra absence or dorsal presence remaining hypapophyses. PR the The FMNH or determine hypapophysis. 2297 to PR FMNH 2298 either in sufficiently not preserved is d4 unfortunately d1–d3, of centra the process. ventral transverse just the depression to shallow a bears centra anterior h oslcnr.I sucerhwfrback far how unclear is It be to on centra. parapophyses looks the dorsal vertebrae of the dorsal the presence of lateral the few with surface first right correlated anterior the the the in of centra preserve swelling The nicely surface. vertebrae These in 59. Figure i-oslvrer,FN R20.C otro oslvrer,FN R20.D adlvrer,FN R20.Scale 2308. PR FMNH vertebra, Caudal D, 2304. PR FMNH vertebra, dorsal Posterior C, 2305. PR FMNH vertebra, Mid-dorsal Poorpsb .Leonard). C. by (Photographs 306 itntprppye r rsn ntecnr of centra the on present are parapophyses Distinct of surface ventral the on presence are Hypapophyses .tsangatsangana A. .H Turner H. A. rrpscu tsangatsangana Araripesuchus h aea ufc fthe of surface lateral The . .tsangatsangana A. oprsno sltdvrerei neirve.A neirdra etba MHP 36 B, 2306. PR FMNH vertebra, dorsal Anterior A, view. anterior in vertebrae isolated of Comparison . individual. h rnvrepoessaepredclrt h long the to perpendicular are processes transverse anterior. the the transverse to situated slightly the directed In are ends crocodylians, dorsals distal their posterior-most In with the ribs. of the attach- processes ligamentous their of from at be ment may rugose rugosity are remains The with processes ends. vertebrae morphology transverse the broad their dorsals, PR long, isolated so FMNH the In (in damaged, uncertain. processes is these transverse 2298) of The The end slightly. narrow mid-dorsals. distal-most dorsals posterior the the of by processes anteropos- broad and longer teriorly become but narrow initially are are processes These facets process. transverse contact the capitular on located and both distal tubercular surfaces. The the contact d6, cylindrical. diapophyseal By circular nearly bear are surfaces 58) Figure 2298; d6 a and to d3 2298). between PR due vertebrae (FMNH centra dorsal the preserved on of lack present are parapophyses the h rnvrepoesso 1d FN PR (FMNH d1–d3 of processes transverse The oouhsterrestris Notosuchus hscniini o present— not is condition this , ¼ cm. 1 Downloaded By: [Columbia University] At: 05:34 17 September 2007 otro osl n h otro osl of dorsals posterior as of species the interpreted gomesii and vertebrae A. isolated dorsals but posterior 2298, PR dorsals posterior The 2005). of (Pol vertebrae the of axis 60. Figure ,Mddra etba MHP 35 ,Pseirdra etba MHP 34 ,Cua etba MHP 2308. PR FMNH vertebra, Caudal D, 2304. PR FMNH vertebra, dorsal Posterior C, 2305. PR FMNH Scale vertebra, Mid-dorsal B, terrestris ecie h otro alo h ersini which in depression the of wall now posterior of and the margin broader describes becomes anterior ridge dorsal the The past centra. the extend The to transverse the the failing dorsally. to and process, closer lies robust, face whole more a as and prezygapophysis and shorter become enlarge prezyga- processes the facets posteriorly, Moving a pophyseal spine. surface and neural lower the the process lateral onto of transverse a extending the ridge bear dorsal meet and distinct to trunk running the anterior ridge throughout the robust processes prezygapophyseal past are The centra. The project the of facets. barely margin dorsomedially-facing just oval zygapophyses with physes neiry h oslvrerehv prezygapo- have vertebrae dorsal the Anteriorly, .tsangatsangana A. ¼ m Poorpsb .Leonard.) C. by (Photographs cm. 1 nti respect. this in rrpscu tsangatsangana Araripesuchus Araripesuchus AN 45)sgetta hs two these that suggest 24450) (AMNH r orypeevdi FMNH in preserved poorly are lsl resemble closely oprsno sltdvrerei otro iw ,Atro oslvrer,FN R2306. PR FMNH vertebra, dorsal Anterior A, view. posterior in vertebrae isolated of Comparison . Notosuchus rgascu aznarezi Uruguaysuchus paradoxus respect, this unlike In horizontal. tsangatsangana truly A. become never they posteriorly, diminishes facets postzygapophyseal extent the the of orientation to oblique laterally the Although, extend crocodylians. not in seen do border but posterior centra, the postzygapophyses the beyond the of well of the expand margin and posterior, enlarge posterior Moving the past centra. just extend physes zygapo- These facets. ventrolaterally-facing with small in crocodylians. dorsals modern in posterior seen extent the tsangatsangana of A. rests. prezygapophyses vertebra preceding The the of postzygapophysis the rcse.Ti stecniinfudi xattaxa extant in found condition the is This processes. sangana of postzygapophyses the above, mentioned h otyaohsso h neirdrasare dorsals anterior the of postzygapophyses The gi,dfeigfo h aa mesoeucrocodylians basal the from differing Again, oouhsterrestris Notosuchus New r oiindhge hntetransverse the than higher positioned are W n us19b VPV24,and V8274), IVPP 1996b; Sues and (Wu Araripesuchus ssmlrt otnouhasand neosuchians most to similar is ontetn u aeal othe to laterally out extend not do Rsoi13)(iue60). (Figure 1933) (Rusconi Pl2005), (Pol rmMadagascar from Chimaerasuchus .tsangat- A. 307 Downloaded By: [Columbia University] At: 05:34 17 September 2007 i-oslvrer,FN R20.C otro oslvrer,FN R20.D adlvrer,FN R20.Scale 2308. PR FMNH vertebra, Caudal D, 2304. PR FMNH vertebra, dorsal Posterior C, 2305. PR FMNH vertebra, Mid-dorsal Poorpsb .Leonard.) C. by (Photographs Mo 91 n nerycooyoopssc as such terrestris Notosuchus crocodylomorphs early in Dibothrosuchus and 1921) (Mook 61. Figure 308 nag.I oencooyin (e.g. but them. crocodylians modern postzygapophyses dorsals In enlarge. the anterior between and the sacrum, the in sulcus toward shallow deepens very triangular is sulcus medial This a generally form the dorsals. to posterior due the of be size simply larger posteriorly. however, prominent may, the more This although slightly vertebrae, appear dorsal laminae the in reduced greatly processes. transverse the with level postzygapophyses are rmtemda hnlmn n h oslegsof edges dorsal 61). the (Figure and foramen is triangular sulcus lamina the the thin in medial fossa. spine the neural triangular from the small of exposure median only the the The posterior of contact midline The to the spine. down sulcus descends neural lamina thin the extend medial contact they to as dorsally laminae and postzygapophyses suprapostzygapophyseal the the from formed are sulcus the is morphology its in seen but tsangatsangana that from present different is significantly structure similar npseirve,tetopostzygapophyses two the view, posterior In but present are laminae Suprapostzygapophyseal In rrpscu tsangatsangana Araripesuchus .H Turner H. A. rrpscu tsangatsangana Araripesuchus W n hteje19) ntx uhas such taxa In 1993). Chatterjee and (Wu . and hmeauhsparadoxus Chimaerasuchus oprsno sltdvrerei oslve.A neirdra etba MHP 36 B, 2306. PR FMNH vertebra, dorsal Anterior A, view. dorsal in vertebrae isolated of Comparison . h aea al of walls lateral the , liao sinensis Alligator Araripesuchus ,the )a sntwl eeoe in developed lamina well thin not medial is suprapostzygapophyseal posterior The no present. are laminae because defined poorly Alligator iein formed, are structure like the of walls lateral The deep. very Fgr 0 eoe in becomes small 60) a in (Figure was foramen postzygapophyses—what triangular the spine neural the between of the extension full With of past two. spine, in neural sulcus the spine the by division, literal neural a in Indeed, is the happens postzygapophyses. is what the of it of laminae limit as posteromedial lateral lamina the the by of bisected Therefore, not tsangatsangana A. 1921). is (Mook sulcus crocodylians the in absent or eoemr rnucdi h otro dorsals. posterior and the spine in the pronounced on and present more Anterior spine. are become the laminae of thin rest thick- posterior the in to similar in compared like relatively ness flare remains not does sacrum. but spine crocodylians, the the of toward terminus dorsal moves The broaden one spines The as taxa. extant anteroposteriorly in seen those as tall as vertebra. the of posterior the on groove In h erlsie ftedra etba r about are vertebrae dorsal the of spines neural The liao sinensis Alligator oee,tedra deo h ucsis sulcus the of edge dorsal the however, , Araripesuchus nta,wa cusi h extension the is occurs what Instead, . h ucslk tutr snot is structure sulcus-like the , ytepszgppye.In postzygapophyses. the by , Alligator rrpscu tsangatsangana Araripesuchus Alligator n sotnreduced often is and ag “V”-shaped large a ¼ cm. 1 Downloaded By: [Columbia University] At: 05:34 17 September 2007 aiua n ueclrfct r iclrand circular are the preserved, facets and tubercular present and When remains. paucity capitular the rib to due of trunk the throughout morphology flange. similar a possessed d4 whether of unclear rib is the It flanges. both anterior distinct former three bear have first these ribs the of shafts ribs The two. with the of four longer processes, the being first tubercular and These capitular 2297. PR FMNH tsangatsangana identifiablefrom Fewdorsalribsarepreservedand/or ribs Dorsal This dorsals. anterior and posteriorly. the disappears prezygapophysis depression in the postzygapophysis between of surface the spine lateral the neural on present the is depression small A 62. Figure ,Mddra etba MHP 35 ,Pseirdra etba MHP 34 ,Cua etba MHP 2308. PR FMNH vertebra, Caudal D, 2304. PR FMNH vertebra, dorsal Posterior C, 2305. PR FMNH Scale vertebra, Mid-dorsal B, itecnb adgnrlzn h hnei rib in change the generalizing said be can Little ¼ m Poorpsb .Leonard.) C. by (Photographs cm. 1 rrpscu tsangatsangana Araripesuchus h iso 1d r rsre in preserved are d1–d4 of ribs The . oprsno sltdvrerei etltrlve.A neirdra etba MHP 2306. PR FMNH vertebra, dorsal Anterior A, view. lateral left in vertebrae isolated of Comparison . A. and ieypoie agratcmn ufc for surface attachment larger muscles. a but intercostal smooth provided most not This flange. are likely anterior-directed d4, broad of to short a surface posterior bear anterior ribs, The dorsal crocodylians. the in modern pronounced in as is of not it swelling is distal the shaft This than rib shaft. larger the in the slightly and of is circular head that portion end the preceding roughly flattened near a and broad to are taper curved dorsal distinct ribs the gently The a of cross-section. all shafts dorsals, are The posterior present. ribs not more is costal the tuberculum the ventrally-directed In as interpreted shallow, groove. Separating is a on that tuberculum. is depression lies triangular the processes typically than two capitulum these process The longer size. a in subequal rsn nboth in Present .gomesii A. New Araripesuchus AN 45)aesalconical, small are 24450) (AMNH .tsangatsangana A. rmMadagascar from FN R2298) PR (FMNH Araripesuchus 309 as Downloaded By: [Columbia University] At: 05:34 17 September 2007 iue6)cnit ftosca vertebrae—the crocodylomorphs. sacral most two of of condition consists conserved 64) Figure tsangatsangana A. diinly nsm pce of 1973). species Gasc some in and Additionally, (Hoffstetter originate of processes uncinate crocodylians the where is which shaft, rib the surface of posterodorsal the with) association in found (or swt other with As Sacrum 1921). (Mook slightly ossify to known are processes 36 ,Mddra etba MHP 35 ,Pseirdra etba MHP 34 ,Cua etba MHP 2308. PR FMNH vertebra, Caudal D, 2304. PR FMNH vertebra, dorsal Posterior C, 2305. PR FMNH Scale vertebra, Mid-dorsal B, 2306. htte“pks of “spikes” the fact the that is living interpretation in this present Supporting crocodylians. processes uncinate cartilaginous anterior to the homologous the likely are with spikes These vertebrae. associated dorsal “spikes” tipped acutely 63. Figure 310 ¼ .H Turner H. A. m Poorpsb .Leonard.) C. by (Photographs cm. 1 rrpscu tsangatsangana Araripesuchus Araripesuchus U 77 MHP 2301; PR FMNH 8767, (UA Araripesuchus ar,tescu of sacrum the sacra, Alligator oprsno sltdvrerei ih aea iw ,Atro oslvrer,FN PR FMNH vertebra, dorsal Anterior A, view. lateral right in vertebrae isolated of Comparison . r oae on located are h uncinate the n ue ihtescn arlvertebra. sacral second the straight with fused is centrum and the of surface anterolateral contact posterior centrum. the The the of the by parapophyses flanking largely the in of expansion formed surface. expanded concave is deeply concavity greatly a This is forms and centrum plane articular horizontal the anterior of the however, surface dorsals, of shape posterior and size the the roughly is vertebra this of centrum the distinct. of and spines unfused remain neural at vertebrae the and two centra though their at zygapophyses, both their other, in each to fused noted largely been therein). discussion has see however, 2005; this (Pol vertebrae to terrestris exception Notosuchus rare The h ih ieo h rtsca smsl isn.The missing. mostly is sacral first the of side right are The 2301 PR FMNH of vertebrae sacral The hc osse he ue sacral fused three possesses which , Downloaded By: [Columbia University] At: 05:34 17 September 2007 eta E iw Scale view. (E) ventral iue6.FN R2301, PR FMNH 64. Figure h ufc rao h cotyle. the of parapophyses. area of surface two-thirds the shifted nearly form posteriorly parapophyses the by Together, and part, enlarged in broad formed, very the being is narrow, It dorsoventrally 1921). and of that (Mook unlike posterior crocodylians is cotyle the modern The and cotyle. deep flat a is bears surface surface anterior is vertebrae The two the than similar. of length wider the much though first, is the of vertebra that sacral cen- second The the of groove. trum oriented anteroposteriorly shallow h epycnaeatro ufc ftefis sacral first the of surface anterior concave deeply the h armof sacrum The a by marked is sacrals both of surface ventral The .tsangatsangana A. ¼ m Poorpsb .Leonard.) C. by (Photographs cm. 1 rrpscu tsangatsangana Araripesuchus sitrsig with interesting, is ata armi neir() otro B,lf aea C,rgtltrl()and (D) lateral right (C), lateral left (B), posterior (A), anterior in sacrum Partial . otepeyaohsso h eodsca,though sacral, like second size in the reduced not of prezygapophyses fused the are first to the of postzygapophyses The preserved. pseudo- be, posterior opisthocoelous. would the in and articulation were pseudo-procoelous way, anterior condyle a the in a then of case, second the presence with fact the the articulation If of for vertebrae. condyle case these the developed the a of had and (in sacral) sacral) centrum vertebra first the the succeeding of case the the whether the of (in unclear posterior vertebra preceding the is on It cotyle deep second. the and vertebra h rzgppye ftefis arlaenot are sacral first the of prezygapophyses The New Araripesuchus oouhsterrestris Notosuchus rmMadagascar from Pl2005). (Pol 311 Downloaded By: [Columbia University] At: 05:34 17 September 2007 al ftecnrm epdpeso separates depression deep A centrum. the of lateral the of walls continuations as appear are which surface ridges, ventral two the of length entire the anteropos- along Running teriorly slender midpoint. and their long at rather constricted are and 2298 caudal PR FMNH preserved of All centra slender. more become the caudal, centra like second the broad the to are Posterior centra centra. dorsal caudal posterior two first the 24450), etba dnie,i ean nla h hp of In shape centra. the caudal unclear anterior remains caudal the it anterior identified, definitive vertebrae Without amphicoelous. are vertebrae. which respective of their two with 2298, associated PR chevrons FMNH three with preserved Only are only missing. with 1991). portion tail, known distal-most (Hecht complete the nearly vertebrae specimen a caudal reflects complete number 21 This preserves most it the and is 24450) sltdcua etbai eta iw Scale view. ventral in vertebra 2308, PR caudal FMNH Isolated 65. Figure any isolated for vertebrae Araripesuchus caudal five of number 59D– this, total The (Figures 65). to prepared 63D, fully PR addition and found FMNH In were caudals in 65). preserved (Figure are 2298 vertebrae caudal Nine vertebrae Caudal the poster- to enlarged fused greatly firmly the parapophysis. is iorly-shifted the and rib of process area sacral substantial transverse medial second the a The of in ilium. much with resulting surface a extension contact with broad is posterior rib attached long its and crocodyli- process most Like this 2301. forms, PR process FMNH in transverse preserved right is posterior the Only ribs. sacral and posterior pre- the the of arch, those dorsals. to neural similar the are on postzygapophyses location and size In Poorp yC Leonard.) C. by (Photograph 312 l adlcnr r mhpaynt weakly to amphiplatyan are centra caudal All ohsca etba aknua pns(iue64). (Figure spines neural lack vertebrae sacral Both expanded via ilia the to connected is sacrum The .H Turner H. A. pce sntknown. not is species rrpscu tsangatsangana Araripesuchus .gomesii A. .gomesii A. (AMNH (AMNH ¼ 1cm. . iia owa sse in seen prezygapophysis, is the what process of to the edge of similar posterior edge the anterior contacts The slightly. posterior 2298. PR FMNH in obscured are for most known because of not progression is height The spine present. neural are laminae thin anterior distinct caudals some Posterior in vertebrae. though absent, dorsal are the laminae thin of spines the as broad as h igs hsmrhlg ssmlrt htse in seen that to similar (e.g. crocodylians is modern morphology This ridges. the eoee a nioae etsauaadcoracoid and scapula Also left damaged. isolated complete. an margin scapula was left nearly anterior-most complete recovered the is the nearly only a of right has with right portion 2334 the a complete PR but only FMNH nearly is blade, and scapula scapular left scapula partial The a right coracoid. preserves 2297 and in PR intact FMNH left slabs. preserved the is of girdle any shoulder articulated No preservation and form girdle—general Shoulder skeleton Appendicular Description: view. demonstrates anterior in centra, “V”-shaped are characteristically chevrons caudal The the intersegmental. are the chevrons on the that of facets surface the articulation ventral with their conjunction directed in with anteroventrally This, position lack 66). “articulated” (Figure and roughly vertebrae in process are two transverse the presacral of the laminae. level dorsal suprapostzygapophyseal of located the those are to postzygapophyses like The similarly vertebrae. column—roughly are postzygapo- presacral The physes dorsomedially. the facing of facets circular those to gradually caudals length the caudals. but succeeding trans- the long, the in diminishes are caudals, five processes anterior-most verse the In 2005). amlac nltrlve.Scale view. lateral 2299, in PR arch FMNH Haemal 66. Figure .Leonard.) C. h rnvrepoessaenro n ln othe to slant and narrow are processes transverse The not are spines The spines. neural tall bear caudals The ftetrecern rsn nFN R2298 PR FMNH in present chevrons three the Of anterior the on similar are prezygapophyses The liao sinensis Alligator oouhsterrestris Notosuchus rrpscu tsangatsangana Araripesuchus ¼ c.(Photograph1cm. by .tsangatsangana A. ). (Pol . Downloaded By: [Columbia University] At: 05:34 17 September 2007 Scale oe tapaslkl httesnhnrssbetween synchondrosis the that likely appears There- it elements. two fore, the between one interdigiting any of lacking with sign smooth, relatively are articulate scapula ventral the and of that coracoid surface the coracoid of surface dorsal and The another. scapula consists 2313 a PR of FMNH specimen but articulation, scapula. with smooth. the is by surface coracoid, formed glenoid fossa The the the of one-third of roughly comprised predominately hmeauhsparadoxus Chimaerasuchus h ld a ebodri eea nsm basal some in general in (e.g. broader scapulocoracoid be although mesoeucrocodylians the can crocodyliforms, blade as other the most broad in as is articulation the blade about of scapular length generally The the articulation. to compared scapulocoracoid blade scapular broad scapula. the of edge anterior lacking the only from condition, good material in are These 67). (Figure 2313, PR FMNH 67. Figure osauao oaodi rsre nfull in preserved is coracoid or scapula No is and posteriorly opens fossa glenoid The h cpl srmral nisproportionally its in remarkable is scapula The ¼ m Poorpsb .Leonard.) C. by (Photographs cm. 1 hni crocodylians. in than ) rrpscu tsangatsangana Araripesuchus oouhsterrestris Notosuchus riuae etsauaadcrci nltrl()adpseir()view. (B) posterior and (A) lateral in coracoid and scapula left Articulated . , h w lmnswsoe nalpreserved all in open was elements tsangatsangana two the cplrbae oprdt h eta margin. ventral the to narrow seen compared In relatively the shape have blades this, the which scapular from crocodylians, from living Apart significantly in the 68). of differs wall (Figure scapula dorsal the fossa bears glenoid that expansion from ventral blade dorsal a broad a separating area constricted a with crocodylomorph, characteristically is scapula The Scapula for mesoeucrocodylians. condition other primitive for these the true holds if maturity Crocodylia that is, past that advanced indicates 1995), greatly (Brochu most not were at An individuals 2000). al. synchondrosis et Clark as open 1993, crocodylomorphs Chatterjee early and in (Wu seen well is and 1921) 1915, Mook (Reese crocodylians among condition pervasive itnto ewe cpl n oaodi the is coracoid and scapula between Distinction .tsangatsangana A. New Araripesuchus individuals. h ld ftesauais scapula the of blade the , rmMadagascar from 313 A. Downloaded By: [Columbia University] At: 05:34 17 September 2007 aaagscu insignis Mahajangasuchus concave strongly a form margin. anterior to convex, and anteriorly high steeply is sloping edge dorsal The broad. extremely 2313, PR FMNH 68. Figure Scale 314 h hp ftesauai eysmlrt htof that to similar very is scapula the of shape The ¼ .H Turner H. A. m Poorpsb .Leonard.) C. by (Photographs cm. 1 rrpscu tsangatsangana Araripesuchus U 64.Bclyand Buckley 8654). (UA etsauai aea A n eil()ve,soigmsl scarring. muscle showing view, (B) medial and (A) lateral in scapula Left . te eourcdlashv ra cple(e.g. scapulae broad have mesoeucrocodylians Other o nunmdcaecontaining clade diagnostic as unnamed articulation twice scapulocoracoid an blade for the scapular as a wide regarded (1999) Brochu Araripesuchus ersuia and Peirosauridae , Mahajangasuchus Trematochampsa . , Downloaded By: [Columbia University] At: 05:34 17 September 2007 ertsmsl cr hc slkl h attachment (Fu the posterior scapulohumeralis likely M. is the for which point superficial scar, the from muscle distinct serratus scar, ovular scapula, small the is in superficialis. constriction there serratus the near M. and this the to Ventral of attachment the from hs aateepnino h cplrbaei o as not is blade in scapular as extensive the of expansion the taxa these swl ssm aa liaoodcrocodylians alligatoroid basal some as (e.g. well as nitntrde(r)sbiie h ufc fthe of surface the The and subdivides sulcus. low scapulae. posterior (sr3) very second A ridge dorsalis broad. partial but indistinct an shallow M. is a imply sulcus the may posterior forms sulcus of and deep enlargement This deep anterior ridge. a The very longitudinal (sr2). with is glenoid continuous the nearly sulcus above and ridge (sr1) smaller distinct very is coilceti rsn neiry lhuhits although an anteriorly, whether present unclear in is is presence It crest glenoid. dorsal acromial the the forming of and one-third coracoid the contacting base figured also (1996b) blade. Sues scapular and paradoxus figure Wu Chimaerasuchus not did presence. (2005) Pol its but obs.) (pers. ridge subtle otegeod(o 05 speetbti is it but present is in as 2005) pronounced (Pol as glenoid not the to hnaeaeridge. distinct more average a posses than not blade. do An than scapular crocodylomorphs broader other the scapulae directly 2). have of be that crocodyliforms to expansion Plate Other not an 1999; appears with ridge correlated Brochu this of and enlargement longitudinal (Buckley prominent relatively ridge a has indeed which rs iia owa sse in In seen 68A). is (Figure what to insignis similar crest likely. presence h taheto .trsmjradM dorsalis a M. simply and in major is condition teres it (Fu M. as from scapulae of formed subtle attachment sulci very shallow the the is between ridge, posses which demarcation crocodylians longitudinal ridge, Living this prominent surface. the its subdividing for noteworthy of identification rendering problematic. scarring specimens muscle is all scapula in the of missing margin anterior-most The 1876). ger AN 45) Again, in extent 24450). lesser a to (AMNH and 2000) al. et (Ortega aiyi cplrmrhlg with morphology scapular in larity oouhsterrestris Notosuchus hmeauhsparadoxus Chimaerasuchus h otro agno h cpl ssihl rugose slightly is scapula the of margin posterior The iial itntrdei rsn in present is ridge distinct similarly A h aea agno h cplrbaeis blade scapular the of margin lateral The etal,tesauaepnsit rectangular a into expands scapula the Ventrally, oslt h lni,teei distinct a is there glenoid, the to Dorsal Wannaganosuchus U 64 u otnigfrhrdorsally farther continuing but 8654) (UA ¨ .tsangatsangana A. .patagonicus A. bigr17) otatn hsi the is this Contrasting 1876). rbringer Araripesuchus oouhsterrestris Notosuchus , oouhsterrestris Notosuchus aaiuhsmwakayasyunguti Malawisuchus and and Araripesuchus .tsangatsangana A. Diplocynodon and ihaltrlysmooth laterally a with and rgascu aznarezi Uruguaysuchus nti ao,teridge the taxon, this In . Mahajangasuchus .gomesii A. Mahajangasuchus Mahajangasuchus rs dorsal crest a , .patagonicus A. ,hwvr in however, ), hrssimi- shares a very a has .gomesii A. ae its makes Living . ¨ rbrin- . ) , , hf nteatcmn fteM rcp scapularis of triceps glenoid M. oriented Araripesuchus posteriorly the more of the reflects attachment likely the the posterior which in This in 1992). in shift (Brochu crocodylians Additionally, laterally faces in rugosity glenoid. condition the the trasting to tsangatsangana A. dorsal ridge and .tiessauai Fue 89 rcu20) In 2003). tsangatsangana Brochu A. 1879, (Fauvel scapularis (pers. of triceps attachment ridge-like glenoid M. the with the associated a is above rugosity This although presence obs.) is ridge, tubercle this rugose lack crocodylians h xlso fatoposau of the exclusion of with neosuchians that the all for scapula than synapomorphy subequal a coracoid greater a and considered slightly (1994) is Clark length scapula. the coracoid Indeed, the long. very number of is a coracoid in The ways. crocodyliform of typical the from differing eprcs eta otegeodthat glenoid the the to have sangana ventral however, taxa, recess these of deep None 16). fig. 1933: oae in foramen terrestris suchus the for point to likely attachment seems (Fu coracoscapularis the This ancones swelling. M. to small the very correspond On a edge, posterior is subscapularis. straight there the M. along point, the ventral-most to corresponding ml oaodfrmn iia otecniinin aznarezi condition Uruguaysuchus the a paradoxus to a in by similar perforated Chimaerasuchus scapula foramen, is 69). area the coracoid This line. small (Figure contacts straight less region distally or more dorsal expands wide narrow enlarged that The a an by shaft of ventrally tubular followed comprised region gracile, glenoid dorsal is coracoid The Coracoid Pl20)and as 2005) well posterior-facing as (Pol 1996b), enlarged Sues in and of (Wu An noted half was proximal shaft. facet the glenoid bordered down coracoid is fossa is medial the glenoid glenoid the which from the extending the recess, of lamina edge deep thin under a very by area medially a This by dorsal shaft coracoid. marked the the the from is along separated portion of completely ventral swelling the and while small pendulous scapula, the a with is contact poster- is faces There facet and glenoid iorly. concave, the slightly of expanded, surface greatly The fossa. glenoid region. the glenoid the to close very situated is 2313) MCP 6) and 269), PV (MUC h eilsraei akdb epdepression deep a by marked is surface medial The h oaodof coracoid The h oaodcmrsstevnrltotid of two-thirds ventral the comprises coracoid The oouhsterrestris Notosuchus New Fgr 9)or 69C) (Figure .gomesii A. . Araripesuchus Pl20) lolk hs aa the taxa, these like Also 2005). (Pol a elas well (as h uoiyfcspseir con- posterior, faces rugosity the rgascu aznarezi Uruguaysuchus .tsangatsangana A. AN 24450), (AMNH .tsangatsangana A. ,ti uoiyi rsn nthe on present is rugosity this ), Rsoi13) and 1933), (Rusconi .gomesii A. hmeauhsparadoxus Chimaerasuchus W n us1996b), Sues and (Wu rmMadagascar from s fdrsuisand dyrosaurids If rs. aaagscu insignis Mahajangasuchus ¨ bigr1876). rbringer oouhsterrestris Notosuchus has. .patagonicus A. sinteresting, is FN PR (FMNH .tsangat- A. (Rusconi Noto- 315 Downloaded By: [Columbia University] At: 05:34 17 September 2007 crig Scale scarring. h 99 rcue l 02.Tepeec fthe of presence The non- 2002). al. within a trait et Brochu as 1999, chu Bro- optimized and than (Buckley this) is synapomorphy of mesoeucrocodylian discussion character for 2002 this al. et Brochu 1994; Pholidosaurus 2313, PR FMNH 69. Figure nagdadpoone etal iein of In expansion the end. like is distal interesting the Most ventrally facet. the is under pronounced facet tsangatsangana glenoid A. and The shaft. tubular enlarged narrow the have eemnto ean qioa.Itrsigy the Interestingly, so of damaged coracoid equivocal. is coracoid remains the of determination end distal the taxa both 316 ale n oegaulythan gradually more and earlier h etrsdsusdabove. discussed features the togyepne n uvsatrol.Aan it Again, in present paradoxus is anteriorly. is morphology curves edge this appears and anterior this expanded asymmetrical—the however, crocodyli- strongly most crocodyliforms, is in most expansion case Unlike the is forms. than as gradually expanded, more distally and tsangatsangana earlier shaft A. the broaden taxa length. to both its in begins but of morphology, this much approach for tubular paradoxus Chimaerasuchus nearly and narrow in earlier occurs Neosuchia. than feature Mesoeucrocodylia the scenario unconstrained lhuhln,tecrci of coracoid the long, Although h hf ftecrci sas itnta tis it as distinct also is coracoid the of shaft The .H Turner H. A. and .gomesii A. Araripesuchus ¼ m Poorpsb .Leonard.) C. by (Photographs cm. 1 r eoe rmaayi seClark (see analysis from removed are rgascu aznarezi Uruguaysuchus h eta n ftecrci is coracoid the of end ventral The . oee,i ak h eprecess deep the lacks it however, , .gomesii A. ifr from differs rrpscu tsangatsangana Araripesuchus ugssta vnudran under even that suggests and h xaso begins expansion the , rgascu aznarezi Uruguaysuchus .tsangatsangana A. .gomesii A. .tsangatsangana A. Chimaerasuchus oee,in however, , Hsisosuchus osnot does etcrci nltrl() eil() n otro C iw hwn muscle showing view, (C) posterior and (B), medial (A), lateral in coracoid Left . in . in projecting within suchus and variation further pronounced Whether posteriorly. strongly edge the with posterior asymmetrical, not is expansion the Moreover, h rcp.Ti sucer u h aiaroughly in lamina triceps the of the head this but of origin unclear, the with topologically is corresponds This triceps. of coracoideum and caput the the enlargement tubercle for an surface attachment this reflect the may of to glenoid the brachii. dorsal under triceps located the lamina of prominent head long The the to appears coracoid correspond above the to described to tubercle attach The brachii 1879). triceps (Fauvel three M. the the of of two heads crocodylians, In 1879). Fauvel 1876, h .tiesbahise ncooyin (Fu crocodylians for in seen surface brachii attachment triceps M. the the to located corresponds Most likely tubercle shaft. the This of 8741. large margin posterior the UA moderately down halfway roughly and the 2313 is prominent PR FMNH coracoids, ftelrebcp rci uce(Fu muscle brachii anterolaterally biceps attachment large the large the from of formed a tubercle rugose have located which modern Unlike crocodylians, area. attachment coracobrachialis the costicoracoi- the muscle. the deus of received likely a remainder This is the margin. tubercle, coracoid down triceps extending the sulcus to shallow ventral but rim, posterior rcu1992), Brochu eea pcfi ucesasaevsbeo w fthe of two on visible are scars muscle specific Several nltrlve,wa titosdfieteotieof outline the define striations weak view, lateral In .patagonicus A. xssi neti,a ocrci spreserved is coracoid no as uncertain, is exists Alligator .tsangatsangana A. . Fue 89.As ln the along Also 1879). (Fauvel ak ueceand tubercle a lacks ¨ bigr1876, rbringer ¨ rbringer Araripe- Downloaded By: [Columbia University] At: 05:34 17 September 2007 .0wsslce eas aatpclyvee as viewed and typically of 0.10 taxa value (e.g. This because the higher. “gracile” selected or less 0.10 was is is 0.10 value a length the if midshaft considering long-axis between “robust” am ratio and its I if length “gracile” convention, be a to humerus As while taxa. morph importance, phylogenetic elements little importance. skeletal descriptive of certain retaining of be gracility the may result, clades. a specific distinc- of As context arbitrary the is within this diminish “robust” may although tion and arbitrary, “gracile” between likely draw most line 0 a between 1, continuously the considered and ranges Since is above “robust”. discussed what considered ratio is between what exactly line and is the “gracile” ambiguity draw of to point case where this Another element—in humerus. particular the the a be and of width to midshaft length taken or between long-axis here ratio robustness is the of and in reflected independent, degree size the reflection is However, a gracility as size. robust their more appear of be to larger tend have can and will animals bones larger gracility as determine to Humeral difficult crocodylians. modern Araripesuchus h ueiof humeri The Humerus II. digit of II, phalanges MC three as the interpreted and three IV, are slab were III, These on 2328). elements another PR one (FMNH podial with associated of Additional half preserved distal as the radialia, metacarpal. with right ulnare and two a radiale radius, right right a a a as of of well half two-thirds distal proximal the the ulna, was slab isolated that this Recovered phalanges manus. from the various ulnare, with as associated radiale, be well radius, may as a carpal of pisiform right portion and distal the consists isolated ulna, 2324, on an PR an of nearby FMNH near forelimb, located second to a The 2327) slab. exposed. PR pertain and (FMNH surface may end) humerus posterior forelimb proximal the This with (the ulna radius complete obscured, end partial distal its 2324 with a humerus PR left FMNH a of recovered. consists were 2324, 2326) PR PR (FMNH elements forelimbs FMNH podial partial Two and The scarcer. Epipodial are E). 2298, 2325–2327). (Appendix PR predominantly PR intact and preserved FMNH FMNH well typically are 2297, 2302, humeri PR PR the FMNH FMNH of recovered 8721, were any humeri right in (UA four preserved and left was Three slabs. forelimb complete No preservation and form Forelimb—general 8741. either UA on or apparent 2313 is PR attachment FMNH biceps for region no rclt ftehmrsvre mn crocodylo- among varies humerus the of Gracility aa r eygaiecmae to compared gracile very are taxa, .tsangatsangana A. ersrscu gracilis Terrestrisuchus swl sother as well as , )fallbelow neiroe sntdb o 20)on (2005) the Pol than by longer noted As and one. larger anterior depression depressions—with small posterior form, the ridges partially These and surfaces. run enclose, posterior and ridges on anterior humerus the supracondylar the both of end two distal the 2000), near proximo-distally al. et Ortega pce ( species Wle 1990), (Walker ta.20)a ela in as well as 2000) al. et h eaiesz n oiino h rs ssmlrto similar is crest the gracilis profile. of Terrestrisuchus position anterior and concave size a relative The and profile lateral crest. the convex of apex are anterior the There on and humerus. proximal head the the near humeral crest roughly the of to dorsal lateral distally length found tuberosities the extending the of humerus, on one-third the located is of and crest margin deltopectoral smooth, The generally 1991). thin, Hecht obs.; pers. aznarezi hf iemdr rcdlasehbt similarly A in exhibit. noted was crocodylians humerus mid- straight modern bend lateral like the nearly shaft lacks and narrow, humerus straight, The crest is cylindrical. deltopectoral shaft The prominent 70). a (Figure bears and ends both in seen that to similar humeri robust have ezi oee,hsgaiehmr oesmlrt the to similar 1999), more Pol humeri in 1997, seen gracile Gomani dimensions has 1996b, however, Sues and to (Wu related closely as regarded cally mwakayasyunguti Malawisuchus aaagscu insignis Mahajangasuchus aznarezi Uruguaysuchus paradoxus Chimaerasuchus ftehmrlha in head surface humeral posterior the the surfaces. of on anterior found and are posterior depressions Similar the both on end proximal- most the posteriorly below just slightly depression ovular slants an bears and head humeral The surface. obs.). pers. of 2005; (Pol crest shifted medially strongly insignis 32 eeln eea neetn etrs sin As PR FMNH features. and interesting 8721 terrestris several UA Notosuchus in revealing preserved 2302, well is but 93,and 1993), aiswl bv .0 o 20)ntdta some that aznarezi noted Uruguaysuchus (2005) as Pol such mesoeucrocodylians 0.10. basal above well ratios “robust” as viewed typically taxa “ (e.g. while value, this and uei rcdlaslike Crocodylians humeri. nltrlve,teceti raglri hp iha with shape in triangular is crest the view, lateral In h ueu of humerus The h rxmlha swdrta h itlarticular distal the than wider is head proximal The h itledi aae nms rsre humeri preserved most in damaged is end distal The al rcdlmrh uhas such crocodylomorphs Early Rsoi13:fi.1)adother and 11) fig. 1933: (Rusconi Crocodylus rcdls robustus Crocodylus” BclyadBoh 99,bti niethe unlike is but 1999), Brochu and (Buckley New .gomesii A. Rsoi13:fi.1)and 12) fig. 1933: (Rusconi Terrestrisuchus Araripesuchus Mo 91 aerbs ueiwith humeri robust have 1921) (Mook Dibothrosuchus AN 24450; —AMNH Terrestrisuchus Pl2005), (Pol .tsangatsangana A. oouhsterrestris Notosuchus Cuh1984), (Crush and Rsoi13:fi.1) and 12), fig. 1933: (Rusconi BclyadBoh 1999). Brochu and (Buckley alaoeti value. this above fall ) .gomesii A. Cuh18)hv gracile have 1984) (Crush Alligator hmeauhsparadoxus Chimaerasuchus W n us1996b), Sues and (Wu rmMadagascar from .patagonicus A. rcdlfr typi- crocodyliform a , rgascu aznar- Uruguaysuchus W n Chatterjee and (Wu oouhsterrestris Notosuchus and oouhsterrestris Notosuchus oouhsterrestris Notosuchus FN 197946) (FMNH Mahajangasuchus .patagonicus A. AN 24450 (AMNH Araripesuchus sepne at expanded is Uruguaysuchus Sphenosuchus Araripesuchus Notosuchus Pl2005), (Pol Crocodylus (Ortega 317 — . , . , Downloaded By: [Columbia University] At: 05:34 17 September 2007 h eilrdein ridge medial the Scale views. (F) 2302, anterior PR and FMNH 70. Figure aea ufcso h itlpr ftehmrsare humerus the of in as part flat, distal and broad the The of lateral surface. surfaces posterior the the lateral on while prominent surface, more is anterior ridge the on prominent 93 g 1 n other and 11) fig. 1933: rdmnnl aeatrol,o nlf position, life ground. the to in In parallel held or is humerus anteriorly, ventrally—the face predominantly posterior the and by humerus. anterior the formed the of are is surfaces on depression which condyles ridges the trochlea, two supracondylar deep with The moderately continuous a surface. by farther articular separated slant medial extends the to anterolateral convexity to Although slight a convexity. broader medial creating distally, and medial the larger the smaller, is than capitellum anteriorly The developed. 2000). al. et Ortega 24450; (AMNH aeatrol u sdflce etal truhya roughly at ventrally deflected 45 is but anteriorly face 318 rcdlas hrfr,sgetn oeupright more for living a posture than forelimb suggesting pronounced Therefore, more is crocodylians. but face ventral, condyles articular directly humeral whose 1984), crocodylomorph (Crush basal nlvn rcdlas h riua surfaces articular the crocodylians, living In equally not are condyles articular two The 8 .tsangatsangana A. nl.Ti snta eeeacniina nthe in as condition a severe as not is This angle. .H Turner H. A. rgascu aznarezi Uruguaysuchus .tsangatsangana A. h riua ufc osnot does surface articular the , ¼ .tsangatsangana A. rrpscu tsangatsangana Araripesuchus m Poorpsb .Leonard.) C. by (Photographs cm. 1 e rsrscu gracilis Terrestrisuchus Araripesuchus . (Rusconi smore is species ethmrsi rxml() itl() aea C,pseir() eil(E), medial (D), posterior (C), lateral (B), distal (A), proximal in humerus Left . odro h etpcoa rs Fgr 1.Teright The 71). (Figure crest posterior deltopectoral the the on of major border teres the for scar attachment the etpcoa rs sietfidhr stemuscle the the major. as teres on M. here the depression identified for scar is rugose crest the deltopectoral muscles, topographical humer- and close deltoid, oradialis triceps, the the Given with correspondence 2003). (Fauvel brachii Meers triceps the the between 1879, of area heads medial the likely and with is lateral just associated ridge scar This terminates muscle humerus. ridge the the of The midpoint gently the path. low curving past its a diaphysis, along is the depression anteriorly along rugose extending the deltoideus of ridge border M. ventral the of insertion 2003). proximal Meers 1992, (Brochu the scapularis as here interpreted rugosity this elongate an Bordering is head. proximally, humeral rugosity, the of deltopectoral back the the on near crest located is posteriorly, depression present rugose is a but tubercle No aspects. of number of 2003). head gana humeral Meers the on 1992, scarring (Brochu Muscle crest from the head deltopectoral near the of the head surface humeral posterior the the delimiting of angle surface position) (dorsal life posterior the in on tubercle a by indicated is major All from distally projecting and insertion this Opposite teres M. of insertion the crocodylians, in Ancestrally ifr rmtecniinse ncooyin na in crocodylians in seen condition the from differs Araripesuchus aawt ecie ueibear humeri described with taxa .tsangatsan- A. Downloaded By: [Columbia University] At: 05:34 17 September 2007 yC Leonard.) C. by hteje(93 oe ersini similar a in depression left a in the location noted on (1993) and depression Chatterjee and Wu crest Additionally, ovular 12). fig. 1933: deltopectoral (Rusconi an humerus the figured on Rusconi scar muscle similar of humerus Scale of 8721). view (UA distal of Medial humerus view B, right Lateral C, of Posterolateral 2325). 8721). condyle (UA PR A, humerus (FMNH right scarring. of head muscle condyle distal humeral humeral left of of Detail view 71. Figure tahetsraei oae oepoial in proximally more located tsangatsangana is A. This surface 2003). of (Meers attachment insertion caudalis the humeralis to scapularis M. corresponds concavity deep which a 71A), bears (Figure condyle, proximal the to distal hr sa nagdrgo o h tahetof attachment the for (Fu region brachioradialis enlarged M. an is there 89 rcu19,Mes20;Fgr 71C). Figure 2003; Meers ridge 1992, distinct Brochu a (Fauvel arm bears the 1879, of muscles surface flexor the condylar to corresponding the of edge h otro ufc ftehmrlha,just head, humeral the of surface posterior The itly nteatro ufc ertecondyles, the near surface anterior the on Distally, Dibothrosuchus rgascu aznarezi Uruguaysuchus hni xatcrocodylians. extant in than ¨ bigr17) h medial The 1876). rbringer . per opse a posses to appears ¼ m (Photographs cm. 1 lcao,bttepeec fti structure an this of of rudiments presence the the in like but looks olecranon, what preserves ftepoia end. proximal portion FMNH posterior the the but preserve of damaged, 8758 UA slightly and is PR2339 end proximal The lvto fteae vrtesurrounding the crocodyli- over Other as present. such area forms, appears the surface articular of elevation lahsasot rcla ufc,aportion a surface, in as trochlear anteroventrally deflected smooth is which a The of 72). has (Figure preserves bone ulna, the ulna right of a two-thirds proximal 8758, the UA 2326–2327, 2339). to PR PR FMNH belonging FMNH 2324, PR right) FMNH 2298, three PR and left tsangatsangana A. (three ulnae Six Ulna digitorum flexor the and the teres muscles. This between longus pronator distinction nerve. the a of ulnar heads represent the also of may branch groove may a surface groove this transmitted the but through have ambiguous, passing is groove It deep proximodistally. a is There n rcu19,Bclye l 00 place 2000) al. et Buckley 1999, Araripesuchus (Buckley Brochu analyses phylogenetic and Some olecranon. an paradoxus Chimaerasuchus wihpsessa lcao)ta to than olecranon) and an possesses (which Fgr 3.Tedsa n fteun ssmall is ulna 2326 the anterior The PR of end). proximal the end FMNH of size distal the half in The (about 73). edge (Figure and 2324 posterolateral PR FMNH in the exposed is surface medial the the and 8758 for UA making 2326. scar both PR smooth in FMNH A absent the is difficult. is ligament shaft scars lacks interosseous muscle the crocodyli- and of of other delineation narrow surface many The is in forms. The found shaft muscle. bowing the prominent supinator of the remainder for an providing surface of striated part and attachment anterior rough is the region humer- of trochlear edge the M. lateral The of of 2003). quadratus Meers origination pronator 2003; the (Brochu oulnophalangei marking 2326) PR striated, FMNH to 8758, is ventral (UA just region “olecranon” surface to the posterior proximal the The due the toward region. of dorsally articular is flares shaft and narrow The proximally is ulna preservation. narrowness during this compression of some rock the 2003), in preserved (Brochu be record. necessarily olecranon not would cartilaginous which from crocodylians a inference modern possess on mature based Moreover, olecranon phylogeny. an possess may h itlatclrsraei beti A8758, UA in absent is surface articular distal The although narrow, mediolaterally is ulna the Overall, .tsangatsangana A. Chimaerasuchus New Araripesuchus lsrto closer oouhsterrestris Notosuchus eercvrd(A85,FMNH 8758, (UA recovered were ugsigthat suggesting , .gomesii A. W n us19b,lack 1996b), Sues and (Wu sabgosaslight ambiguous—a is aaagscu insignis Mahajangasuchus rmMadagascar from AN 24450) (AMNH Pl20)and 2005) (Pol Araripesuchus Notosuchus .gomesii A. 319 . Downloaded By: [Columbia University] At: 05:34 17 September 2007 h itled h itledi nldslightly angled the is in seen end of of extent the distal that axis to The with not long but parallel end. anteriorly, The is distal surface ends. the articular both proximal and at mid-diaphysis at expanded cross-section in elliptical bone, h aisbnssihl eilyaa rmthe of that from to of similar away radius of more the medially is shaft respect, this slightly the In margin, bends ulna. to rod-like anterior radius prior and the but the on end slender proximal tuberosity is the the Nearing radius 74). the (Figure shape, general oencooyin.Terdu n laof ulna and radius The Dibothrosuchus crocodylians. modern R23)adtrergt(MHP 2298, outline, PR In 2327). (FMNH PR right of FMNH radius three the 2324, PR FMNH and 2329, FMNH PR 2339) PR FMNH (FMNH 2326, PR left PR FMNH recovered—four 2298, were radii Seven Radius make distal anterior radiale the the of the and surface ulna. medial radius of the distal with surface The contact bone. posterolateral the the from dorsal of posteriorly than out surface pronounced projects the and more convexity is anterior is convexity in condition seen posterior that paradoxus This like exactly groove. suchus not but shallow convexities to a small similar by by formed are separated edges posterior and 8758, UA 72. Figure Scale 320 ¼ .H Turner H. A. m Poorpsb .Leonard.) C. by (Photographs cm. 1 .tsangatsangana A. eani ls rxmt o most for proximity close in remain rrpscu tsangatsangana Araripesuchus W n us19b.The 1996b). Sues and (Wu Dibothrosuchus sarltvl simple relatively a is .tsangatsangana A. hni sto is it than Alligator ata ih lai eil() otro B,ltrl() n neir()views. (D) anterior and (C), lateral (B), posterior (A), medial in ulna right Partial . Chimaera- .In a orsodt h itlms xeto the ulnor- triangular smooth M. of a as for expressed extent and insertion distinct is distal-most the adialis Distally, which the scar. radiale, is to the “ventral” area for correspond roughened facet contact may slightly the humeroradialis and near present shallow the a by in in present although be to intersected present not appears This was not brevis. of that medial is the scar” insertion humeroradialis for the the tsangatsangana A. ridge of for A brevis. head margin ridge humeroradialis This lateral (rs2). M. diaphysis the the of marks surface lateral the orsod otelcto o h neto of insertion the radius, for the location (Fu of humeroradialis This the M. half margin. to posterior proximal the along corresponds the present is on tubercle a still but to Distal (rs1). this, present humeroantebrachialis is brachii M. biceps M. of and inferior tendon the common scar of the the surface, surface articular for the lateral to the inferior just on and radius Proximally ulna. the mildly the facet is for proximal the surface of for ventrally this deflected condyle of and concavity—a expanded margin radial mild lateral the The a humerus. to bears corresponding surface surface This outline. in and (Wu radius the of crocodylians, bending 1993). modern medial Chatterjee in the as as bowed well as strongly as not is ftedsa w-hrso hi egh.Like lengths. their of two-thirds distal Dibothrosuchus the of natrpseirysiaigrdei oae along located is ridge spiraling anteroposteriorly An h rxmlatclrsraei ogl rectangular roughly is surface articular proximal The , .tsangatsangana A. rcu(92 ecie “ventral a described (1992) Brochu . ¨ bigr17,Mes2003). Meers 1876, rbringer a nun that ulna an has Araripesuchus , Downloaded By: [Columbia University] At: 05:34 17 September 2007 yC enr.Poorpsi yMhjnauhsEllison.) Mahajangasuchus by B in Photographs Leonard. C. by A hstedmyb u otelc faunique a of lack the to due be suggested may (1992) trend Brochu this the to carpus. given been crocodylomorph has attention little very Historically, Carpus tuberosity small humeroradialis. and M. crest of contact low insertion a the the border is marking anterior there to the visible radius near the Distal and of impress humerus radius. the not for the surface do upon brevis correlates humeroradialis and medialis M. humeoradialis M. of margins medial The surface radiale. entire the The anterior. contacts partially the to angled margin and be convex to mildly appears anterior is surface 2329—its the Nevertheless, broken. PR (Brochu damaged FMNH slightly crocodylians is in in surface articular as distal The bone, 1992). the in depression Scale 2339). PR (FMNH complete radius Nearly the B, of 2324). surface PR proximal (FMNH the pisiform as and well radiale as ulnare, damaged ulna, humerus radius, of left obscured end Partially Distal A, forearm. morphology. right Forearm 73. Figure h eilsraeo h aisi esinteresting. less is radius the of surface medial The n ppdas(iue7) ieteeioil,the epipodials, the Like 73). ulnare (Figure complementary epipodials its and with the preserved 2310), PR being FMNH latter 8737, (UA right two and 2324) tsangatsangana A. Crocodylomorpha. proximal of synapomorphy elongate a of Clark as and presence carpals Benton the (in considered Clark 1988) and (1970) Walker In to crocodylomorph. fact, back ancestral the recalls of habit crocodylians crocodylo- cursorial the modern the and in radiale found way, forelimb elongate ulnare The Either interesting. the is muscles. carpus of morph pelvic descrip- Romer’s of description of the tion that early to by comparable an musculature driven of perhaps forelimb—an absence broader crocodylomorph phenomenon, a the from interest the unfortunate in stem interest of to could of however, lack comparable lack carpus, wrist The the in the complex. in astragalocalcaneal joint intracarpal orrdai eercvrdprann to pertaining recovered were radialia Four New Araripesuchus tolf U 76 MHPR FMNH 8736, (UA left —two rmMadagascar from ¼ m Poorp in (Photograph cm. 1 321 Downloaded By: [Columbia University] At: 05:34 17 September 2007 xesr o ii I h eilrgst (rds2) rugosity medial The the II. lies of origin digit (rds1) the marks for and rugosity facet extensors ulna lateral the to The adjacent margin. proximal is ulnare. that and the point small for acute facet distal the disposed an intersects ventrally The to and ulna. tapers anterolaterally facet directed the the for of a and is portion facet there Laterally facet, edge. rectangular radius the lateral to roughly angle the right than a facet at level extending the of lower shaped edge a “boomerang” medial at the is largest, facet with surface concave, the mildly The The and is radius. surface, surfaces. the proximal for articular entire the three comprising proximal has The 75). (Figure end element slender a is radiale Scale view. (F) 2329, distal PR and FMNH 74. Figure 322 natro iw w uoiisaepeetna the near present are rugosities two view, anterior In .H Turner H. A. ¼ m Poorpsb .Leonard.) C. by (Photographs cm. 1 rrpscu tsangatsangana Araripesuchus etrdu nmda A,pseir() aea C,atro D,poia (E), proximal (D), anterior (C), lateral (B), posterior (A), medial in radius Left . ailaaesotrta htof that than stouter their although are forms radialia these in seen is sulcus similar a and hs w uceatcmn onslecoe together closer in I radiale, lie than points the digit attachment of muscle for two slenderness these the carpophalangei Given extensor 1992). carpometa- (Brochu M. extensor and M. of carpus attachment the indicates and for known are Radialia ii I n h aisrdaelgmn Boh 1992). (Brochu ligament radius-radiale the and II, digit flexor of carpometacarpals flexor and M. I, carpometacarpus digit of carpophalangei flexor M. the the mark of surfaces side, attachment These either (rds3). On rugose ridge. slightly median is bone a by divided is shaft h otro ufc sls ope.Teproximal The complex. less is surface posterior The hmeauhsparadoxus Chimaerasuchus Alligator eutn naslu ewe h two. the between sulcus a in resulting , oouhsterrestris Notosuchus W n us1996b), Sues and (Wu .tsangatsangana A. Pl2005) (Pol . Downloaded By: [Columbia University] At: 05:34 17 September 2007 apleeeti urnl nnw for unknown that currently although is centrale, element carpal the contacted likely than more itly h riua ufc ssihl ocv,a in radiale as Right concave, view. slightly (E) paradoxus Chimaerasuchus is proximal and surface (D), articular lateral the (C), Distally, Scale view. anterior (I) (B), lateral medial and (A), (H), posterior anterior in (G), medial 2310) (F), PR posterior (FMNH in radiale 8736) Right (UA 75. Figure ocv o t otc ihtepsfr carpal. or pisiform more a the in slightly pisiform with the the very preserves contact than 2324 is PR medial its FMNH wider surface and for is proximal concave lateral surface The the distal proximal. without The to but shape processes. in radiale narrow similar is slightly—very the flares diaphysis end the The each simple, radiale. and the relatively of is length element the two-thirds than (UA slightly longer ulnare are ends right proximal large 76). of and (Figure a radiale damaged distal and a whose the 2324, 8759) 8736, complementing PR UA ulnare FMNH of right radiale the small complementing ulnare h laei hre hnterdae slightly radiale, the than shorter is ulnare The left small a recovered, were ulnaria three Only and oouhsterrestris Notosuchus Araripesuchus ,and . rprinlysmlrt htse in forelimb, seen the that of to length total similar the stouter proportionally of be to less tend comprise taxa these and in ulnare and radiale The and 1933), (Rusconi h apshsbe ecie nfwohrsmall- other few doxus in mesoeucrocodylians— described been bodied has carpus The h oegaieadeogt ap eni earlier in seen of carpi that elongate especially and crocodylomorphs, gracile more the esatcltdsaewt h lae itet no of to forelimb Little pisiform. the ulnare. in seen the be can with however, detail, state articulated less apsof carpus uadCatre 93.Mroe,terdaeand radiale the Moreover, 1993). Chatterjee and Wu opooy smr ugsieo h apsof carpus the of suggestive more Dibothrosuchus is slender and taxa. morphology, elongate aforementioned overall their the with combined than This, length forelimb the of iemn te set ftesole and shoulder the of aspects other many Like W n us1996b), Sues and (Wu ¼ New m Poorpsb .Leonard.) C. by (Photographs cm. 1 Araripesuchus .tsangatsangana A. Araripesuchus or rhscu stormbergi Orthosuchus oouhsterrestris Notosuchus opie agrproportion larger a comprises rmMadagascar from rgascu aznarezi Uruguaysuchus h apsresembles carpus the , hmeauhspara- Chimaerasuchus Dibothrosuchus Alligator Ns 1975, (Nash Pl2005). (Pol The . 323 . Downloaded By: [Columbia University] At: 05:34 17 September 2007 Poorpsb .Leonard.) C. by (Photographs iue7.FN R2312, PR FMNH 77. Figure of ulnare 8736, UA 76. Figure h eaapl,a in as metacarpals, the aea D iw Scale view. (D) lateral notntl,ltl a esi eadn h manus the regarding said of be can little Unfortunately, Manus stormbergi 324 .tsangatsangana A. .H Turner H. A. Araripesuchus Ns 1975). (Nash rrpscu tsangatsangana Araripesuchus ¼ ned hsetnst h taxon the to extends this Indeed, . m Poorpsb .Leonard.) C. by (Photographs cm. 1 Dibothrosuchus r osdrbylne than longer considerably are rrpscu tsangatsangana Araripesuchus or etunr natro A,mda B,pseir() n aea D,ve.Scale view. (D), lateral and (C), posterior (B), medial (A), anterior in ulnare Left . Orthosuchus ih eaaplI n hln natro A,mda B,pseir() and (C), posterior (B), medial (A), anterior in phalanx and II metacarpal Right . hlneleeet aebe ecie o h other the for Araripesuchus described been have elements phalangeal Araripesuchus ie httemjrt fti aeilwsisolated, or 77). difficult was was material Figure manus the this to 2328; assignment of unambiguous majority PR the FMNH that Given 2312, PR material (FMNH specimen–bearing the among from recovered ubro eaoil n hlne were phalanges and metapodials of number A species. ngnrl eiga omtcra or metacarpal no as seeing general, in ¼ cm. 1 Downloaded By: [Columbia University] At: 05:34 17 September 2007 soitdwt ii I Scale II. digit with associated n tue,edn nasaladsihl recurved slightly and small a phalanx. in ungual shorter ending become stouter, phalanges and dorsal the in distally, proximal appearance Moving clover view. a The end proximal concavities, element. the conjoined giving three manual of The comprised is preceding concavity constriction. reception the the midpoint for of concave distinct is surface the articular a proximal to have slightly appearance and twisted laterally slight rotated 78). (Figure a the trochlea metacarpal and in the trochleated resulting is of small end another plane distal is triangular shallow there The and a which depression. is to broad end medial The anterior relatively proximal depression, the the III. is on of Present III surface slightly. MC very with MC medially of the overlap of flares its end cross-section, portion at proximal thins in proximal-most which triangular the of portion roughly medial trochlea. is slight a end in terminates MCIIislessslenderthanMCIV.Theproximal portion of distal phalanges The three are the and 2328 II. II–IV, MC PR metacarpals FMNH consists specimen of this impossible. metacarpals, identified only cases the most in 2328, PR FMNH 78. Figure h rxmlpaagsaelne hntedistal the than longer are phalanges proximal The end. proximal its lacks but slender, is IV MC ¼ m Poorp yC Leonard.) C. by (Photograph cm. 1 rrpscu tsangatsangana Araripesuchus ata au rsre eaapl I I,adI,a ela phalanges as well as IV, and III, II, metacarpals preserved manus Partial . MHP 30i iatcltd u preserves complete nearly but a another. and pubis. ischium, disarticulated, one left left partial is a with ilia, 2330 both associated PR not nearly elements, FMNH two a were 8767), These and 8768). (UA 2301, however, (UA PR ilium sacrum FMNH left of partial complete that a than was smaller ilia. right recovered and left Also well-preserved is sized, consists rib. 2303 moderately PR ilium FMNH of sacral pubis. exposed partial the poorly a and right but ilium large left and a posterior preserves 2297 ribs PR the FMNH sacral 2301) from and PR anterior exposed disarticulated (FMNH sacrum individual the partially larger The lacks a is a hindlimb. of though pubis ilium right right 2298, right the PR the underneath FMNH of in pubis portion preserved left and by not ischia obscured The is are exposed. sacrum poorly the and of matrix left surface and dorsal right The well- complete ilia. a to with attached pelvis sacrum complete preserved most the preserves 2298 ayn ttso rsrain eercvrdadare and to recovered were referred in preservation, 2331) of 2330— states PR PR varying FMNH 2303, FMNH PR FMNH 2297—2298, 2301, PR UA FMNH 8767—8768, (UA 8776—8778, pelves seven least at of Portions preservation and form girdle—general Pelvic New rrpscu tsangatsangana Araripesuchus Araripesuchus rmMadagascar from MHPR FMNH . 325 Downloaded By: [Columbia University] At: 05:34 17 September 2007 Poorpsb .Leonard.) C. by (Photographs iue7.FN R2303, PR FMNH 79. Figure 326 h ui i o ietycnatteiim(e.g. ilium the contact directly not that gomesii indicates A. did pubis the pubis for a ilium the the of on absence facet The contact in 1992). seen (Brochu be can individuals fusion mature of among degree a condition not though pervasive had crocodylians, ilium—the ischium the the with that fused indicates facet ischia. recovered smooth of The the paucity a of and facet preservation ischiac anterior poor to difficult generally remains given facet anterior characterize the but ilium, the on h otro loshai otc ufc ssmooth is surface contact ilioischiadic posterior The .H Turner H. A. , .patagonicus A. rrpscu tsangatsangana Araripesuchus n te crocodyliforms). other and ih lu nvnrltrl() eil() n osl()ve.Scale view. (C) dorsal and (B), medial (A), ventrolateral in ilium Right . ta.(00 in (2000) al. et uln n eyde.Hct(91 oe h same the noted (1991) in Hecht condition deep. very and outline opooyi enin seen is morphology h ctblmi vrugb ag supra- large a by that suggesting overhung crest, is acetabular crocodyliforms. acetabulum non-neosuchian The other and 1996b), esdo a tlatcpbeo eaieyupright the relatively of a morphology the of with capable consistent is least This at posture. was or sessed es obs.), pers. h ctblmi lotcmltl iclrin circular completely almost is acetabulum The hmeauhsparadoxus Chimaerasuchus .gomesii A. .patagonicus A. oouhsterrestris Notosuchus AN 45)adOrtega and 24450) (AMNH ieie hssame this Likewise, . Araripesuchus W n Sues and (Wu Pl2005; (Pol ¼ pos- cm. 1 Downloaded By: [Columbia University] At: 05:34 17 September 2007 us1996b), Sues oyin,e.g. codylians, pectblr rcs sgetyrdcda in as reduced greatly gomesii is anterior A. the well-developed process a but (preacetabular) bears process, (postacetabular) 79–81) posterior (Figures ilium The Ilium of characteristics posture. limb showed erect also which girdle, shoulder 2303, PR FMNH 80. Figure Poorpsb .Leonard.) C. by (Photographs , .patagonicus A. hmeauhsparadoxus Chimaerasuchus oouhsterrestris Notosuchus rrpscu tsangatsangana Araripesuchus n te mesoeucro- other and , Pl2005), (Pol W and (Wu etiimi etoaea A,mda B,addra C iw Scale view. (C) dorsal and (B), medial (A), ventrolateral in ilium Left . otro aigeg ftepseirpoeso the blade iliac of in dorsal process developed seen the posterior lacks ilium the the The of onto ilium. continues edge and facing crocodylians, posterior in as heavily as mor- thalattosuchians. in this present not is although phology a crocodyliforms, trait protosuchid this considered pusillus Theriosuchus 1999), insignis Mahajangasuchus h oslsraei titdaogiseg,btnot but edge, its along striated is surface dorsal The rgascu aznarezi Uruguaysuchus Alligator New Araripesuchus en oesmlrt htse in seen that to similar more being , W ta.19) lr (1994) Clark 1996). al. et (Wu yaoopyfrnon- for synapomorphy rmMadagascar from BclyadBrochu and (Buckley Rsoi13)and 1933) (Rusconi ¼ cm. 1 327 Downloaded By: [Columbia University] At: 05:34 17 September 2007 ean eryitc euceta scmaal to comparable is that gomesii peduncle 8768 A. intact ilia—UA nearly most a on anterior retains preserved The both. poorly on is smooth peduncle are one, facets anterior contact the than 2303. with larger PR is FMNH peduncle and teres posterior 8768 The the UA on for visible scar is A ligament peduncles. posterior and anterior 81). immedi- (Figure process, acetabulum the posterior of the outside on ately of supraacetabular ilium 2301 right prominent PR the on FMNH found the is foramen below nutrient A crest. ilium surface the entire the of comprises which acetabulum, deep te auenncongopcooyiom and right partial crocodyliforms of view group Lateral immature B, non-crown 8768. UA mature ilium left other view Ventrolateral A, scarring. muscle and Scale size 2301. ilium PR between FMNH Comparison ilium 81. Figure 328 h ctblmi efrtdvnrlybtenthe between ventrally perforated is acetabulum The .H Turner H. A. . Alligator Boh 92.Teiimbasa bears ilium The 1992). (Brochu ¼ m Poorpsb .Leonard.) C. by (Photographs cm. 1 and rmnn,frigatinua pog nthe Both on ilium. the “prong” of triangular surface a anterodorsal forming prominent, euce oslt h ctblrproain(e.g. anterior perforation the acetabular to the Alligator to posterior 1921), dorsal lies (Mook peduncle, process process anterior the reduced however, crocodylians a Living in have peduncle. nearly also anterior but the to, dorsal with, located line is process The 1999). ta.1996), al. et paradoxus n MHP 33 h tutr sgetyreduced in greatly is as structure The 2303. PR FMNH and lhuhsal h neirpoesi very is process anterior the small, Although h neirpoesi elpeevdi A8768 UA in preserved well is process anterior The aaagscu insignis Mahajangasuchus oouhsterrestris Notosuchus MH174)(iue80). (Figure 197946) FMNH ; W n us1996b), Sues and (Wu rgascu aznarezi Uruguaysuchus Pl2005), (Pol BclyadBrochu and (Buckley Theriosuchus Rsoi1933) (Rusconi Chimaerasuchus .gomesii A. (Wu Downloaded By: [Columbia University] At: 05:34 17 September 2007 rcs a ietdpseirywtotany (e.g. without deflection posteriorly ventral and or ilium. directed this the dorsal crocodylomorphs was of in length that process total appears the it Ancestrally, one-third iliocostalis than M. more of origin is 81). the (Figure process mark 1923) the (Romer may of and surface rugose The process. “prong”-like oouhsterrestris Notosuchus paradoxus Chimaerasuchus Ns 95 and 1975) (Nash i o figure not did like shaped not paradoxus Chimaerasuchus W ta.19;IP 163.Teanterior The V10613). IVPP 1996; of al. processes et (Wu AN 45)and 24450) (AMNH aeti ae“rn”lk neirpoes sdoes as process, insignis anterior Mahajangasuchus “prong”-like same this have MCP 6) hrfr,Iitrrtteposterior in the ilium interpret the I of therefore, process 267), PV (MUC h oslsraenal bet iia odto is condition whole, similar the A seen a absent. nearly of surface curve as the dorsal and the pronounced at less is, process projection the of ventral process end the the the with broader 81B), dorsoventrally PR Figure (FMNH specimen 2301; larger the In the process. of margin posterior dorsal the on ventral curve slight small process a with the combined a of to portion posterior-most however, the due, on projection be may This aspect. posteroventral slight a have to appears process the 8768) uraeaua rs htcrepnst h rgnof individuals. origin the the larger of to corresponds margin specimens, in that posterior crest the supraacetabular than smaller on present distinct is In depression less A margin. is dorsal scarring the along centimeter. a are of and half another a one about contact by separated not The do distinguished. facets be contact still two can outline semicircular ilium.its preserved, and the well less two is of surface facet the surface anterior the Although the of of longer two-thirds the nearly is occupies facet posterior The and ss2). (ss1 2303 are PR FMNH ribs of ilium sacral right the two on preserved the for facets the of The end process. the posterior on striations light few a only with smooth, the approximately its length. for its along of dorsolaterally half twists final face process to The length ventrolaterally. crest, supraacetabular faces the near originates in it as as process, of well ilia the as in present is morphology 93 h rcs sdrce otrvnrly nthe In posteroventrally. of directed ilia smaller is process the 1933) otro rcs fteiimin ilium the of process posterior deflection. dorsoventral any without posteriorly h otro rcs seogt,cmrsn slightly comprising elongate, is process posterior The nltrlve,msl tahetsarn svisible is scarring attachment muscle view, lateral In predominantly is ilium the of surface medial The o 20)ntdadroeiltiti the in twist dorsomedial a noted (2005) Pol Terrestrisuchus .gomesii A. .tsangatsangana A. rgascu aznarezi Uruguaysuchus oouhsterrestris Notosuchus .tsangatsangana A. .gomesii A. .I oebslcooyiom (e.g. crocodyliforms basal some In ). AN 45)and 24450) (AMNH rgascu aznarezi Uruguaysuchus Pl2005), (Pol .patagonicus A. Araripesuchus U 64 and 8654) (UA W n us19b are 1996b) Sues and (Wu and W n us1996b), Sues and (Wu FN R20,UA 2303, PR (FMNH rhscu stormbergi Orthosuchus Notosuchus .patagonicus A. n ucn (1933) Rusconi and , .tsangatsangana A. Pl20)and 2005) (Pol MCP 267) PV (MUC spsesn a possessing as Dibothrosuchus ob directed be to .patagonicus A. Theriosuchus similar A . (Rusconi The . ietecniinse in seen condition the like neiryo h eta ufc fteposterior the of in than surface ilium the ventral of the process more positioned on is anteriorly scar caudofemoralis the V10613) eetalre tahetsraefrtecaudofe- the for In surface simply attachment may moralis. and larger ilium a the process, on reflect posterior higher the located of is this end but the on notch a possesses fsc crin presence scar the a but such scar, of similar a rim. prominent bears enlarged 24450) very an (AMNH is by ventrally scar bounded the and 2301), PR (FMNH ilium oc sfiue yRsoi(93 g 6 for 36) fig. (1933: Rusconi by similar aznarezi A figured ilium. Uruguaysuchus the is of remainder notch the and it between in 2301). PR larger (FMNH proportionally specimen is posteroventrally large flange is projects the The that ilium. surface the flange and from attachment a Carrano as This 1923, developed of 2002). (Romer surface process Hutchinson ventral the posterior on located the is brevis process. femoralis) the of end posterior the at scar tibialis a condition The leaving attach process. process, the in FTE on posterior end and the unscarred smooth FTI on the forward further 1992), Brochu 197946; FI)adetru FE.In (FTE). and externus attach- internus Carrano and tibialis the flexor the as (FTI2) as of two head area well for surface this ment as figured (2002), (1923), Hutchinson cortical Romer the of preservation bone. poor to due determined be U ietdvnrlya 45 a the at of of end bottom ventrally the the directed with on “U”-shaped, “U” situated is is It two process. the the of dorsal more The muscles. 3 the iliotibialis common in and is iliofibularis present scar both is the PR to that division indicate (FMNH a may and that specimens depression, 8768) some UA M. in and (Brochu 2301 appears of crocodylians older It origin in seen 1992). Hutchinson be the can and that for Carrano 2002) corresponds 1923, area (Romer however, the iliofibularis region, to This topographically M. 3. to related scarring iliotibialis the be may that concavity ovular taxon. present the scar in 1 circular discerned iliotibialis M. be to distinct cannot more A respectively, head. second the smaller, posterior, in of that and have muscle anterior heads this attachments the of 3) for 2) iliotibialis M. surface iliotibialis of attachment (M. Alligator large head origin a Hutchinson second the and depict to Carrano (2002) 1923). corresponds (Romer and iliotibialis of scars largest along the is the crest rugosity supraacetabular the The of margin 1923). the (Romer iliofemoralis M. hr tpoet rmteiim oc spresent is notch a ilium, the from projects it Where (coccygeo- caudofemoralis M. of origin The h otro rcs er w ucesa sites. scar muscle two bears process posterior The an is there crest supraacetabular the to Posterior Araripesuchus h rt(.iitbai )adtid(M. third and 1) iliotibialis (M. first The . New Theriosuchus Araripesuchus .patagonicus A. ifr rmti,wt h flexor the with this, from differs . aaagscu insignis Mahajangasuchus W ta.19;IVPP 1996; al. et (Wu rmMadagascar from Araripesuchus Alligator MCP 6)cannot 267) PV (MUC 8 nl.I h larger the In angle. Alligator .Tog the Though ). ie more (i.e. .gomesii A. (FMNH 329 Downloaded By: [Columbia University] At: 05:34 17 September 2007 Poorpsb .Leonard.) C. by (Photographs shu,ltrlve.B eticim eilve.Scale view. medial ischium, Left B, view. lateral ischium, 330 rsre Fgr 2.I slreyitc,though intact, largely is It been 82). have (Figure to appears preserved ischium one only Interestingly, Ischium ilium. the of remainder the and is it notch between no present ventrally, developed moderately is surface 2330, PR FMNH 82. Figure .H Turner H. A. rrpscu tsangatsangana Araripesuchus ,Left A, . ¼ cm. 1 hn ihteiicpoessls asv hnin than massive less processes iliac like neosuchians the with thin, proximo- running 1991). fracture (Hecht it a from through distally ischium has known 24450) only (AMNH the and them h shu,acniinsmlrt htof that insignis to similar suchus condition a ischium, the of blade the from symmetrically diverge not do processes Alligator nelre peia ed In head. spherical with enlarged process cylindrical an this lack 18) Figure 1933; coni ecie ie that given described edlk in spherical like enlarged an in head ending cylindrical, and long is rcse htcnatteiimadadistally a greater a and from to proximal Ischia medially extent. ilium lesser of curves or the which pair blade, contact a expanded possesses that it processes as crocodyliforms hsmsl speetee nhatchling for in scar even a present because is interesting muscle is this which 2330, FMNH in discernable PR flexor is M. scar of such three No head internus. of tibialis bears origin the process to iliac related posterior scarring the to ventral ischium the marks muscle. likely this and of present bone, origin is the the striations on of (Romer present set femoris not a is although the adductor rugosity M. as distinct of A serves The two 1923). blade head puboischiotibialis. ischial for M. the origin of of pit side origin shallow ischium, posterolateral a the the is to there of process, related iliac side anterior anterolateral the limb below of the number a On of muscles. origin the for surface attachment posterior a the on whether present process. unclear is is surface It acetabular evaluate. distinct to difficult element tsangatsangana A. was pubis acetabulum. the the from that surface excluded indicating ventral completely process, the anterior on located the is of pubis the for contact rxmledi agl natwt nyteproximal damaged. the processes only iliac the with of intact ends largely broken be The is is end. end not distal-most blade proximal its could missing The and blade midshaft matrix. distal around of the cleaned of completely portion a laterally yidia nsae rjcigatrol ihlittle ilium. the with contacts process anteriorly the as projecting inclination dorsal generally shape, is shaft in remaining the cylindrical the of but end broken, proximal the is The process ilium. from the diverges of process margin anterior iliac anterior The 1996b). in condition n us19b and 1996b) Sues and rcu19) Both 1999). Brochu vrl,teicimi lne n mediolaterally and slender is ischium the Overall, h shu of ischium The nms rcdlmrh,tepseirmri of margin posterior the crocodylomorphs, most In the as serves ischium the of surface lateral The in unpreserved largely is process posterior The In .gomesii A. Mo 91 es b.FN 996.The 197946). FMNH obs. pers. 1921; (Mook hmeauhsparadoxus Chimaerasuchus aaagscu insignis Mahajangasuchus AN 45) h neirprocess anterior the 24450), (AMNH and Theriosuchus edrn h opooyo the of morphology the rendering , .tsangatsangana A. .patagonicus A. Alligator hmeauhsparadoxus Chimaerasuchus rgascu aznarezi Uruguaysuchus Araripesuchus u otatn the contrasting but , W ta.19)and 1996) al. et (Wu .tsangatsangana A. osntpreserve not does stpclo most of typical is W n Sues and (Wu Bclyand (Buckley Mahajanga- r o well not are .gomesii A. Alligator (Rus- the , (Wu Downloaded By: [Columbia University] At: 05:34 17 September 2007 ui hf,hwvr sntrttdmdoaeal at The mediolaterally ischium. rotated not the is however, pubis contact shaft, the of pubic to shaft posteriorly the with extending dorsally faces blade pubic aznarezi BclyadBoh 1999), Brochu and (Buckley ui ssihl atndpoial n rotated In and blade. the proximally of plane flattened the from slightly medially is pubis 1996). ale ihncooyiom,e.g. crocodyliforms, paradoxus in It within occurred 1990). apomorphy (Clark this earlier Clark that and however, Eusuchia Norell appears, the now 1988, for Clark from con- and synapomorphy previously Benton excluded been a therefore has sidered feature is blade This and triangular acetabulum. flat, ischium with articulates a the pubis the into cylindrical earlier, described nearly As expands distally. slender outline which a of obturator in neck, an consists lacks paddle-shaped and bone foramen the generally Proximally, 83). is (Figure pubis The Pubis of 1923, origin the (Romer iliac with 1986). internus Rowe posterior associated puboischiofemoralis the likely of M. is margin are that dorsal striations process small PR the of set FMNH on A of present exposed. well surface less is medial 2330 The 1992). (Brochu 2331, PR FMNH 83. Figure ui,vnrltrlve.B eticim osmda view. dorsomedial ischium, Left B, view. Scale ventrolateral pubis, h ui sitrsigbcuetesato the of shaft the because interesting is pubis The ¼ m Poorpsb .Leonard.) C. by (Photographs cm. 1 Rsoi1933), (Rusconi W n us1996b), Sues and (Wu rrpscu tsangatsangana Araripesuchus aaagscu insignis Mahajangasuchus Sunosuchus Chimaerasuchus Uruguaysuchus Alligator W tal. et (Wu ,Left A, . the , l in all rsn in present ogl vli hp.Tetopt rsn nthe on present pits two in The surface and shape. articular smooth in is oval surface roughly damaged the being preserved, latter the Where of slightly. FMNH surface the and with 2330 2331, PR PR FMNH except specimens all 83). in (Figure the blade of the length into the softens of it one-third before distinct first pubis a the as about blade for the towards ridge out extends shaft the of nte truhya45 a one roughly oppose shaft at the another of portion flattened the and pubis ml oto nisdsa riua surface. a articular lacking distal femur its the on only portion with elements small preserved hindlimb well these frac- very places, and of Although are number largest a complement. in the right tured hindlimb preserves distal intact preserved 2300 most poorly PR a FMNH of femur. FMNH consists and proximal respectively 2335 femur, the right PR and preserved left PR 2337 a and FMNH of PR present. proximally half FMNH not broken is and is fibula 2297 left tibia the and left distally The obscured matrix. remains damaged femur by left is the femur of much right and The midshaft hindlimbs. right and left recovered. also 8781, were (UA 2335) 2330, PR PR fibula FMNH FMNH left 2300, PR two FMNH 2298, and PR FMNH right three PR therefore, PR and FMNH FMNH 2335, 2337) are, 2298, PR left FMNH PR 2330, and PR (FMNH Three FMNH 2300, tibiae individuals. paired right four four were than and more these of of indicative although all 2337), PR FMNH not 2330, (UA PR PR FMNH recovered FMNH 2298, 2300, were PR FMNH femora 2297, PR right FMNH 8734, four preserved. and are left fibulae Four and tibiae, femora, Numerous preservation and form Hindlimbs—general the pubis. on the of located ridge is the ambiens on surface M. facing of ventrolaterally one its attachment head the and blade, for for shaft pubic surface pubic the made the to relation of be in flattening rotation slight cannot the to possibilities Due of these tsangatsangana point of A. exclusion or one the between described of Distinction as ligament (1992). well Brochu pubis-ischium by a as Carrano for 1923, 2002), attachment (Romer ambiens Hutchinson M. attachment and of of corres- two site head rugosity the for shallow with problematic. topographically a more ponding the is proves along there pubis the Anteromedially, scarring of of muscles end heads articular the two among the Distinction PIFE2). for and (PIFE1 blade externus surface puboischiofemoralis the M. attachment of as much 2002), served Hutchinson and Carrano h shai otc ufc sboe n missing and broken is surface contact ischiadic The MHP 28peevsprilbtarticulated but partial preserves 2298 PR FMNH 1992, Brochu 1923, (Romer crocodylians in As Alligator New Araripesuchus Araripesuchus .In .tsangatsangana A. ae ntepeevdmaterial. preserved the on based Alligator 8 . nl.Tefltee portion flattened The angle. rmMadagascar from Boh 92 r not are 1992) (Brochu h ld fthe of blade the , 331 Downloaded By: [Columbia University] At: 05:34 17 September 2007 Scale iue8.FN R2300, PR FMNH 84. Figure 332 hp,tog o osrnl imi as sigmoid strongly so not in though sigmoid generally shape, Like 84). is (Figure femur bone the gracile crocodyliforms, and most long a is femur The Femur 2337 PR 8780 pes. complete FMNH UA nearly metatarsal. a and second preserves IV, 2338 the metatarsal PR of consists isolated FMNH of an metatarsals, PR half preserves FMNH two recovered. distal preserves was the is IV 2330 and of III metatarsus consisting II, metatarsals The 2319) metatarsus A PR preserved. specimens. PR right of (FMNH FMNH number well a a in by is of represented present and is surface 2297 calcaneum articular A distal astragalus. the is preserves fibula end. left distal the its but but all preserved, preserving is complete, tibia more is left the femur proximal of the Only end right head. the only The lacking intact, femur more left is femora. The trochanter. fourth right the to distal and obscured left partial, asleeet r oerr.FN R2330 PR FMNH rare. more are elements Tarsal but preserved, well possesses 2330 PR FMNH ¼ .H Turner H. A. m Poorpsb .Leonard.) C. by (Photographs cm. 1 rrpscu tsangatsangana Araripesuchus Alligator etfmri aea A,pseir() eil() n neir()view. (D) anterior and (C), medial (B), posterior (A), lateral in femur Left . and n etatrmdal tnal 25 a nearly at anteromedially bent and scars. internus puboischiofemoralis tsangatsangana A. robusta naris ahcelscu trinquei Pachycheilosuchus FN 197946), the (FMNH trochanter, like neosuchians in fourth than straighter much and is constrictionshaft head the to femoral distal the that between fact the from stems This n aa rcdlmrh like crocodylomorphs basal and te escin ie oesmlrt crocodyliforms a to such similar more (i.e. neosuchians other odlsbigofe rmoeaohr hsofe is offset This 65 distal (roughly another. crocodylians and in one large proximal from the offset of being planes condyles long condylar the its in along lateral twisted resulting is the axis femur the of of shaft remainder The surface. fibular the the from delimits groove condyle shallow is A condylar condyle condyle. distinct lateral fibular subtle very a The possesses two and medial the 2298). than larger forms slightly PR and (FMNH expanded surfaces is end .tsangatsangana A. owtsadn,tefmrlha ssilexpanded still is head femoral the Notwithstanding, oouhsterrestris Notosuchus W ta.20) oevr h eoaof femora the Moreover, 2001). al. et (Wu aktecntito ttelvlo the of level the at constriction the lack hsofe sol rud45 around only is offset this Sunosuchus Rgr 03,and 2003), (Rogers and rgascu aznarezi Uruguaysuchus 8 ersrscu gracilis Terrestrisuchus ars 96,btin but 1986), Parrish ; W ta.1996), al. et (Wu 8 nl.Tedistal The angle. 8 ermino- o n i m r Te . Alligator ). Downloaded By: [Columbia University] At: 05:34 17 September 2007 nepee hssrcuea evn oeyfrthe for coccygeo- solely (CFL; longus femoralis serving caudofemoralis as of insertion structure this caudo- interpreted M. for point anterior 1923). insertion the (Romer longus the of femoralis portion as a serves least flange, at and trochanter, the aznarezi chus insignis oouhsterrestris Notosuchus patagonicus A. h uceisrin noteflnemypoemore prove may flange the onto insertions muscle The eu.Tesraeo h rcatri uoe(tr rugose is trochanter the the of femur shaft head, of the femoral surface down the way the The the to femur. of of close quarter one surface located than is less medial It 85). the (Figure on prominent Leonard.) (t C. by 1, trochanter (Photograph PIFI 1. fourth brevis; internus caudofemoralis puboischiofemoralis with CFB, 2300, longus; associated caudofemoralis CFL, PR scar attachment FMNH Muscle 85. Figure srgs.Asmlrsrcuei rsn nteother the in present is structure flange similar Araripesuchus the of A edge rugose. the is and striated finely the of is surface depression The the 94). of Figure shaft 103-1; the (character flange from femur (cdfemf). prominent anteriorly extends very outline that a process in on or located circular is depression nearly The and broad very 2002). Hutchinson of and weakly insertion 1992, Carrano Brochu is the 1923, (Romer to brevis trochanter caudofemoralis corresponds M. the and to shaped crescent posterior rugosity The tr 2 o 20)adBclyadBoh 19)have (1999) Brochu and Buckley and (2005) Pol h ersinatrmda otetohne is trochanter the to anteromedial depression The h orhtohne (t trochanter fourth The ,adi sbree nete ieb depressions. by side either on bordered is it and ), BclyadBoh 99,and 1999), Brochu and (Buckley sensu Rsoi13) h uoesraeof surface rugose The 1933). (Rusconi pce ( species MCP 6)a ela in as well as 267) PV —MUC h rvosymnindauthors). mentioned previously the Pl2005), (Pol .gomesii A. 4 rrpscu tsangatsangana Araripesuchus swl eeoe and developed well is ) AN 24450; —AMNH Mahajangasuchus Uruguaysu- 4 1 ). , . rcus(92 nepeaini orc hni is If it in than scar. flange correct the the that is from likely interpretation separate (1992) completely Brochu’s and insertion CFL the to distal of being 1 PIFI with 1992), Brochu 19)fiue h neto fPF ifrnl in differently 1 PIFI mississippiensis Brochu of CFL. insertion A. the to figured anteriorly (1992) surface slightly a shaped forms insertion crescent 1 inserting PIFI The M. on shaft. 1) longus femoral the caudofemoralis figure M. (PIFI to distally (2002) and 1 anteriorly Hutchinson internus and puboischiofemoralis (1879), Carrano Fauvel (1923), and Romer 85). (Figure complex hsinsignis chus odrdltrlyb lgtyrgs ig (t ridge rugose slightly depression a striated by a laterally is bordered femur, the of surface posterior tuberosity”. dorsal (1992) “proximal Brochu the that termed tuberosity. structure lateral rugose the and to the raised corresponds of a This is center femur, the the near of surface positioned apart and farther scars, spaced 2 are (rf1), in distally than proximally one 2 one scars, and internus two (rf2) The puboischiofemoralis 1923). Romer M. 2; indicating (PIFI of scars muscle of insertion set a the is femur, the of 1 surface PIFI increased on capability. mark adduction may overall and flange size the of in size and the resulted contraction CFL likely modern during The rotation flange medial in 2000). increased the of whole-limb Gatesy phase displacement and anterior ancillary swing (Hutchinson some the crocodylians create 2) 1 during to PIFI PIFI adduction appears of with it function in although (along The insertion ambiguous, their above. in remains discussed shift anterior taxa an CFL the the saw both 1 case, that PIFI the indicate and would is structure this this If then are interpretation. margin The this its of with (2002). rugosity consistent the Hutchinson and flange the and of Romer location Carrano of conclusions and for the given surface (1923) 1 attachment PIFI and common CFL a both as flange this anterior for insertion the CFL of retractor. femoral trochanter movement primary the would, fourth anterior the an of This to and CFL. enlargement relative surface for an attachment attachment indicate the therefore, as simply aznarezi n ssc ucisnadGts 20)reference (2000) Gatesy and clades Hutchinson archosaurian such other as in and trochanter not greater is corresponds the surface structure to declined this the Topographically, (1992) because tubercle. “trochanter” a Brochu of and use “trochanter”, the (1956) a as Romer it to while and referred (1984) is Carrano Crush structure mixed. this 1986, for Nomenclature 1923, puboischiofemoralis 2002). Hutchinson Romer M. of (PIFE; insertion externus the marks area meitl itlt h eoa ed ln the along head, femoral the to distal Immediately lateral the on flange, anterior the to Proximal the consider to however, unreasonable, not is It Alligator New a encretyitrrtdadserved and interpreted correctly been has , oouhsterrestris Notosuchus Araripesuchus Boh 92.Ascae ihPIFI with Associated 1992). (Brochu PF M (PIFI Araripesuchus rmMadagascar from sensu and , oe18 and 1986 Rowe , Uruguaysuchus Mahajangasu- G .This ). 333 Downloaded By: [Columbia University] At: 05:34 17 September 2007 334 xed ate ontefmrta in than femur the and down shaft farther femoral extends the of surface anterolateral the along h ii of tibia The Tibia (Brochu gastrocnemius 2002). condylar which Hutchinson M. and condyle, lateral of Carrano fibular 1992, origin the the the on to represents proximal present just a is surface attachment it and rugose tubercle muscle making appear small surfaces 2297 distinct condylar The PR discern scarring. FMNH to in difficult fractured heavily trochlea. medial well-developed and a lateral forms articular distinct and distal hemicondyles into developed The is appear- crocodylians. condyle early hatchling its the in noted as and ance ridge scar” the adductor to femorotibialis “primary referred M. M. (1992) of of Brochu origin internus. insertion the and the femoris between adductor as boundary structure the this marking the interpreted of (1923) expansion Romer the to condyles. prior trochanter distantly fourth diminishes the and near begins ridge The femur. the towards condyle. laterally the curving and distally prominent becoming more proximally, distinct not is ridge iliofemoralis rsn in present iliofemoralis M. of the insertions in the near denote beginning scars, 2 scars PIFI Two ridges. proximodistal in tsangatsangana shaft femoral Araripesuchus The ridges. running proximodistally ufc,tetbai akdb w ml,incons- of this small, insertion (interior two internus the to tibialis by flexor indicates 2298 distal M. marked One PR Just tuberosities. is FMNH picuous tibia surface. of the rugose well tibia surface, is its right but M. 2300, the illustrating PR iliotibialis, on FMNH The in M. preserved 1923). damaged of (Romer is ambiens surface tendon M. and common femorotibialis, the for crocodylians. in (1992) Brochu by proximalnoted pit” “proximal distinct The a the lacks 2300 surface. to PR FMNH medial proximally in end the extends than and extent lateral greater the than larger surface is articular medial The condyles. femoral the articulation with for surface is contact the end forms proximal it slight as The very expanded shaft. a its with in slender curve and anteroposterior long is bone The 2005). aa eourcdlassc as insignis such other mesoeucrocodylians as well basal as crocodylians modern of those to similar aznarezi hssraei rcdlas aeteial,as parenthetically, crocodylians, trochanter”. in “greater surface this h itli aae nFN R20 and 2300 PR FMNH in damaged is distal The of posterior the on runs ridge longitudinal second A rxmly h neireg evsa h insertion the as serves edge anterior the Proximally, Alligator Alligator .H Turner H. A. BclyadBoh 1999), Brochu and (Buckley Rsoi13)and 1933) (Rusconi rrpscu tsangatsangana Araripesuchus rrpscu tsangatsangana Araripesuchus femoraaremarkedbyanumberof Rmr12) nyoesc ig is ridge such one Only 1923). (Romer oouhsterrestris Notosuchus osse nytwo only possesses sensu Mahajangasuchus Uruguaysuchus ti located is It . Alligator Fgr 6 is 86) (Figure oe 1923, Romer The . (Pol rcdlas n oesmlrt h odto in condition the to in seen similar insignis that Mahajangasuchus more than and weaker much crocodylians, is of is and anterior prominent tibialis insertion not the internus with associated tibialis the scarring flexor proximal the the scar, with the indicates the As anterior. of This tibialis to M. lateral surface. distal more immediately articular The is 1992). tuberosities Brochu 197946; rxmlha evsa h taheto the of flaring attachment the the of lateral as surface in serves view lateral head when broad proximal fibula sharp The proximal a the aspect. producing to posteriorly, bend developed strongly of fibula The mesoeucrocodylians. other head. slender and crocodylians proximal and tsangatsangana long distinct very A. very fibula is a right element complete with The a 87). preserves (Figure 2300 PR FMNH Fibula ligament tibioastragalar the to medial 1992). (Brochu corresponds circular of which smooth lateral attachment surfaces, a and articular anterior medial bears the distal The tibia between a face. distal centered depression, the contact bears of astragalar tibia surface the lateral distal and of medial the edges the of between depression surface appearance. triangular curved posterior a tibia portion. the The of lateral the projects margin giving the laterally, lateral projects distal surface than fibula the extent facing astragalar edge greater The the a to of more distally the the margin portion Additionally, medial lateral. distally, medial the the than Viewed with developed strongly crescentic astragalus. is the surface with surface longus lateral midshaft. the digitorum near tibia pierces the flexor foramen of nutrient surface interosseus the small the A and to origins. correspond insertion and would smooth cruris along that present is shaft typically scarring shaft the muscle any The of fibula), flattened. devoid the (facing slightly edge lateral being the only with section, just anterior. tibialis sits the for ridge scar This insertion the individuals. to proximally the smaller of in presence even ridge the the slightly preserves illustrating 2298 completely, is PR structure FMNH depression but 2300, in The border PR anterior FMNH the ridge. of portion a small with along damaged, anterome- a bordered by depression dially shallow a bears end, rmtetba hf,a ti in is it raised as proximodistally sharply shaft, as is nor tibial scar distinct, the as The from not is shaft. the and from the elongate medial the of slightly on tibia, midline the positioned of is surface tuberosity anterior This 1992). Brochu h blrha smdoaeal opesdand compressed mediolaterally is head fibular The contact the form to expands tibia the Distally, cross- in circular nearly is tibia the of shaft The proximal the near tibia, the of surface lateral The ifr rmta fmodern of that from differs BclyadBoh 1999). Brochu and (Buckley Alligator (FMNH Downloaded By: [Columbia University] At: 05:34 17 September 2007 ogltrllgmn Boh 1992). (Brochu 197946), ligament (FMNH lateral long 2300, PR FMNH 86. Figure and insignis Mahajangasuchus iia to similar itl()ve.Scale view. (E) distal swa nteeohrtx n ismlrfo the from dissimilar and taxa of other these morphology however, in bend, the weak posterior to is bend The and head. expansion fibular posterior proximal slight a show all gomesii rgascu aznarezi Uruguaysuchus and .tsangatsangana A. .patagonicus A. .tsangatsangana A. ¼ m Poorpsb .Leonard.) C. by (Photographs cm. 1 Sunosuchus rrpscu tsangatsangana Araripesuchus BclyadBoh 1999), Brochu and (Buckley Rsoi13:fi.22) fig. 1933: (Rusconi hsfaueappears feature This . oss blrheads fibular possess W ta.1996), al. et (Wu Fgr 98). (Figure Alligator ih ii npseoeil() aea B,atrltrl() rxml() and (D), proximal (C), anterolateral (B), lateral (A), posteromedial in tibia Right . A. rcatri o yia o rcdlfrsi is iliofibular crocodyliforms—it the for of typical location not The is fibula. proximal the trochanter the contact of to edge trochanter the from extends bone the on the wrinkle inconspicuous near An margin. very proximal head, fibular the on proximodistally located smooth, ridge elongate a is trochanter The fibula. of the surface lateral the on feature distinct a is 1991) Sereno fibulae. any preserve nqeto unique h loblrstohne atro rcatrof trochanter (anterior trochanter iliofibularis The New Araripesuchus Araripesuchus —though rmMadagascar from .wegeneri A. osnot does 335 Downloaded By: [Columbia University] At: 05:34 17 September 2007 uretfrmni rsn eilyna midshaft. small near a medially though present and is scarring, foramen smooth muscle nutrient distinct is of shaft devoid the Generally, mediolaterally. flexor M. of 1992). origin (Brochu the longus flange, to digitorum the on corresponds the rugose of structure not margin the is anterior bone the the along Although flange fibula. thin a is there 1991]). [Sereno Mahajangasuchus n itl()ve.Scale view. (F) distal and ontesato h bl (e.g. fibula length the the of of shaft quarter the one down about located generally 2300, PR FMNH 87. Figure 336 h hf ftebn ssedradcompressed and slender is bone the of shaft The trochanter, iliofibularis the to anterior Immediately .H Turner H. A. n ueosohrarchosauriforms other numerous and ¼ m Poorpsb .Leonard.) C. by (Photographs cm. 1 rrpscu tsangatsangana Araripesuchus Alligator ih bl nltrl() otro B,mda C,atro D,poia (E), proximal (D), anterior (C), medial (B), posterior (A), lateral in fibula Right . and raglri rs-eto,btlcigadsa hook distal twisted a lacking a but ( cross-section, fibula in the triangular of shaft the appearance. the giving of portion proximal shaft the than crease surface anteriorly sharp This more bone. a faces the and of fibula surface posteromedial the the to on fibula posticus edge forming the anterior flattened, tibialis very sharp of is a tibia) M. two-thirds the facing site distal of portion (the This The origin head. 1992). the fibular (Brochu the the indicates on to likely wrinkling distal surface subtle medial reveals examination Close sensu h itledo h bl selre and enlarged is fibula the of end distal The rcu19)lk htse in seen that like 1992) Brochu Alligator . Downloaded By: [Columbia University] At: 05:34 17 September 2007 elhsohrcooyoops ossso the the on of “peg” a of consists morphology consists This 1978). terjee Chat- 1971, crocodylomorphs, (Bonaparte condition normal” other “crocodile has well n eta F iw ih srglsi eta iw() Scale (G). view ventral in astragalus Right view. (F) ventral and aslmrhlg of The morphology 1992). Brochu tarsal 1991, Sereno 1986, Parrish 1984, Tarsitano and Hecht 1982, Chatterjee 1980, Brinkman 1979, (Cruickshank signifi- tarsus archosauriform systematic the of and cance morphology the on been has conducted work of amount vast a carpus, the to contrast In the preservation and on form Tarsus—general tubercle 1992). associated a (Brochu cruri is interosseus is M. of tubercle there origin the This with fibula, surface. the posteromedial of end xso h blrsatb bu 5 about by shaft fibular the main of the axis from anterolaterally displaced end distal The 2297, PR FMNH 88. Figure rrpscu tsangatsangana Araripesuchus rrpscu tsangatsangana Araripesuchus 8 rxmlt the to Proximal . etclaemi neir() aea B,pseir() eil() osl(E), dorsal (D), medial (C), posterior (B), lateral (A), anterior in calcaneum Left . ¼ as m Poorpsb .Leonard.) C. by (Photographs cm. 1 e eaiet ttoayataau n tibia. and astragalus stationary a and to calcaneum. calcaneum relative the pes the of on motion rotational “socket” for allows a This into fits that astragalus h itlsraeo h srglsi eysmlrto of similar comprised is very It is 88G). astragalus (Figure the crocodyliforms other of surface distal The Astragalus taxon. the tarsal for recovered with were elements or tarsal no distal Moreover, fibula. articulated or tibia the speci- tarsus to articulated these elements the of Neither preserve 2298). mens PR PR (FMNH FMNH recovered and well- Two also 2297 elements. were the calcanea of preserved exposing surface and posterior matrix, the only rock still within 2330), PR contained FMNH partially 2297, PR (FMNH astragalus The rrpscu tsangatsangana Araripesuchus New Araripesuchus rmMadagascar from rsre two preserves 337 Downloaded By: [Columbia University] At: 05:34 17 September 2007 eta roeo ucste“epgov”o Hecht deep of a groove” “deep forming sulcus—the prominent, or the and groove of central sided edges high lateral are and tuber medial in The to 1933). present similar (Rusconi and condition calcaneum the the of morphology overall tuber. of calcaneal margin the medial trough the past the extending medially posteriorly, articulates. while bends trochlea anteriorly faces astragalar trough the The which upon facet foramen. and nutrient small smooth a is by pierced surface appears lateral The in 197946). seen is (FMNH what than thinner ball the of surround- bone the ing leaves This 1999). Brochu and in (Buckley square deep—as and roughly large is very socket the is calcaneum, socket the of size This the to Relative peg. shaped. reception astragalar the for the socket deep of a bears ball medial flattened the The of distal 1992). surface Brochu a the 1980, with (Brinkman forming IV contact tarsal for one-third surface remaining ventrally-directed the smooth and and rounded surface contact tibial the of thirds tuber. directed posteriorly the parts and peg, distinct ball, astragalar anterior three an 1992)—the into Brochu 1980, divided of (Brinkman be can calcaneum calcaneum the aznarezi resembling Uruguaysuchus element, narrower tsangatsangana Araripesuchus acnu Fgr 8.Lk h srgls this astragalus, crocodyliforms. to other the of that Compared to Like left similar very isolated is 88). element complete (Figure a preserves calcaneum 2298 PR FMNH Calcaneum pc hog hc h edno .flexor in seen second M. is A what of than 1992). deeper tendon (Brochu the slightly groove, passes supplies the longus groove which digitorum This through trochlea. space distal the the from divides groove roller astragalar deep con- the a is Posteriorly, and roller, trochlea. distal Protruding the from (1983). laterally of tinuing Tarsitano hollow” “anterior and for fossa—the Hecht squarish surface a element. revealing articular tarsal distal the third the is and II the I, roller metatarsals and The distal 1984) 1984). Tarsitano Tarsitano medial and (Hecht and trochlea Hecht astragalar 1979, (Cruick- shank roller distal medial surfaces—the distinct two hspgde o ae oad h aea margin— lateral the in towards condition the taper unlike not but does sides, peg sub-parallel has this calcaneum, the into inserts 1992). Brochu 1980, (Brinkman metatarsal-distal the ligaments for tarsal groove attachments the This of anteriorly. one more marks surfaces two the divides 338 h ue svr arwcnitn ihthe with narrow—consistent very is tuber The trough-shaped a is socket calcaneal the to Medial two- with narrow, mediolaterally is ball anterior The oto fteatro ufc sexposed is surface anterior the of portion A h srglrtohe,wihfrstepgthat peg the forms which trochlea, astragalar The .H Turner H. A. Alligator Alligator Rsoi13:fi.2) The 24). fig. 1933: (Rusconi acna h acnu of calcaneum the calcanea, rgascu aznarezi Uruguaysuchus aaagscu insignis Mahajangasuchus . samediolaterally a is Alligator Alligator , r etrkoni other in in represented known poorly better is are pes the tsangatsangana Araripesuchus manus, the Like Pes than theless, ere fcmltns U 76 A88,FMNH varying with 8780, UA in 8736, along (UA phalanges completeness preserved of and degrees are (MT) metatarsals 2337) isolated FMNH 2319, PR gomesii A. nla fteeeeti ssrnl compressed strongly as in is is it element as the mediolaterally if unclear iia ewe h w pce.Tetbrof tuber The of are species. that like tuber two and gomesii the ball Araripesuchus anterior between the similar of proportions to The similar very are and r nw from known are cus n astn (1983). Tarsitano and per httemriso h w tutrsare it structures two Ventrally, the another. trough. one of with the confluent margins nearly from the of tuber 1992). that surface the appears Brochu separates posterior 1983, bone the little Tarsitano very and Medially, to attach (Hecht not does it but this tuber. crocodylians through modern the passes in on groove tendon groove (Achilles’) deep calcaneal and The sided high similarly a eaaslI U 70.Atro ntelf n otro nthe on Left posterior B, and right. left the the on on Scale Anterior posterior right. 8780). and (UA left II the (FMNH metatarsal III on metatarsal Anterior Left A, 2338). morphology. PR Metatarsal 89. Figure oclaemi nw for known is calcaneum No rcuigacmaio ihta ao.Calcanea taxon. that with comparison a precluding , Alligator .gomesii A. ¼ AN 45) w ata e (FMNH pes partial Two 24450). (AMNH rrpscu tsangatsangana Araripesuchus m Poorpsb .Leonard.) C. by (Photographs cm. 1 FN 996 rcu1992). Brochu 197946; (FMNH rrpscu gomesii Araripesuchus ospsesaclaemnarrower calcaneum a possess does er eppseirgroove posterior deep a bears rgascu aznarezi Uruguaysuchus rrpscu tsangatsangana Araripesuchus .tsangatsangana A. nietemns pes manus, the Unlike . Araripesuchus rrpscu patagoni- Araripesuchus AN 24450) (AMNH oee,i is it however, , taxa—i.e. Never- . possess . Downloaded By: [Columbia University] At: 05:34 17 September 2007 .Leonard.) C. nasih wse pernet h metatarsal. body trochlea. the the into the gently of curving to trochlea, surface the posterior the appearance above with present is twisted depression a slight Anteriorly, a resulting slightly laterally in rotated the trochlea and the trochleated of are plane is ends distal surface The cross-section. proximal the of surface. articulating half the medial into MT thinned the in the while of only lamina, two-thirds thin III the nearly forms II, surface MT prece- proximal that In the lamina of metatarsal. distinct surface ding a proximoposterior proximal the into the main underlies noticeably medially the surface. as while thins thick metatarsal, surface fan-shaped as the nearly a of is shaft articular into surface proximal PR this expanded The FMNH Laterally, 89). and is (Figure II III surface MT MT large tibia. a the a 2338 of preserves length 8780 the UA half than more long—slightly are that elements assignable. clearly those and on complete elements solely specific based of therefore Details are given preservation. difficult of proves level elements assign- this these Unambiguous of many 2337). of PR ment FMNH 2330, PR Scale 2319) 2337). PR PR B, (FMNH right). (FMNH the pes (on pes right phalanges Partial and Partial left) the A, (on morphology. metatarsals the Pedal showing 90. Figure h hfso h eaasl r erycrua in circular nearly are metatarsals the of shafts The very and slender are metatarsals the Generally, ¼ m Poorpsby (Photographs cm. 1 Poorpsb .Leonard.) C. by (Photographs eta ufc ftebn.Gvntergeneral their Given bone. is the the but only of exposes surface other surface ventral the ventral while and places, in dorsal damaged for osteoderms both the of preserved preserves One 2333). PR are (FMNH recovered elements tsangatsangana armor Araripesuchus dorsal Few preservation and form armor—general Dorsal phalanx ungual 91). recurved (Figure slightly and small stout a more in and ending shorter become proximal Moving phalanges view. the proximal The distally, in appearance clover a element. concavities, it conjoined giving three pedal of comprised The is preceding concavity 90). reception the the midpoint Figure for of concave distinct 2311; is surface a articular PR proximal having (FMNH distal, constriction the than robust hlne ndra lf)advnrl(ih)ves Scale views. (right) ventral and (left) dorsal in phalanges iue9.FN R2314, PR FMNH 91. Figure iue9.FN R2333, PR FMNH 92. Figure iatcltdotoem,soigvnrladdra morphology. dorsal Scale and ventral showing osteoderms, disarticulated h rxmlpdlpaagsaelne n more and longer are phalanges pedal proximal The ¼ m Poorp yC Leonard.) C. by (Photograph cm. 1 New Araripesuchus rrpscu tsangatsangana Araripesuchus rrpscu tsangatsangana Araripesuchus rmMadagascar from w sedrswere osteoderms Two . ¼ Pedal . .Two cm. 1 339 Downloaded By: [Columbia University] At: 05:34 17 September 2007 340 Fgr 2.Tepeec falniuia ig as ridge longitudinal a of in presence The 92). (Figure gensis hs lmnsaercaglr n akthe crocodyliforms basal lack in and seen (e.g. rectangular, process are anterolateral elements These Osteoderms armor. the dermal the from of likely rows are anterior osteoderms preserved the outline, ouigo aa eourcdlasgnrlyand generally (2005), Calvo mesoeucrocodylians Araripesuchus and basal Turner on and focusing (2000) al. from et derived was Buckley regime across taxon-sampling scored The taxa. were al. 32 characters matrix et These the upon (1994). Buckley largely Clark based by turn from The in al. is primarily 2005). which et (2000), adapted Calvo Sereno and was 2003, Turner dataset Pol 2004, 2000, Brochu Turner 1996, and al. 2003, Buckley al. et 1997, et Buckley al. 1999, et Ortega from Wu 1994, gathered 1997, Gomani was (Clark characters, studies the new with previous two characters, of morphological addition 129 of dataset A sampling taxon and coding, character Dataset, subtle placement Phylogenetic by marked being two with the osteoderm, lateral the of striations. of the posterior surface mediolat- ventral toward run the the grooves so Two on elements. erally less the of become margin and the medially by overlap of surface osteoderms. the preceding ver- indicating osteoderm preserved the smooth, articulated of margin is not an anterior The with are column. tebral associated osteoderms and two Orientation articulated that the elements. to for because determined damage preserved be cannot the the patterns given in determined surface be cannot owal eeoe otrol 0 rvr strongly very or (0) posteriorly developed straight weakly head fibular to of portion 128—Proximal-most Calvo and new. Turner are 129 from are and are 128 Characters 124–125 126–127 (2005). 119– and Characters Characters 122 (2003). (1997). al. 120, et al. Sereno from et modified modified is Wu 121 from Character from (2000). al. modified et unmodified Buckley are from are 115–117 Characters (1997). 111–114 Gomani al. et Characters Ortega from modified (1996). are 123 Characters and (1999). 118 while Brochu 102– 107–110, and (2003), Buckley Characters Pol from unmodified. are to 106 are according 2–101 modified characters is 1 acter h eta agn fteotoem r concave are osteoderms the of margins ventral The h oe hrcesicue nteaayi are: analysis the in included characters novel The Char- (1994). Clark from are 1–101 Characters rrpscu gomesii Araripesuchus rtscu richardsoni Protosuchus .Tesraeo oei evl sculpted heavily is bone of surface The ). .H Turner H. A. npriua Apni B). (Appendix particular in and and ssscu chungkin- Hsisosuchus oouhsterrestris Notosuchus .tsangatsangana A. a neovdi eeoe l (2000). al. et however, Sereno taxa, in three unresolved was these between relationship The eas ofwaei omni hsadthe and (2000) this al. in examined et common only in Ortega included are analyses. taxa few aforementioned notosuchian so the because including topologies between Ortega in (2000). present al. is et topology converse The suchians. and derived te hnt other to than other patagonicus A. pce of species ihtetoSuhAeia taxa— American South two the with re eel httetretx epnil o the for responsible incomplete wegeneri relatively suchus taxa the three individual are nodes the the collapsed that of reveals these Examination trees of factors regions. provides contributing the and unresolved to consensus is as strict information 18 tree the no same to the the identical of for is consensus trees Adams support An nodal weak. generally overall is polytomies, the three-taxon tree three however, consensus only The with resolved 93). well (Figure the strict of analysis results A the slightly. present summarizes trees vary 18 the clades of larger consensus and the (Buckley although within 2003), Pol details 2000, analyses largely al. et Buckley ingroup previous 1999, Brochu the with among congruent a relationships was postulates of (CI) distribution index pattern character index retention The the consistency 0.6660. and (RI) 0.4396, The was 308. CI(inf) 0.4578, of length with a recovered were trees parsimonious most Eighteen Results poste- a the (1) with 97). junction possesses 96, posterior the (Figures or process at the tubercular spine (0) in projecting lacks rodorsally shaft ribs of cervical process and 98); (Figure 129—Posterior (1) flange distinct forming projecting lcmn ftefu aasae ncommon. with in of taxa reordering shared the complete four taxa nearly in a the considerably four is differ there the Essentially, (2000) of al. that and Topologi-placement et analysis (2001). this Ortega or of al. of clade (2000) et Ziphosuchia al. Sereno the et cally, of Ortega Notosuchia of similar clade the is Ziphosuchia this the sebecosuchians) composition, monophyletic to plus In a (notosuchians recovered. clade and including was taxa crocodyliforms Sebecosuchia of notosuchian clade including large traditional a clade node, this a with a Araripesuchus include Neosuchia, analysis the monophyletic in recovered clades large-scale ti ifiutt aeamaigu comparison meaningful a make to difficult is It h titcnessdmntae httefour the that demonstrates consensus strict The Notosuchus Araripesuchus Sebecus opiigissse ru.Otieof Outside group. sister its comprising bigmr lsl eae oeach to related closely more —being Notosuchus , Bernissartia , h otbslo h sebeco- the of basal most the Baurusuchus Malawisuchus Araripesuchus omamnpyei group, monophyletic a form hl eeoe l (2001) al. et Sereno while , and Iberosuchus en h most the being and pce.Other species. .gomesii A. Simosuchus . Araripe- and . Downloaded By: [Columbia University] At: 05:34 17 September 2007 igenmesidct ea ne o oe ihbosrpvle esta 50. than less values bootstrap with nodes for index decay indicate numbers Single iue9.Pyoeei eainhp within relationships Phylogenetic 93. Figure otprioiu re (length trees parsimonious most ¼ 0,CI 308, ¼ .58.Pie ubr tndsidct otta upr tp n ea ne (bottom). index decay and (top) support bootstrap indicate nodes at numbers Paired 0.4578). Araripesuchus n eourcdlapooe nti td.Te sasrc osnu f18 of consensus strict a is Tree study. this in proposed Mesoeucrocodylia and New Araripesuchus rmMadagascar from 341 Downloaded By: [Columbia University] At: 05:34 17 September 2007 o ifiut ti ot oig oee,ta the that however, noting, worth is Uruguaysuchus It compari- difficult. make the son sampling Again, taxonomic in crocodyliforms. discrepancies of analysis recent a phylogenetic in represented taxa notosuchian of proportion 342 rsn nlsswsntrcvrdi o n Norell’s, and Pol while in recovered not was analysis present hnto than oepoepsil hrce upr o alternate for within support relationships character ingroup possible explore to lent ooois dnia otefis reexcept tree of first placement Two the the most C. for to the Appendix identical of in topologies, one provided alternate for is apomorphies trees of parsimonious list complete A of position phylogenetic The considered. will be groups this these not of of discussion aim further the therefore not study, is relationships sebecosuchian and .Teol ldsdsusdhr r hs directly those Appendix are to here in discussed pertinent clades depicted only The are C. topologies alternative nae clade— Unnamed crocodyliforms. other to relative position taxon’s perne h cplrbae of the blades general twice scapular In is the the articulation. that appearance, on scapulocoracoid blade the scapular diagnosed of broad be length very can a of it basis but characters, derived Discussion (104-1). edge serrated postorbital anterior teeth straight of (96-0), with part osteoderms of bar dorsal dorsal postorbital (29-1), portion from of distinct postorbital part constricted, dorsal of (19-1), broad bar most a postorbital as contacting dorsally extends sheet narrow quadratojugal forming (9-2), maxilla, slit and premaxilla with contact support character Ambiguous in 96). shaft Figure (129-1; the ribs with cervical process posterior tubercular the of articulation junction at scapulocoracoid the spine projecting posterodorsally the 95), Figure of (106-1; length than the greater and twice broad very blade caudofemora scapular 94), Figure for bears femur of flange margin anterior (91-1), well-developed hypapophyses with vertebrae cervical (69-1), septum Synapomorphy Unambiguous Peirosauridae neirmri ssrnl ocv ncnrs othe to contrast margin. in posterior concave straight strongly is margin anterior insignis Mahajangasuchus o n oel(04 aeacnieal larger considerably a have (2004) Norell and Pol .H Turner H. A. Malawisuchus Uruguaysuchus hscaei upre yanme of number a by supported is clade This . 1 Araripesuchus Trematochampsa þ Araripesuchus Simosuchus rrpscu wegeneri Araripesuchus a on lsrto closer found was xlrto fnotosuchian of Exploration . r iia Fgr 5.The 95). (Figure similar are Araripesuchus i ucltr (103-1; musculature lis otu osrce at constricted Rostrum . haa iie ya by divided Choanae . ld eoee nthe in recovered clade eainhp n the and relationships 1 Araripesuchus Mahajangasuchus Araripesuchus eeused were , Notosuchus .Both and 1 iue9.Sseai aito nteatro agno femur. of margin anterior of femur the left in of view variation Medial Systematic A, 94. Figure hwn tt fcaatr13B eilve frgtfmrof femur right of view Medial 103.B, character Goniopholis of 1 state showing Scale richardsoni ih)and right) haai rsn ihngnohld swl as septate well A as clade. goniopholids the within terrestris to present Notosuchus unique is not choana these are Crocodyliformes, of this features context for the diagnostic Within also group. are hypapophyses developed oouha aa rvnstsigtefl xedof within extend distribution. full (especially characters the the testing remains prevents taxa) postcranial notosuchian a of but trait, in mesoeucrocodylian paucity widespread known more being a be as present more is well mwakayasyunguti trait suchus as also The Crocodylia clade. are this in just hypapophyses than widespread cervical developed ett haaadcria etba ihwell- with vertebrae cervical and choana septate A ¼ cm. 1 AN 04 ih)soigsae0o hrce 103. character of 0 state showing right) 3024; (AMNH rrpscu tsangatsangana Araripesuchus AN 2;lf)adlf eu of femur left and left) 620; (AMNH and hscaatrmy indeed, may, character This . aaagscu insignis Mahajangasuchus brscu macrodon Iberosuchus FN R20;left) 2300; PR (FMNH Protosuchus U 8654; (UA Malawi- Well- . Downloaded By: [Columbia University] At: 05:34 17 September 2007 n etcria i of rib cervical left and of iue9.Sseai aito ntebd fcria is ,Ltrlve frgtcria i of rib cervical right of view Lateral A, ribs. cervical of body the in variation Systematic 96. Figure of scapula left of view Medial A, shape. scapula in variation Systematic 95. Figure ucltr n ehp h puboischiofemoralis the caudofemoralis the perhaps of and insertion flange musculature the This 94). to (Figure femur corresponds the of margin anterior rrpscu tsangatsangana Araripesuchus mississippiensis liao mississippiensis Alligator ebr fti ld oss itntflneo the on flange distinct a possess clade this of Members AN 16 hwn tt fcaatr16 Scale 106. character of 0 state showing 1106) (AMNH rrpscu tsangatsangana Araripesuchus AN 16C)soigsae0o hrce 2.Scale 129. character of 0 state showing CA) 1106 (AMNH FN R21;rgt hwn tt fcaatr16 ,Mda iwo etsauaof scapula left of view Medial B, 106. character of 1 state showing right) 2313; PR (FMNH FN R29;lf)soigsae1o hrce 2.,Mda iwo ih evclrib cervical right of view Medial 129.B, character of 1 state showing left) 2298; PR (FMNH ¼ fti smr iepedaogntscin or notosuchians among widespread general. more in mesoeucrocodylians is unclear is this it material, if postcranial little to due again but nens1. internus cm. 1 New ¼ rgascu aznarezi Uruguaysuchus cm. 1 Araripesuchus aaagscu insignis Mahajangasuchus aaagscu insignis Mahajangasuchus rmMadagascar from a iia flange, similar a has U 64 et and left) 8654; (UA U 64 right) 8654; (UA Alligator 343 Downloaded By: [Columbia University] At: 05:34 17 September 2007 eilsraei ohtx.Scale taxa. both in surface medial aea agno ocv eilsraepeeti ohrb lwrarw) pe rosidct rsneo pn lf)adsot margin smooth and (left) spine of presence indicate arrows Upper arrows). (lower ribs in both structure The in (right). present surface medial concave of margin lateral Lomasuchus fti pn nterbsati aibei ohtaxa, both in if uncertain variable the is is It slight. shaft with is rib variation this the position junction but on exact spine and the this size of The at 96). the (Figure ribs of tuberculum shaft cervical the on posterior spine projecting posterodorsally a iue9.Mda iwo evclrb of ribs cervical of view Medial 97. Figure oee,tr u ob uaiesnpmrh o a for synapomorphy putative Mahajangasuchus a be may, to It out as turn clade. however, unnamed optimized larger this unambiguously for synapomorphic currently is character etba.Tec isaentpeevdin preserved is not are rib insignis ribs c6 jangasuchus c3 the preceding The the of on vertebrae. present shaft spines the the with homologized on “dorsal the noted assessment, process. protuberance” homology or present spine the a as Given projecting characterized better posterodorsal is the structure defined and well bifurcated, not is being rib as the of morphology the bisected, in intact Alligator is cervical concavity next insignis the This of process the rib. anterior for the concavity with a of contact in consists crocodyliforms “bifurcated” most of are c5 insignis of and ribs the of c4 bodies the that noted (1999:158) Brochu and 344 otrvnrlposition support posteroventral character Ambiguous crown (121-0). of condyle positioned occipital base quadrate to of ventral at condyle compressed constricted mandibular fossa laterally (117-1), teeth supratemporal teeth cheek the cheek into (116-1), all, posterior at (23-1), not or slightly, synapomorphy Unambiguous patagonicus tsangatsangana rrpscu wegeneri Araripesuchus nae ld (a.k.a clade Unnamed rib. cervical this on spine a lack aaagscu insignis Mahajangasuchus rrpscu gomesii Araripesuchus .H Turner H. A. h eilsraeo h otro process posterior the of surface medial The . ihadsic onigrmlk htse in seen that like rim bounding distinct a with Fgr 7.Sneti ocvt snot is concavity this Since 97). (Figure ,and 1 rrpscu gomesii Araripesuchus 1 Trematochampsa þ Araripesuchus Mahajangasuchus Araripesuchus but Araripesuchus 1 rrpscu tsangatsangana Araripesuchus hc rsreteelement, the preserve which , Araripesuchus 141,proximal-most (114-1), ¼ and rna xed only extends Frontal . ciia odl in condyle Occipital . cm. 1 Araripesuchus and aaagscu insignis Mahajangasuchus aeti etr,so feature, this have ld.Bclyand Buckley clade. Mahajangasuchus Mahajangasuchus Araripesuchus rc 1959)— Price Peirosaurus possess Maha- sbte ecie sasieta iucto ie h otniyo h concave the of continuity the given bifurcation a than spine a as described better is , A and (A) hsfaue oee,i uhmr rvln trait, in prevalent more present much being a is however, feature, This Discussion projecting strongly very 98). Figure head (128-1; flange distinct a forming fibular posteriorly of portion hrceitca yaoopyfor synapomorphy a as characteristic of diagnostic as fossa Araripesuchus supratemporal or the slightly, into only all, at extending not group. frontals the cited to (1959) unique is Price Unfortunately, characters these monophyly. of one only its supporting characters n fteecaatr a ciia odl in condyle and occipital notosuchian in not in (an though taxa, sebecosuchian found characters is position) these posteroventral of One fteqart.Cooyiom ntelatinclusive least containing the in clade Crocodyliforms condyle quadrate. mandibular the the of of position the in seen is forms crowns. tooth uha(lhuhi srvre in reversed is it aznarezi (although suchia oteocptlcnye hsi h odto present condition the is This in ventral condyle. positioned occipital be the to to condyles the Croco- for the for is condition with dyliformes ancestral level The situated condyle. are occipital that condyles mandibular iia to similar h neta odto fcnclteh Moreover, teeth. conical of with teeth of cheek the condition clade, ancestral Ziphosuchia the a as a bonapartei optimized Bretesuchus is for and cheek peirosaurids of synapomorphy in compression crown known Lateral the is taxa. of teeth other base in the seen at are constricted teeth cheek and nuabgosotmzto fteecaatr as characters remaining three these the for of apomorphic optimization unambiguous in clarki CTA piie hma the at them optimized ACCTRAN ol o ecddfor coded be not could eoa of Removal liao mississippiensis Alligator eeslt h rmtv odto o crocodyli- for condition primitive the to reversal A ieie aeal opesdpseircekteeth cheek posterior compressed laterally Likewise, h w hrcesabgosyspotn hsnode this supporting ambiguously characters two The Araripesuchus and and .An .wegeneri A. Araripesuchus hsaayi eoee hstraditional this recovered analysis This . iouhsclarki Simosuchus aaiuhsmwakayasyunguti Malawisuchus . Araripesuchus Alligatorium Araripesuchus rgascu aznarezi Uruguaysuchus B hwn aueo evclsie Note spine. cervical of nature showing (B) rmteaayi hrfr results therefore analysis the from h nyecpina tretains it as exception only the .wegeneri A. ihbslyconstricted basally with , ). Goniopholis ld a numerous has clade n rcdlahave Crocodylia and eeu icaeorhinus Sebecus Araripesuchus .Asaresult,only Araripesuchus Uruguaysuchus Araripesuchus , n Zipho- and Simosuchus node. taxa. are . . Downloaded By: [Columbia University] At: 05:34 17 September 2007 FN R20;D hwtedrvdsae1wt h togyiflce otrol rjcigfiua ed iua r cldt h same the to scaled are Fibulae head. fibular projecting posteriorly inflected strongly the with 1 state derived the size. show D) 2300; PR (FMNH 1 clade— Possible strongly. as posteriorly project heads iue9.Ltrlveso rcdlfr blesoigsseai aito nfiua head. fibular in variation systematic showing fibulae crocodyliform of views Lateral 98. Figure liao mississippiensis Alligator blrha steoeuiu ri for trait unique one 98). (Figure the is head fibular nrtmoa eeta(81,brbtenobtand orbit between bar (18-1), fenestra infratemporal support character Ambiguous synapomorphies. unambiguous nmiuu synapomorphy Unambiguous gomesii in seen extent the to not mwakayasyunguti rnucd raldvlpdposteriorly-directed greatlydeveloped pronounced, A rrpscu patagonicus Araripesuchus Protosuchus aaagscu insignisMahajangasuchus and rrpscu wegeneri Araripesuchus rrpscu tsangatsangana Araripesuchus aeelre blrhas u hsis this but heads, fibular enlarged have AN 16C)so tt fcaatr128. character of 0 state show CA) 1106 (AMNH Araripesuchus ua o-hpdbeneath rod-shaped Jugal . 1 .T Araripesuchus and o otefibular the do nor h e r r eetdvrial ofcltt comparison. facilitate to vertically reflected are ea Malawisuchus Araripesuchus r en o upr o hscaeaeee nw for described and known recovered is even material further are until and ambiguous clade providing this wegeneri thereby for characters support five the uprigti ru r ipybigple down characters pulled the being simply the condition, are from this group Under this supporting node. optimized characters this under any at are only setting fact, ACCTRAN the In synapomorphies. unambiguous choanal Discussion of surface (126-1). (100-0), groove from ventral trunk by absent marked of septum extend osteoderms not part (70-1), does ventral dentary fenestra (32-1), beneath on only sculpturing part with anterior narrow, fenestra supratemporal rrpscu gomesii Araripesuchus ie h urn hrce distributions character current the Given . New .gomesii A. hscaei upre yno by supported is clade This . Araripesuchus rtscu richardsoni Protosuchus AN 45;C and C) 24450; (AMNH þ .patagonicus A. rmMadagascar from AN 04 )and A) 3024; (AMNH oe oeof None node. A . tsangatsangana 345 A. Downloaded By: [Columbia University] At: 05:34 17 September 2007 gi ipybigple onfo the from down pulled are being group gomesii this simply this supporting again Under characters node. the this at condition, previous optimize setting characters ACCTRAN the the any under that only to is it clade, Similar possible synapomorphies. biguous and Discussion mandible of part posteriorly (71-6). ventral attenuating process contact from not retroarticular does projecting nasal (11-1), (4-1), bar lacrimal internarial of any, support character Ambiguous synapomorphies. biguous synapomorphy Unambiguous 1 clade— Possible unlikely. for 1 tsangatsangana clade—Araripesuchus Preferred unlikely. as clade possible this view I distributions r nw for clade known this are for support ambiguous providing characters 346 ihsot aia (104-0). carinae smooth with synapomorphy Unambiguous gomesii rrpscu tsangatsangana Araripesuchus rrpscu patagonicus Araripesuchus rrpscu wegeneri Araripesuchus .H Turner H. A. þ hscaei upre yn unam- no by supported is clade This . .patagonicus A. iue9.Rsoainof Restoration 99. Figure .wegeneri A. rrpscu wegeneri Araripesuchus ie h urn character current the Given . iwti osbecaeas clade possible this view I , oe oeo h three the of None node. et ihu aia,or carinae, without Teeth . rmxlafrsltl,if little, forms Premaxilla . hr r ounam- no are There . 1 Araripesuchus rrpscu tsangatsangana Araripesuchus A. Discussion flange distinct a strongly forming (128-1). very posteriorly head fibular projecting of (114-1), (112-1), position portion skull posteroventral proximal-most the in of broad condyle remainder relatively occipital the snout than (100-0), shorter trunk and of part ventral support character Ambiguous and with clade this for above. discussed topology topology for than alternate stronger favored the is support the character because weak. is analysis is group this this for Nonetheless, support character analysis, the scniee nuabgostatdansn the diagnosing trait unambiguous group. an considered is odto fsrae aia hti rsn nthe in present the is of that relative carinae closest serrated of condition yaoopisfor synapomorphies clade. this of synapomorphy a as snout and broad short relatively a optimizes DELTRAN addition, h rn a lodans hsgop oee,the in however, known group, not this is diagnose trait also may trunk the trait. among the widespread possessing all sebecosuchians nearly more and notosuchians some even with crocodyliforms, are carinae serrated oeti oeicuiecaedpnigo o the how on depending clade inclusive more this nose rgnlilsrto yCrlnMcKee-Freese. Carolyn by illustration Original . ntecneto h oiino hs he aai a in taxa three these of position the of context the In w hrcesdsusderira possible as earlier discussed characters Two h bec fotoem rmtevnrlpr of part ventral the from osteoderms of absence The Trematochampsa .wegeneri A. When . aaagscu insignis Mahajangasuchus rrpscu wegeneri Araripesuchus nti iuto ean h primitive the retains situation this in hi hrdls ftehcarinae teeth of loss shared their , Araripesuchus rrpscu tsangatsangana Araripesuchus Araripesuchus sedrsasn from absent Osteoderms . rprmydiag- may proper Peirosauridae, , ld.I fact, In clade. sicue in included is .In Downloaded By: [Columbia University] At: 05:34 17 September 2007 urtmoa os skont epeetin present the be from to mwakayasyunguti orbit notosuchians known the as the well as is atoposaurids separating fossa in bar supratemporal seen narrow also The is fenestra mwakayasyunguti Malawisuchus infratemporal jugal the rod-shaped The taxa. distinct beneath two these a in seen solely but not taxa both a in septum. septum. the of shallow of choana presence feature the is the of groove is surface The ventral taxa the two on the groove to Unique value. Araripesuchus Discussion from absent (100-0). trunk osteoderms of part (70-1), ventral fenestra beneath support character Ambiguous ventral by marked only septum (126-1). part groove choana anterior of on surface (31-1), sculpturing with supratemporal narrow, and fossa orbit between bar (18-1), fenestra synapomorphy Unambiguous 1 clade— Unnamed distinct a very forming being posteriorly head flange. projecting fibular a of strongly by portion the diagnosed and proximal-most condyle be occipital would positioned posteroventrally group this optimization, hscaeaekoni te rcdlfr aa In taxa. crocodyliform other to in addition known are clade this ntscu minor Anatosuchus patagonicus ru eainhpbtentetoSuhAmerican South two taxa, the sister between a relationship recovered group dataset the Unsurprisingly, results. agscu insignis jangasuchus uhsrichardsoni suchus h oohl of monophyly The of Araripesuchus relationships and membership Monophyly, Conclusions this diagnoses setting. DELTRAN osteoderms the under trunk group an ventral Lastly, of fenestra. mandibular absence the below extend not bec fteecaatrsae det isn data) missing to (due in states character these of absence aznarezi h eann nmiuu hrce upr is support character unambiguous remaining The ieie h hrcesabgosysupporting ambiguously characters the Likewise, rrpscu gomesii Araripesuchus .wegeneri A. .gomesii A. and hrce upr o ot American South a for support Character . otsbcscin swl as well as sebecosuchians most , rrpscu gomesii Araripesuchus iouhsclarki Simosuchus ld sfil toga stebootstrap the is as strong fairly is clade and rrpscu patagonicus Araripesuchus , sitrrtd ne DELTRAN Under interpreted. is , . , Araripesuchus oauscu brachybuccalis Comahuesuchus ssscu chungkingensis Hsisosuchus ntscu minor Anatosuchus .patagonicus A. ua o-hpdbeneath rod-shaped Jugal . etr osntextend not does Dentary . aednaista do that dentaries have n thalattosuchians. and sspotdb these by supported is and hsfidn has finding This . , Uruguaysuchus Malawisuchus Araripesuchus , Maha- Proto- and oooista r o akdysbpia.It suboptimal. markedly place to not steps extra two are requires that topologies itrgopo oouhaso samme of to sister as member position are a to a steps draw and Notosuchia extra as to two or required only notosuchians Additionally, of Notosuchia. group sister nlsssosta ayn h eaierelationships relative the varying between that shows analysis Araripesuchus o n oel20,PladApesteguı and Pol 2003, 2004, Pol 1997, Norell al. and et Brochu Pol (Wu 2000, analyses al. Other et 2002). al. Ortega et 2000, al. et Buckley Brochu 1999, and (Buckley matrices is character taxon and topology similar sampling having This analyses sense). previous with to broad consistent is the it than (in neosuchians to notosuchians related closely more is that nwrn hsqeto ean ifiut nthe In difficult. remains analysis, question present this Answering yaoopis t oiina lsrto closer as þ position Its synapomorphies. .gomesii A. Is champsa ntecae nti ae ihdnyi o an not as is such closely taxa ziphodonty other related with shared case, of condition retention plesiomorphic this a the simply but In species, the of clade. autapomorphy in the dentition in ziphodont of wegeneri A. phylogenetic presence preferred the study’s hypothesis, teeth this serrated of In presence (ziphodonty). the on to largely status focused taxonomic have preserved species this phenetic against are The made signal. arguments phylogenetic characters coherent a of provide number sufficient neovdsse aat hsSuhAeia clade. American South this to taxa Norell sister unresolved and Pol species 2003, sangana new Pol The 2000, 2004). al. two 2003, these et at (Ortega looking analyses taxa other by indicated been ntelc fuabgossnpmrhe sup- of synapomorphies topology alternate unambiguous the of based porting lack is preference the This on analysis. this from topology lsrt h ot American South the to closer as e o n Apesteguı and Pol see (also ooyeof holotype ihntecaeursle,teerslsare results taxon these the of in inclusion unresolved, continued the with clade consistent the within en h itrtxno other of taxon scenarios sister preferred namely possible the (2000), the being the al. et of fact, Ortega by one In suggested to and 2002). Prasad conforms 2000, Broin topology al. et de Ortega Lapparent 1990, al. et Michard The lhuhcaatrspoti ekadisposition its and weak is support character Although Araripesuchus .patagonicus A. .tsangatsangana A. . New n h African the and Araripesuchus þ sesl eocldgvnisbslposition basal its given reconciled easily is .wegeneri A. Araripesuchus .patagonicus A. Araripesuchus lsrt oouhas Sensitivity notosuchians. to closer Araripesuchus notosuchian? a than rgascu aznarezi Uruguaysuchus Mahajangasuchus sfamnay eetees a Nevertheless, fragmentary. is .wegeneri A. n oouhasrslsin results notosuchians and swl upre ssse to sister as supported well is cnr rnte l 1990, al. et Brunet (contra .wegeneri A. ´ iha aya nine as many as with , 05.T esr,the sure, be To 2005). a rmMadagascar from Araripesuchus samme faclade a of member a is rrpscu tsangat- Araripesuchus Araripesuchus Araripesuchus .wegeneri A. stepreferred the is ´ Araripesuchus 05 place 2005) a r on as found are and .wegeneri A. .wegeneri A. .gomesii A. taxa. Tremato- species u of out sthe as being 347 . Downloaded By: [Columbia University] At: 05:34 17 September 2007 348 aooi cp n aae ie ubrof number A size. dataset and scope taxonomic reconstruction). geographic (depending the Antarctica on well potentially as and America Madagascar South as in Maastrich- likely and occurrences Albian with the tian, between gap temporal the yohsspeit h rsneof unsampled presence the of predicts amount hypothesis substantial fairly Araripesuchus phylogeny a present The The implies Cenomanian. Aptian. and Albian the the American in South derived occurring more member, occurring h rsneof presence The of biogeography Historical dataset. particular a outside of ambiguous context the remain will mesoeucrocodylians ae h oiinof position is the comparison taxon comprehensive made, and a sets such Until character sampling. their various these in between analyses disparity conflicting the alleviate not does hna4 ilo erdsaiyeit ewe the basal) between exists (but disparity younger year More million relatively sampling. 40 future a than potential about predictions nw rmteMatihin n h older the and Maastrichtian, the Araripesuchus from known eprlocrecsof known the occurrences with along temporal here, presented phylogeny The thoughts Final 2004). (Turner affected overall not the exam- is mesoeucrocodylians data), ined the present from recovered the signal with inconsistent not more Araripesuchus the wegeneri if of pesuchus relationship Even the “traditional” indicate biogeographically and landmasses. that America South Indo-Madagascar scenarios between contact geographic provides prolonged it biogeographic for as crocodyliform interesting, support is overall conclusion This this pattern. with ent among topology test the extremely in to disagreement Unfortunately, among opportunity species. related the closely scenarios rare biogeographic important a during different provides an biogeography It it Gondwanan Cretaceous. make for Madagascar taxon and America, eto oteecuino fia h preferred The Africa. of for exclusion hypothesis phylogenetic the con- to a faunal indicated nection American—Indo-Malagasy data the South study, that close examined In was (2004). Turner Cretaceous by Mid-Late the during lians clade’s the interpreting problematic. history makes biogeographic trees parsimonious h otsgicn ekessi hssuyare study this in weaknesses significant most The naprn rwakt h rsn td sta it that is study present the to drawback apparent An h igorpi inlaogmesoeucrocody- among signal biogeographic The .H Turner H. A. aai nedtecrettplg i is (it topology correct the indeed is taxa itr.Tepeerdphylogenetic preferred The history. species. ssse oteSuhAmerican South the to sister as Araripesuchus Araripesuchus Araripesuchus .wegeneri A. Araripesuchus Araripesuchus Araripesuchus Araripesuchus .tsangatsangana A. nArc,South Africa, on aai h most the in taxa Araripesuchus steoldest the is mn other among aa makes taxa, sconsist- is cu in occur Arari- in , u) akNrl,DeoPladJh Maisey John Fernando and History); Apesteguı Pol Sebastian Natural Novas, of Diego Museum Norell, (American Mark hue); Paleontologı areLoadto h atmjrt fthe of majority in filled vast Ellison Mick the while Group. within, used took photographs Discussion Iowa Leonard of Systematics Carrie University and and the Smith Paleontology from Nathan Storrs, individuals James, Glenn Rich numerous Lindsay Brett, Maisano, Sumrall, Carl Jessica Colin Colbert, Sereno, Matt Paul Ketchem, Apesteguia, Callery, Sebastian John Calvo, Jorge Bell, Makovicky, Chris Forster, Pete Cathy Diego Upchurch, Scott Paul Carrano, Clark, Matt Sampson, Krause, James David Norell, from Mark Pol, assistance correspondence, conversations, general for and/or grateful am Sereno I list, Paul and and Simpson Museum); W. Chicago). of (Field (MNHN) (University Broin Resetar Buenos de Naturales, A. L. Ciencias F. de Aires); Argentino (Museo og av n .Slao(ue eGeologı de thank (Museo to Salgado like would L. and and to Calvo indebted Jorge am I material, committee—Jonathan my of Buckley. Greg and members Adrain to grateful other also am the I am my Brochu. is Christopher people I supervisor, these among them, of foremost part thesis, To large Masters a my represents individuals. work this numerous As indebted. deeply of without help accomplished been the have not could work This Acknowledgements crocodyli- in problems these on paleontology. form light shed methods, help biogeographic may of development Recent. coupled further taxa, the with crocodyliform of into census expanding up an ever to incorporating and analyses point phylogenetic Mesozoic rigorous More starting the a biogeogra- during crocodyliform phy provide of questions that America. other examine as South such and relationship Analyses biogeographic Indo-Madagascar non-tradition between a crocodyliforms that suggest found tree-fitting fruitful study is That the analysis 2004). (Turner reconciliation of tree use as such the methods indicates phylogeny form biogeographic crocodyliform overall break-up of pattern. result to the the serve have also changing would could por- which characters They relationships, unresolved tree. current potential the or of of poorly-supported tions and/or number clarify taxa the these could of in in Inclusion represented expansion analysis. poorly is current crocodyliforms the included—e.g. basal diver- the of not Additionally, sity taxa. were notosuchian numerous taxa mesoeucrocodylian rlmnr igorpi nlsso crocodyli- of analysis biogeographic preliminary A ihaooist hs ndetnl etfo this from left inadvertently those to apologies With o cest olcin n aiu undescribed various and collections to access For ´ el nvria ainldlComa- del Nacional Universidad la de a ´ n ljnr Kramarz Alejandro and a ´ ay Downloaded By: [Columbia University] At: 05:34 17 September 2007 nue T 1975. MT. Antunes References Geoscience of Department Iowa Fund. 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S. a ...s. u eainscnlsnotosu los con relaciones sus y sp.) n.g.n. , ˆ Sarcosuchus r mcrocodilı um bre ahcelscu trinquei Pachycheilosuchus ´ esdsAeio aSreBauru Serie da Arenitos dos deos rmteCeaeu fAfrica. of Cretaceous the from Araripesuchus ´ e notossu deo ´ esdlUuuy( Uruguay del ceos ´ oouhsterrestris Notosuchus nooi 88:19–71. ontologie rmtcapaoblita Trematochampsa e procoelous new a , ´ uod Creta do quio ean rmthe from remains Uruguay- ´ quidos (Arch- ´ cico ´ , ipo G 97 nacete ancient An 1937. GG. Simpson akrA.17.Arvso fteJrsi reptile Jurassic the of revision A 1970. AD. Walker crocodyliform sebecosuchian new A 2005. JO. Calvo the AH, Turner during biogeography Crocodyliform 2004. AH. Using Turner Analysis (Phylogenetic PAUP* 2002. DL. Swofford phylogenetic and structure Boston Cranial 2002. MA: HCE. Larsson Boston, H-D, Sues 2. version TreeRot, 1999. MD. Sorenson uX C htejeS 1993. S. Chatterjee -C, extant X. Wu in structures facial of Homology 1995. LM. 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Gondwana graphical of Evidence Cretaceous: Sunderland, 4.10b. Inc. Version Associations, Sinauer Methods), MA: Other and Parsimony crocodyliform 22:112A. enigmatic the rebouli of relationships University. 965:1–20. Novitates Museum Am Patagonia. aaaa nuaiiyadnslcnhe Morphol J conchae. nasal to and reference 225:269–327. special pneumaticity with crocodilians), paranasal and (birds archosaurs 30:321–336. Michigan Museum Univ Contrib Paleontol, (Pakistan). Pab Balochistan Cretaceous Vitakri, Upper of Formation the from Mesoeucrocodylia) (Reptilia, Occas 106:1–37. Kansas Africa. Univ southern Hist, Nat (Basutoland), Museum Lesotho Pap of Triassic the Vert J Madagascar. 24:128A. Paleontol Formation, Maevarano Cretaceous Upper 19:375–514. Berlin in Museum Zoologischen dem aus A Abteilung Palaeontogr 138:133–165. Bayerns. Oberjura-Plattenkalke der 257:323–372. Biol Philos Lond: crocodiles. Soc of R classification Trans the on remarks with (Marsh), omnso t hlgntcrltosis etPaleontol Vert J relationships. phylogenetic 21:492–514. its on comments and Canada, Saskatchewan, from material New Crocodyliformes): etPlsaia32:166–180. Palasiatica Vert oop rmteLwrJrsi rCiaadtepyoeyof 13:58–89. phylogeny Paleontol the Vert J and Sphenosuchia. China the or Jurassic Lower the from lomorph oso ne ogla(epesRpbi fCia.CnJEarth 33:599–605. J Sci Can China). of Republic (People’s Mongolia Inner of Cretac-eous Lower the from Crocodyliformes) (Archosauria: suchian Paleontol Vert J China. Hubei, of 16:688–702. Cretaceous Lower the from of Vert J China. Yunnan, of 16:42–48. lower Paleontol Jurassic) the (Lower from Formation Crocodyliformes) Lufeng (Archosauria: 1944 Young, oms omScun(hn) n h oohl of 1:1–87. Sin Paleontol monophyly Acta China. the and 17:89–103. (China), Paleontol Vert J Sichuan Protosuchia. Crocodyli- form (Archosauria, protosuchian formes) Cretaceous Early new hmeauhsparadoxus Chimaerasuchus New rmteCeaeu fMrco etPaleontol Vert J Morocco. of Cretaceous the from Araripesuchus nuuulcooyiomreptile crocodyliform unusual an , iohouhselaphros Dibothrosuchus rmMadagascar from ihaouhsshuhanensis Sichuanosuchus scincooiefrom crocodile usuchian erminonaris i r a n o n i m r Te ltontu hsui Platyognathus Hamadasuchus alpsvictor Hallopus (Archosauria: crocody- a , Hsisosuchus 351 ,a . Downloaded By: [Columbia University] At: 05:34 17 September 2007 pedxA1 hrce itue in analysis used phylogenetic list Character A.1: Appendix 0 rnasnro ewe ris(iia nbreadth in (similar orbits between narrow Frontals 20. a only contacting dorsally, narrows Quadratojugal 19. temporal lateral beneath flattened transversely Jugal 18. (0) part posterior as broad as jugal of part Anterior 17. (0), bar postorbital on jugal to anterior Postorbital 16. contact not does prefrontal of (1) process Descending not does or (0) premaxilla 15. (1) contacts not Nasal does or (0) border narial in part 14. takes Nasal 13. (0) only edge medial along (1) nasal contacts not Lacrimal do or (0) lacrimal 12. contact Nasals palate on meet 11. not do maxillae of ends Posterior 10. 352 .Peail n ail ihbodcnato face, on contact broad with maxilla and Premaxilla 9. or (0), face on maxilla overlies loosely Premaxilla 8. posterior meet not do premaxillae of parts Palatal 7. laterally nares vertical, premaxilla of part Dorsal broad, or 6. (0) nares to anterior narrow Premaxilla 5. internarial of half ventral least at forms Premaxilla tubular nearly or 4. (0) wide than higher Rostrum broadening orbits, 3. to anterior narrow Rostrum 2. mandibular and cranial of surfaces External 1. onsl)()o r ra,aottienasal twice about broad, are or (1) breadth (0) nasals) to quadratojugal (1) bar postorbital or of of portion most (0), postorbital contacting sheet broad postorbital a as dorsally of extends part small (1) fenestra beneath rod-shaped or (0) fenestra (1) part posterior as broad as twice about or postorbital or (1), (2) jugal jugal to lateral to medial postorbital (2) suture robust in palate contacts or (1), palate contacts or (0), palate (1) edges anterior and medial on or (1) meet do ends or (0), palatines to anterior con- directed laterally (3) cavity broad, forming maxilla, and with premaxilla with contact at contact constricted rostrum or at (2), slit narrow constricted forming maxilla, rostrum and premaxilla or (1), illa premax- and maxilla between broad, notch or open (0), laterally contact at narrow not does rostrum (1) joint butt along together sutured maxilla and premaxilla along (1) posteriorly maxillae meet with contact or (0) foramen incisive to dorsally or dorsolaterally (1) nearly oriented premaxilla nares of part horizontal, dorsal or (0), oriented (1) nares to lateral part the to breadth in similar (1) bar internarial of any, if little, forms or (0) bar (2) high than wider or (1) (1) throughout broad or (0) orbits at abruptly [Pol 1] (2) character pits 2003; and grooves deep with heavily or ornamented, (1), grooved slightly (0), smooth bones .H Turner H. A. 0 aaa ufc fpeyodsot 0 or (0) smooth pterygoid of surface Palatal opens or 40. (0) palate from ventrally opens Choana 39. suspensorium, and palate to restricted Pterygoid 38. narial below palate on meet not do Palatines or 37. (0), flat nearly squamosal of edge Posterior 36. 5 otrdra onro qaoa qae off, squared squamosal of corner Posterodorsal 35. distinct a (dermosupraoccipital) Postparietal 34. separating region sculpted broad, with Parietal 33. with or (0) portion occipital broad without Parietal 32. broad fossa supratemporal and orbit between Bar 31. dorsally, broadens bar postorbital of end Dorsal 30. lateral and anterior with postorbital of part Dorsal or (0) 29. process anterolateral without Postorbital of part 28. dorsal of edge lateral on opening Vascular 27. unsup- flattened, transversely bar Postorbital sculpted 26. surface lateral weak, bar Postorbital 25. 4 urtmoa ofwt ope dorsal complex with roof Supratemporal 24. or (0) fossa supratemporal into well extends Frontal with or 23. (0) flat parietal and frontal of surface (1) Dorsal fused or (0) paired 22. Frontals 21. cltd(1) sculpted (1) depression midline into posteriorly (1) laterosphenoid of quadrate contactand to edge sutured to strongly ventrolateral foramen, dorsally trigeminal form extends and pterygoid laterosphenoid or (0), overlapping basisphenoid and quadrate with joints (2) palate not secondary of do part that forming passage, shelves tonarial meetventral palatal meet(1),or form or (0), passage (1) process long a as poster- iorly extends squamosal of edge posterolateral lmn 0 rntdsic fsdwith “lobe” withunsculpted (fused or (0) “lobe” extra lacking distinct not or (1) parietal?) (0) element supra- between crest sagittal (1) fossae with temporal or (0) fossae (1) portion occipital broad part anterior on (1) sculpturing only with narrow, bar or (0), surface dorsal sculpted broadly with solid, and (1) postorbital of part dorsal distinct constricted, from bar postorbital or of (0), part postorbital dorsal of part dorsal with continuous edges (1) facing anterolaterally with or (0) only edges (1) process anterolateral with (1) present or (0) absent bar postorbital bar postorbital (1) ectopterygoid or by supported (0), columnar, ectopterygoid by ported robust, bar postorbital (1) or unsculpted (0), sculpted) skull (if ufc 0,o oslyflt“kl al”developed, table” “skull flat dorsally or (0), surface (1) all at not or slightly only extends (1) ridge midline narrow (1) xedn aeal eodqart otcs(1) contacts shelves quadrate beyond flat laterally extending with postorbital and squamosal with Downloaded By: [Columbia University] At: 05:34 17 September 2007 0 tciia ihu ag etoaea atventral part ventrolateral large without Otoccipital 60. large common through pass IX-XI nerves Cranial 59. descending concave laterally without Otoccipital bilateral 58. well-developed without Basioccipital 57. brain- of surface ventral on exposed Basisphenoid 56. basioccipital, to length in similar Basisphenoid 55. mova- forming prominent, process Basipterygoid 54. slen- process) between (cultriform rostrum Basisphenoid enclosed 53. not tubes Eustachian 52. contact not does quadrate of part Ventromedial 51. edge ventral flat with quadrate of ramus Pterygoid 50. meet not do otoccipital and squamosal, Quadrate, 49. quadrate with otoccipital of contact Ventrolateral 48. with articulates quadrate of head primary Dorsal, to 47. medial broad quadrate of edge Posterior single 46. with or (0), fenestrae without Quadrate 45. pterygoids of edge anterior near situated Choanae enclose 44. or (0) choanae enclose not do Pterygoids 43. of one-fourth than less size, in moderate Choana 42. are or (0) choanae to posterior separate Pterygoids 41. rlpr (1) ventrolat- part large eral with or (0) process paroccipital separate to in XI and X nerves (1) passage to medial IX passes nerve cranial or (0), otoccipital in vagi foramen with or (0) (1) flange process subcapsular to ventral flange (1) tubera pendulous large, with or (0) tuberosities (1) basioccipital and pterygoid surface ventral by or from excluded virtually (0), or (0) case surface ventral concave (1) basioccipital than shorter or basisphenoid flat with basipterygoid joint or (1) basipterygoid suturally with closed (0), absent, or pterygoid small process with joint ble (1) expanded dorsoventrally or (0) der entirely or (0) (1) enclosed basisphenoid and basioccipital nerves (1) cranial IX-XI for passage enclose form to and artery otoccipital carotid contacts or (0) otoccipital (1) edge ventral along groove deep with or (0) meet or (1), skull (2) passage of to lateral edge broadly enclose lateral (0), near passage passage cranioquadrate enclose to (1) broad or (0) narrow very with or (1) laterosphenoid (0) and prootic prootic and otoccipital, squamosal, strongly contact, otoccipital (1) concave to dorsal edge posterior narrow or (0), concave gently tympanum, on fenestrae more (2) surfaces or posteromedial and three dorsal with or (1), fenestra (1) pterygoids of middle in or (0) anteriorly) (or (1) choanae (1) breadth skull of half nearly extremelylarge, choana or (0), breadth skull (1) fused 0 neirdnaytehopst premaxilla-max- opposite teeth dentary Anterior 80. of edge lateral with homodont, teeth Maxillary to 79. size in similar teeth premaxillary two Posterior 78. involved or (0), symphysis in involved not Splenial 77. does posterior pterygoideus M. arched for area Insertion or (0) 76. flat surangular 7 5 n of d .Ma i edge b u l Dorsal a rf e74. n e s t r ap r e with s articulating (1) process e medial n absent without t( Articular or 0 )o (0) ra present b73. s Prearticular e n t( 172. ) (0), robust and short very process Retroarticular 71. mandibular beneath (1) posteriorly septum extends by Dentary divided or (0) confluent 70. Choanae than 69. longer much fenestrae Supratemporal 68. about or (0), orbit as (1) large confluent as fenestra or Antorbital (0) divided nares 67. External 66. two or (0), orbit in present palpebral small One (0) 65. flat nearly supraoccipital of surface Posterior 64. supraoc- into extend not does antrum Mastoid 63. foramen of edge dorsal forms Supraoccipital 62. pseudo- fenestrae between interfenestralis Crista 61. te etr et 0 rmr hntiethe twice of than length more the or (1) twice length (0) than teeth more dentary no other tact con- illa teeth maxilla or of waves two (1), (2) in (“festooned”) edge curved loci maxilla of edge with these the and enlarged at in row, outward enlarged tooth extending teeth of or middle (0), straight maxilla (1) longer much or (0) teeth anterior (2) symphysis in extensively involved or (1), symphysis in slightly or (1) angular (0) of angular surface lateral of onto surface extends lateral onto extend not (1) dorsally process with (1) or (0) basisphenoid and otoccipital attenuating 1994] and Clark projecting from mandible [modified of (5) posteriorly part ventral or and from (4), and curving projecting ventrally posteroventrally paddle-shaped posterodorsally and or robust, (3), or elongate short, (2) or situated (1), absent or fenestra beneath extend not (1) does or (0) fenestra than shorter or to length in (1) much orbits equal or or (0) (1), orbits (3) absent orbit or the (2), orbit of than diameter smaller the large half one or (1), (2) present present palpebral palpebrals large (1) prominences posterior bilateral with or (1) regions ear middle in connect canal to transverse supraoccipital through extends or (0) cipital (1) supraocci- foramen separating from magnum, pital foramen the dorsal meet or to broadly otoccipitals or (0) (0), magnum vertical nearly ovalis (1) horizontal and rotunda New Araripesuchus rmMadagascar from 353 Downloaded By: [Columbia University] At: 05:34 17 September 2007 0.Atro agno eu ier() ranterior or (0), linear femur of margin Anterior 103. glenoid of wall lateral the only dorsal forms Surangular on 102. keels longitudinal or with trunk Osteoderms of (0) 101. part ventral only from osteoderms absent or Osteoderms dorsal 100. (0), with imbricated Tail osteoderms all 99. or Some 98. longitudinal parallel, two in osteoderms Dorsal 97. edge anterior straight with osteoderms Dorsal 96. 354 5 oslosteoderm Dorsal 95. amphiplat- or amphicoelous vertebrae caudal All amphiplatyan 94. or amphicoelous vertebrae Trunk 93. amphiplatyan or amphicoelous vertebrae Cervical 92. well-developed without vertebrae Cervical 91. as vertebrae cervical posterior on spines Neural 90. long oras phalanges thanlong(0) broader (1) intercentrum Atlas without or (0) for with digit 89. facet pedal Fifth lateral large with 88. femur for end Distal of 87. edge ventral of half anterior forms Pubis or 86. (0) end distal expanded without rodlike, to Pubis length 85. in similar ilium of process Anterior scapula of 84. length the half than more no Coracoid 83. symmetrical edges scapular posterior and Anterior pre- 82. opposite tooth to posterior teeth Dentary 81. agno eu er ag o caudofemoralis for flange bears femur of margin and [Buckley 102] approximately character (1) 1999; fossa forms Brochu glenoid the surangular of or one-third (0), fossa (1) keels without or (0) surface (1) present or (0) (1) osteoderms by surrounded completely (1) another longitudinal one to two sutured osteoderms than more in (1) or rows (0) rows anterior on laterally (1) process edge anterior with or (0) (2) square or (1), long than broader rectangular, te adlvrert rceos() rall or (1), (2) with procoelous procoelous vertebrae biconvex, caudal vertebrate vertebra caudal caudal first other or (0), yan (1) procoelous or (0) (1) procoelous or (0) (1) physes hypapo- well-developed with or (0) hypapophyses (1) rodlike narrow, or anteroposteriorly (0) vertebrae cervical anterior on those as broad (1) broad as (1) facet small very with or (0) fibula the (1) of ischium process anterior an by acetabulum the from excluded partially least at pubis or (0), acetabulum (1) end distal expanded with the less (1) or process posterior one-quarter of length or (0) process posterior (1) scapula to length in equal about or (0) (1) strongly edge more posterior edge than concave anterior or (0), view lateral in (1) row tooth maxillary in teeth smaller opposite enlarged or (0) homodont contact maxilla-maxilla .H Turner H. A. rounde s ,oae() or (0), ovate d, 2.Qartjglde o ompr ftequadrate the of part form not does Quadratojugal 120. its with midline along parasagittally run Palatines 119. curved gently or straight view dorsal in row Tooth 118. or (0) crown of base at constricted not teeth Cheek laterally 117. or (0), conical teeth cheek Posterior palate 116. on (1) exposed not or (0) exposed Vomer or 115. (0) position posterior in condyle Occipital 114. multi- (0), multicusped not teeth cheek Posterior 113. shorter and broad relatively or (0), long Snout 112. or (1), short and narrow or or (0), broad Prefrontals (0), 111. symphysis to posterior thin Splenial 110. without continuous, dentary of surface transversely Lateral 109. or (0), dentary Compressed 108. edge dorsal or (0), straight dentary of edge Dorsal 107. of length the twice than more no blade Scapular 106. (0), alveolus seventh to lateral smooth Dentary 105. carinae smooth with or carinae, without Teeth 104. 1 Tre n av 05 hrce 120] character 2005; condyle Calvo quadrate and [Turner the (1) of part forms or (0) condyle palatines (1) 119] character caudally 2005; or Calvo rodlike and [Turner (0) becoming laterally passage diverge nasopharyngeal the of surface ventral the in participating length entire al. et Ortega 15] character from 1996; [modified (1) sigmoidal or (0) character117) 2000; constricted(1)(Buckleyetal. 116] character 2000; al. et [Buckley (2) spatulate strongly or (1), compressed 115] character 2000; al. et [Buckley from [modified 32] character 1997; (1) Gomani position (2) posteroventral row one than 46] multi- character more 1997; or Gomani in from (1), [modified cusps row single with in cusped cusps with cusped [Gomani (2) 6] skull character 1997; of remainder the and narrow than or shorter (1), skull the of remainder the than 4] character 1997; [Gomani (2) long and narrow 7] character (1) 1996; al. symphysis et Ortega to from [modified posterodorsal robust splenial 5] of character 1996; [modified al. surface (1) et Ortega groove lateral from longitudinal or with (0), dentary groove longitudinal (1) high 2] character 1996; as al. et wide Ortega from as [modified almost dentary, expanded 1] character 1996; al. et Ortega from (1) [modified waves concave two with sinusoidal, dentary of 106] the (1) character twice 1999; articulation Brochu than scapulocoracoid and [Buckley the greater of and length broad very scapular blade or (0), articulation scapulocoracoid the to lateral 1999; pit 105] Brochu character and occlusion [Buckley (1) large alveolus seventh with dentary or Brochu and [Buckley 104] (1) character serrated 1999; teeth or (0) 1999; Brochu and 103] [Buckley character (1) musculature Downloaded By: [Columbia University] At: 05:34 17 September 2007 ?? ?? 0? ?? 01 01 10 10101 01001 ?001? ?0010 ????? ?0?? ?00?? 01000 11000 51000 ????0 03100 ?1??1 0???? 00??0 12110 11110 00010 11120 1?0?1 1100? 11?11 11000 0000? 01101 2000? icaeorhinus Sebecus 0?100 10100 01??? ?0011 11101 ?0?1? 5?010 ?1?? ????? 03?01 ?2?10 0???? ?11?? ????? 0???? 00??0 0???? 0?011 11?10 1??11 11120 11010 11??1 ??00? 01111 10000 pacheoi Baurusuchus 10010 01011 00000 ?0??0 ????0 00?0 ?110? 00001 000?? 5?010 1111? ????0 ?110? 0101? 00?10 11?10 01120 02?10 00000 11002 0?011 11111 11010 01001 21101 10210 terrestris Notosuchus analysis phylogenetic in used Codings A.2 posterior the in shaft of process Posterior 129. straight head fibular of portion Proximal-most 128. slightly only or same the tooth maxillary Second 127. slightly or smooth septum choanal of Surface 126. row tooth upper of portion anterior of Position 125. or (0) absent lower) (upper, diastemata Median 124. absent or small oralis intramandibularis Foramen 123. 2.Atrol,pltnsaetasesl a n do and flat transversely are palatines Anteriorly, 122. positioned quadrate of condyle Mandibular 121. evclrb ak 0 rpsess()a (1) possesses [new] process or junction tubercular the the (0) with at spine lacks projecting posterodorsally ribs cervical very or distinct [new] (1) (0) forming flange posteriorly, posteriorly projecting developed sharply weakly to character 2005; Calvo maxillary and other 127] [Turner much (1) or than (0) teeth (hypertrophied) teeth maxillary larger other 2005; than (1) larger Calvo groove and by Turner 126] marked character from or [modified 2003; (0) al. depressed et [Sereno row 75] tooth character dentary from ventrally (1) and labially offset or (0) to adjacent 74] character 2003; al. et [Sereno (1) present 8] character 1996; al. et Ortega from [modified (1) slot-like and large or (0) o hrl iietemxla 0 raenarrow are maxillae the or dividing sharply “arrow”-shaped, (0) and maxillae the divide sharply not character 1997; with al. level et 124] [Wu on (1) or condyle (0) occipital condyle occipital to ventral 1 Tre n av 05 hrce 122] character 2005; Calvo and [Turner (1) 0? ?0 ???? 00110 0000? 0??00 00100 00000 00??0 00000 10001 3?000 000?1 1200? 111?0 10000 ?1?00 00101 12000 01011 11110 10100 00110 1000? 00110 02110 11101 011?0 Teleosauridae 00100 10000 10001 ???? 00001 00000 00010 00??0 11?00 00001 00000 01101 00000 30000 1200? 12000 ?1000 01001 20101 11110 00110 10000 00110 12110 11101 211?0 Pelagosaurus ?00? 10011 00001 4?001 ?0000 01100 0210? 00000 00000 11001 10010 ??1?1 011?1 11??0 00??? 11011 11011 11110 01010 10000 ?11?0 10001 01001 02110 1?131 21210 pusillus Theriosuchus 10?00 ??000 010?? 01100 00000 ?00? ?0010 11011 ????1 01?10 0???0 4?001 11100 100?1 0?1?? ????1 1?00? 00??? 1??10 10111 ?11?? ??00? 10000 0000? 1?03? 21210 Alligatorium ?0?? ?0000 11?0? ??0?? ?000? ?00?? ?1100 ????? ?2?1? ????? ???10 ?1010 ????? ????1 ????? 02?1? ????? ??1?? 0???? ??1?? ????? ????? ???2? ??002 1?121 212?? wegeneri Araripesuchus 1?001 01011 11100 1011 00100 00000 11010 ?0000 51000 100?1 0?11? 1?1?0 11?01 ?1111 00??0 11?10 12??0 11?20 10010 10011 10001 10111 11110 00002 1?121 21200 gomesii Araripesuchus 11?00 0101? 1?100 ???1? 00100 51000 1?11 000?0 ?2111 00000 10??? ?1?01 12??0 ?1??? 00??0 10010 ?1?11 10011 11?10 10111 1?1?? 11110 10001 00??? ????1 212?? patagonicus Araripesuchus 11000 01110 11100 0011 ?0100 00000 ????? 11010 1000? 61000 11?00 02110 11111 ?1101 00??0 11?10 121?0 1112? 00010 10011 10001 10111 11010 1?002 1?121 21210 tsangatsangana Araripesuchus New Araripesuchus rmMadagascar from 355 Downloaded By: [Columbia University] At: 05:34 17 September 2007 356 10 10 0? 01 00 00 00 00000 ?0?? 20000 ?00?? 00100 0000? 00000 ?000? 10010 011?0 000?1 11100 ?1?01 01000 00??1 11111 00?00 01100 00010 00001 00000 00110 10010 1?000 00110 200?0 strombergi Orthosuchus ?0?? ?00?? 0?000 0200? ????0 0000? ????? ?0000 ???01 ?00?1 2?11? ????1 ?11?? ??1?1 ??002 00101 ???01 11?11 0011? 0?001 0?00? ?0??0 10010 ????? 0?1?? ?00?? leali Hemiprotosuchus 20110 011?1 ?000 0200? 01101 00000 0000? 00000 00001 10011 00101 000?1 11100 01000 11111 01101 01101 00101 0011? 00002 00001 00110 00010 0000? 00010 00000 richardsoni Protosuchus 10000 01011 13100 11111 ?000 10000 11100 11101 00000 00010 00001 10010 11111 01100 11001 11110 11010 10011 11120 31000 11000 10110 00002 02110 11131 21201 Crocodylia 0?010 00001 4?001 00001 ?0?? 01100 131?? 10000 00000 01011 ?1??? 00021 02??? 11??0 100?0 000?0 11?1? 11?10 1?001 1112? 10?11 11000 ??000 ??00? 11131 212?1 fagesii Bernissartia 00000 00001 01100 ?0?0 10000 ?1010 10000 100?1 4?001 ?00?1 1311? ?1??0 ?1?01 11?1? 10?10 02?10 11?10 00010 1?001 11120 10111 1000? 11000 ?010? 1?131 212?1 Goniopholis 00??0 00001 01100 ?0000 ????? ???0 01010 1??00 4?000 ??0?1 12110 ?1??0 ?1?0? 111?? 10??0 11?10 01110 00010 11120 1?00? ?000? 10011 11000 00102 11101 212?? delfsi Eutretauranosuchus 00??0 ?0001 00100 ?0000 ??0? ????0 0??00 000?? 12000 30001 ?11?0 1200? 00101 ?1?00 01011 11110 02110 00110 00110 ?000? 00000 10000 20101 0011? 11101 011?0 Metriorhynchidae .H Turner H. A. ?? ?0 0? 1? 01 00 11 21000 11110 ?0000 00?10 00?0 010?1 00000 ?0000 000?2 41010 02111 ???01 ?1?01 0???? 00011 01??? 11?10 00010 1?110 0?011 11000 11011 00010 1?00? 01100 11201 clarki Simosuchus 00011 10?10 1101? ???00 ?0?00 00?? ????? ?1010 ????? 51000 12111 ????? ?2110 ?1?0? ????? 100?? 00?10 11?11 000?1 11120 10111 11001 11?10 1?002 2?101 2121? minor Anatosuchus ?0?? 00011 10010 ????? 11011 ????? ??00? 0???? ???00 ?1010 ????0 ????? ?1??1 1?1?0 02??? 1???? ?0010 ????? ????? ????? ??1?? 111?? ????0 1?00? 2?101 11??0 brachybuccalis Comahuesuchus 0000 00000 11?00 1121? ?0??? 1??0? 0???? ?111? ????? 41?00 100?? ?2110 ????? 12??0 ????1 00010 00??? ????? ?011? 0?11? 11111 10002 11110 0100? 0?101 110?0 mwakayasyunguti Malawisuchus 0000 ?0000 21000 ????? ?011? 1???? ?0??? 0??1? ??11? ??000 ????0 ????? ?111? ?0001 ????? 12?1? ????? 1??1? ????? ??01? 1???? 1???? ?10?1 ??00? ??10? 111?0 aznarezi Uruguaysuchus ???? 1???? 000?1 ????? ???0? ?1??? ?1100 5?0?? ?1111 ?10?? 1???? 0?0?? 10002 ?1?0? 00010 ????? ?0000 1?011 11??? 10011 1112? 11??0 ????1 00??2 011?1 212?0 Trematochampsa 00000 2000? 000?? 00?00 ?1?10 ?0011 ?0?? 1?000 00000 ????1 ?1001 1???? ?1?02 12110 00?0? 010?0 00010 11?10 ?0011 ????1 01?11 10011 10001 00010 1?00? 11?1? ????1 200?? ????? chungkingensis Hsisosuchus ????? ????? 1?1?? ???0? 00?? 00000 ????? 2010? 0(0&1)0(0&1)(0&1) 1?0?? 0?00? ????? 11110 12110 00010 ?1120 1?011 11001 10011 ?10?0 1?002 11121 212?1 Peirosauridae ?001 ??0?? ????? 10000 000?? 11?01 ????? ??0?? 11?11 11001 ????? ????0 ?1111 ????? 51010 010?? ????1 ????? ????? ????? ????? ????? ????? ????? ????? 2???? insignis Mahajangasuchus Downloaded By: [Columbia University] At: 05:34 17 September 2007 A8654) UA insignis 283) MUC-PV Mahajangasuchus 270, MUC-PV 269, PV patagonicus Araripesuchus gomesii Araripesuchus tsangatsangana Araripesuchus 700) wegeneri Araripesuchus GDF603) minor Anatosuchus clarki Simosuchus 1997) Gomani mwakayasyunguti Malawisuchus aznarezi MOZ Uruguaysuchus 31, MACN-N 30, 6131P) MACN-N 202, MUC-PV brachybuccalis 1044, MPCA-PV250) Comahuesuchus MACN-RN 249, MPCA-PV 287, MACN-RN1043, MUC-PV 1042, MACN- 1041, RN MACN-RN 1040, MACN-RN 1037, 1994) terrestris al. et Wu Notosuchus 1994, al. et Li chungkingensis Hsisosuchus stormbergi Orthosuchus leali Hemiprotosuchus 3024) AMNH richardsoni Protosuchus were the that after added specimens are reference. author bibliographic the the by of first-hand reviewed numbers Collection phylogenetic in analysis used taxa Fossil B: Appendix ????? ????? ?1?? ????? ????? ????? 10??? ?21?1 ??0?0 ????? ????? ???1? ????? 0???? ????? ????? ????? ????? ????? ????? ????? ????? ????? ??00? 0111? 1?101 pakistanensis Pabwehshi ???? ????? ??100 011?? 10000 ?1?00 ?1?1? ?00?1 ?10?0 00?1? 0?11? ?0??? ?1??? ?0??? ???1? 1?001 ????0 ????? ????0 ???2? ??0?? 1??1? 11?00 0?002 01111 1000? macrodon Iberosuchus ?1?? ?1?00 0010? ????? ?10?1 ????? ?1011 1000? 0???? ???10 ?1020 ???10 5??10 ????? ????? 03?0? ????? ????? ????? ????? ????? ????? ????? ??00? 01111 10100 bonapartei Bretesuchus ????? ????? ???? ???? ????? ???0? 0???? ????? ?101? ?1??0 ???0? ????? ????0 0?001 ????? ????0 ????0 ????? ????? ????? ????? ????? ????? ????? ????? 2???? enderi Pehuenchesuchus Bclye l 00 A8679) UA 2000; al. et (Buckley Bnpre1971) (Bonaparte Pie15;AN 24450) AMNH 1959; (Price Srn ta.20;MNN 2003; al. et (Sereno Gsaii17;MACN-RN 1971; (Gasparini Bfeat18;MNHN-GDF 1981; (Buffetaut Ns 1975) (Nash Rsoi1933) (Rusconi CletadMo 1951; Mook and (Colbert BclyadBoh 1999; Brochu and (Buckley Otg ta.20;MUC- 2000; al. et (Ortega YugadCo 1953, Chow and (Young peetpaper) (present Cake l 1989, al. et (Clark Bnpre1991; (Bonaparte pedxC pmrh it(iue100) (Figure list Apomorphy C: Appendix 1999) Brochu 1994, (Clark Crocodylia 1990) Clark and fagesii Bernissartia ciliosus 1994; (Clark Metriorhynchidae 5138) 1994; AMNH (Clark Stenosauridae 1994) Clark typus 620) AMNH 5782, Pelagosaurus AMNH 1999; al. et 8028) Goniopholis CMNH 570, AMNH 1994; delfsi Eutretauranosuchus Alligatorium 2000) al. 1955, et Ortega Price pusillus 1991 Theriosuchus 1991) al. al. et et Gasparini (Gasparini Peirosauridae 3160) AMNH icaeorhinus Sebecus 2000) GSP-UM pakistanensis Pabwehshi MAU-PV-CRS-440) enderi Pehuenchesuchus 2000) macrodon Iberosuchus bonapartei Bretesuchus pachecoi Baurusuchus Trematochampsa eie ttsfrcae within clades for states Derived ** icse ndphi h et h reue to used tree 100. Figure The in * text. shown is the apomorphies these in derived depth in discussed Mesoeucrocodylia: 11,23(1), 21(1), 41(1), * oe7 2(1), 7: Node 111(2) 79(1), * 85(1), ** Thalattosuchia: ssscu chungkingensis Hsisosuchus * * * Teleosauridae: 36(0), Teleosauridae ** Pelagosaurus * 91,67(2), 59(1), 76(1), 62,18(1), 16(2), 29(0), 61(1), 61,77(2), 76(1), ¼ 29(1), 30,101(1) 63(0), ¼ CTA piiainonly optimization ACCTRAN MH997) AMNH ETA piiainonly optimization DELTRAN New * ** 99(0), * * Mo 92 lr 96 94 Salisbury 1994; 1986, Clark 1942, (Mook * 78(0), 91,82(0) 59(1), 51,4() 55(0), 49(1), 45(1), * Wlnoe 91 lr 96 1994) 1986, Clark 1971, (Wellnhofer ** 30,6() 66(1), 65(0), 63(0), 30(0), Araripesuchus 51,5() 65(2), 50(1), 45(1), : Bfeat17,Rsusn2002) Rasmusson 1976, (Buffetaut * ** þ * * * 25(1), Bsain n az19,Norell 1990, Sanz and (Buscalioni 78(0), * * 108(1), 16(1), ** 16(2), erohnhde () 33(1), 1(0), Metriorhynchidae: 71(3), 115(1) * Smsn13,Clet1946; Colbert 1937, (Simpson Oe 89 lr 96 1994; 1986, Clark 1879, (Owen ** EdsDsoghms1863, Deslongchamps (Eudes * 3(1), Pie1945) (Price 19(0), Gsaiie l 1993) al. et (Gasparini Atns17,Otg tal. et Ortega 1975, (Antunes 50,4() 50,57(1), 55(0), 47(0), 45(0), 3(1), * Wlo ta.20;cs of cast 2001; al. et (Wilson 6(1), * Tre n av 2005; Calvo and (Turner Mo 97 lr 1986, Clark 1967, (Mook ** * 91,3() 72,39(1), 37(2), 36(1), 29(1), ** 20(1) ** ** ** 99(0), rmMadagascar from 11,16(0), 11(1), : * 115(1) ** 76(1), 01,21(0), 20(1), 7(1), 01,24(0), 20(1), * tnoarsbollensis Steneosaurus () 31,14(1), 13(1), 6(1), erohnhssuper- Metriorhynchus 7(1), ** * Araripesuchus * ** 68(0), 108(1), ** ** 61(1), * 9(0), 80,70(1), 68(0), 80,83(1), 78(0), () 10(1), 9(0), ** ** * * * * * 115(1) 68(0), 59(0), 19(0), 25(1), 25(0), 71(3), 357 are Downloaded By: [Columbia University] At: 05:34 17 September 2007 iue10 reue odrv pmrhe nApni .Ti so fte1 otprioiu re umrzdi iue9.Arrows 93. Figure in summarized trees parsimonious most 18 the of on is This C. Appendix within in topologies apomorphies alternate derive to two used indicate Tree 100. Figure 358 91(1), oe6: Node 100(0) ** 21,84(1), 72(1), oe4: Node 11,107(1), 81(1), oe5: Node Metriorhynchidae: 51,4() 60(0), 49(2), 45(1), .H Turner H. A. ** 62,103(1), 96(2), ** * ** () 52,1() 46(1), 17(1), 15(2), 3(2), 9(), 3(2), * 96(0), ** * 108(1), 19(1), ** ** 6(1), * 112(1) 16(2), * ** 0() 106(1) 104(1), * ** 121(1) 61(0), * 29(), Araripesuchus () 61,79(1), 26(1), 9(2), ** 01,75(1), 20(1), * * 81,71(5), 68(1), 90,69(1), 59(0), . ** 25(0), Araripesuchus 113(1) tsangatsangana Araripesuchus 31(1, * rrpscu tsangatsangana Araripesuchus .patagonicus A. rrpscu gomesii Araripesuchus ** 121(0), 114(1), * 70(1), * 128(1) ** 128(1) 23(1), : ** : 0() 126(1) 100(0), * 100(0), þ * 1() 1() 117(1), 116(1), 114(1), .patagonicus A. () 11,71(6), 11(1), 4(1), : * 104(0), þ .gomesii A. ** :18(1), 112(1), þ Downloaded By: [Columbia University] At: 05:34 17 September 2007 esci:9(3), Neosuchia: ** 71(4), rrpscu wegeneri Araripesuchus oe3: Node Trematochampsa rrpscu patagonicus Araripesuchus rrpscu gomesii Araripesuchus Alligatorium * * 11,35(1), 31(1), Theriosuchus Alligatorium Peirosauridae: ** oe2 5(1), 2: Node ** Eutretauranosuchus * Crocodylia Ziphosuchia: Bernissartia 97(1) 96(0), Mahajangasuchus 71,7() 78(1), 74(1), 67(1), terrestris Notosuchus 112(0) * Uruguaysuchus 70(1) * Anatosuchus * 61,9() 11,9() 93(1), 92(1), 91(1), 90(1), 76(1), Goniopholis Mahajangasuchus 110(1) Eutretauranosuchus ** ntscu minor Anatosuchus rcdla 31,44(1), 43(1), Crocodylia: * ** Uruguaysuchus oe1 () 12(1), 1(2), 1: Node 116(1) ** Anatosuchus ** * ** ** Uruguaysuchus iouhsclarki Simosuchus 113(2) 101(1), 91(1), 79(1), 78(1), 46(0), Malawisuchus ** 112(0), 71,102(1), 97(1), 101(1) 20,49(1), 42(0), 79(1), 55(1), 0() 1() 125(1) 119(1), 100(0), 103(1), 16(0), 121(1) 71,108(0) 97(1), 93(1) 59(1), 95(2), 111(1), () () 6(2), 4(1), 3(2), 60(1), 60,1() 71,4() 45(0), 42(1), 37(1), 17(0), 16(0), 1() 116(2) 113(2), * * ** ** * 51,76(0), 75(1), 23(0), * * 111(1), * þ ** :23(1), : 21,68(0), 42(1), 96(0), 121(1) 113(1), 95(2), () 31,112(1) 23(1), 8(0), : ** þ 12(1), : 70(1), Bernissartia þ þ ** : * * () 01,3() 67(1), 31(1), 20(1), 3(1), : ** () 22(1), 6(0), ** 51,94(2), 75(1), * 50,56(1), 45(0), Theriosuchus 51,83(0), 75(1), Comahuesuchus Comahuesuchus ** þ þ () 18(1), 3(0), () 71,95(2), 67(1), 6(0), : 5(1), 70(1), : : 1() 124(1) 111(0), * * þ ** () 22(0), 1(1), : 0() 105(1), 104(1), () 37(1), 9(0), : Simosuchus Simosuchus ** 51(0), () 51,3() 52,46(0), 45(2), 34(0), 15(1), 2(0), : 70(1), ** ** ** þ ** * * : ** Peirosauridae: 97(1), 114(1) 1() 117(1) 115(0), 108(0), () 5(1), 3(2), * 67(3), 90,27(1), 19(0), ** Goniopholis : ** 31,36(0), 23(1), 91,69(1), 19(1), ** 83(0), ** 74(1), ** 90,120(1) 99(0), : : 11(1), ** 104(1) * ** ** 51,22(1), 15(1), * 94(1), * * * 74(1), ** ** 4(1), : 74(1), 01,71(4), 60(1), 74(1), 45(0), ** 70(1) 23(1), * ** ** * 0() 120(1) 101(0), * 50,71(3), 65(0), 115(0) : 92(1), 75(1), ** þ 100(0), 91,66(1), 59(1), 31,2() 45(2), 28(1), 23(1), 11(1), : ** * ** þ 31,69(1) 13(1), : 112(1), 67(2), Malawisuchus ** ** ** * ** 21,116(1) 42(1), 77(0), * 10(0), * Notosuchus * ** * * 120(1) 10(0), 31(1), 94(1), 96(1), ** 65(0), * * 90(1), * 70,104(1), 77(0), * 90(1), * ** 83(0), 29(1), 51,17(0), 15(1), 5(1), 90,36(0), 29(0), 31,100(0) 93(1), 21,66(0), 42(1), * 101(1) ** 111(0) ** ** 75(0) ** * ** ** ** ** ** ** ** 114(1), 120(1), * * * * * 103(1), * * * * * 101(1) 92(1), 11(1), 31(1), 90(1), 99(1), 90(1), 51(0), 11(1), 95(0), 95(0), 67(3), 67(3), 55(1), 75(0), 42(1), 97(1), 11(1), : : c rsacai parietalis cranii crista frontalis cranii crista crest cotylar ccp ccf capitulum cc process capitate capt condyle calcaneal rostrum capp basisphenoid basisphenoid calc bsr antrum basioccipital bs basioccipital boa tympani bo bulla rib axial right b.tym rib axial left axr.r pleurocentrum axis axr.l spine neural axis axp arch neural axis axns ridge median axna axis intercentrum axis axmr rib atlantal right axint rib atlantal left atr.r arch neural atlas atr.l peg intercentrum astragalar atlas for atna facet trough-shaped atint condyle astragalar canal semicircular astf anterior astc surface articular surface of articular fossa asc of medial fossa lateral articular ar.mf maxilla ar.lf of process ascending ar ilium the of fenestra process antorbital apm anterior AOF angular femur of api ridge adductor an calcaneum of acetabulum ball adr anterior acet ab in used figures abbreviations Anatomical D: Appendix ** rtscu bonapartei Bretesuchus awhh pakistanensis Pabwehshi 123(1), Baurusuchus 116(0) * Baurusuchus Sebecus Iberosuchus Pabwehshi Bretesuchus Iberosuchus Sebecosuchia: Sebecus Baurusuchus * 19(1), 1() 118(1), 110(1), 115(0) New : ** * þ ** 127(1) () 9(1), 1(2), : 74(1), : 90,26(1), 19(0), Iberosuchus Araripesuchus ** þ : þ þ þ * 90,69(1), 19(0), Bretesuchus 3(0), Baurusuchus Pabwehshi Pabwehshi * ** * 2(0), 122(1) 127(1) ** : ** 42(1), þ 1(1), : ** * 3(2), 81,80(1), 78(1), : : Bretesuchus * 70(1), rmMadagascar from ** 3(0), 42(1), þ ** þ 2(0), ** Baurusuchus * 1() 118(0) 114(1), Pabwehshi ** 20,7() 107(1), 79(2), 42(0), 70(1) * () () 77(2), 5(1), 3(1), * * ** 74(1), 1() 120(1) 114(0), 73,109(1), 67(3), 73,104(1) 67(3), þ * 115(0) : ** þ 110(1), * 3(1), 359 Downloaded By: [Columbia University] At: 05:34 17 September 2007 360 e einEsaha opening Eustachian median median the meu for channel anterior fenestra mandibular mea antrum mastoid canal semicircular MF lateral laterosphenoid ma recess lacrimal lsc the of recess opening lacrimal anterior ls opening ligament Eustachian lr.ao lateral lateral external long for lr attachment centrum on depression leu lateral lell lacrimal ridge ldc jugal foramen jugal lac jugal jrd jf fenestra infratemporal j peduncle) cruris (posterior interosseous peduncle for ITF ischial attachment trochanter isped iliofibularis septum choanal ioc internal choanae internal ilft humerus ics ic bulbi levator & h ophthalmic for scapulocoracoid groove of fossa glenoid vagi foramen goV recess frontal gfs fv perilymphaticum foramen fr ovale foramen artery ophthalmic fply for foramen oralis intramandibular fov foramen fibula foa for facet fio metoticum foramen magnum fibf longus foramen digitorum flexor for fme attachment foramen fm carotid posterior condyle fibular fdl foramen carotid fcp anterior aerum foramen fc frontal fac exoccipital fa f meatus auditory external ex recess ectopterygoid eam ectopterygoid roller distal ectr ect crest deltopectoral dr dentary diapophysis dpc barb diap rib cervical canal d cranioquadrate crb foramen coracoid cq coronoid condyle occipital corf cor ceratohyal IV tarsal distal co for surface contact chl transiliens Cartilago femur for cdt4 on contact flange caudofemoralis cdfemf cct .H Turner H. A. utcintube Eustachian nerve trigeminal of branches srobi sinus rhomboid puboischiofemoralis rs for femur on rugosities radiale scars rf1-2 muscle radiale rds1-3 process retroarticular rd of rarp hemicondyle radial the for humerus concavity of hemicondyle radial radhc radius for facet radius radh quadratojugal radf quadrate ra qj phalanx pterygoid q of process anterior pterygoid px canal ptap semicircular posterior sinus parietal pt psc prezygapophysis ps prootic przg pillar prefrontal pro prfp prefrontal recess ilium the prefrontal of ilium prf.rec process of process posterior posterior the prf on flange ppi peduncle) (anterior peduncle pubic ppf postzygapophysis pped postorbital pozg foramen po premaxillary pmxf to (corresponding premaxilla on depression premaxilla pmxd palpebral pmx posterior palpebral pisiform p.plp tuberosity plp dorsal proximal fibula pi of ridge proximodistal fold pdt condylar proximal pdrf parapophysis pcf palatine parp parietal pal otoccipital pa orbit oto opisthotic orb duct nasopharyngeal spine neural opo canal npd neural arch ns neural nasal nc na maxilla the of process n palatal maxilla recess mxpp frontal median mx tibioastragalar mfr medial for attachment sulcus triangular medial fossa mtal triangular small median mts Eustachian median the mstf for channel posterior mep armlopening?) lacrimal ligament tube nens2 internus humerus Downloaded By: [Columbia University] At: 05:34 17 September 2007 A8720 UA Skulls sinus rhomboid recess t otic external rs fenestra supratemporal roe ribs sacral for STF scar ridges recess ss1-2 scapular squamosal squamosal sr1-3 sqr splenial sq lamina suprapostzygapophyseal spozgl fenestra suborbital supraoccipital sp recess SOF subglenoid ridge so supracondylar sgr crest supraacetabular surangular scr sac scars muscle sa radius rs1-2 MHP 2297 PR FMNH u tuberculum trochanter fourth with process associated transverse rugosities tub lamina thin tr1-2 posterior lamina thin tp anterior canal temporal tlp tla tc 4 rnoaitlstr obc focpt? 1.18 1.90 5.31 1.25 ? ventral to jugal supraoccipital of depth, extent dorsoventral ventral-most to postorbital orbits depth, of dorsoventral front in snout ? depth, occiput dorsoventral of back to suture suture frontoparietal frontoparietal to premaxilla of orbits tip to anterior just snout contact width, pmx-max at snout of width orbits quadrate b/w left frontal to of quadrate width right of edge lateral width, STF b/w width it fsultbeaogpseirmri fSF? jugals STF b/w of skull margin of posterior width along table skull of width rnoaitlstr obc focpt2.77 2.27 orbits of front in snout depth, occiput dorsoventral of back to suture frontoparietal eiltrlwdh nobtlfnsr 0.60 0.60 ? supra- of ? margin posterior along table skull of width fenestra antorbital width, mediolateral fenestra antorbital length, anteroposterior fenestra supratemporal width, mediolateral fenestra supratemporal length, anteroposterior i fpeail ofotprea suture frontoparietal to premaxilla of tip it fsota m–a otc ? 0.83 5.30 orbits to anterior just contact snout width, pmx–max at snout of width orbits quadrate b/w left frontal to of quadrate width right of edge lateral width, fenestra supra-temporal b/w width it fsulbwjugals b/w skull of width xeto ciia condyle occipital of extent eprlfenestra temporal orhtrochanter fourth 0.69 3.09 1.20 5.57 ? ? , , 4.58 6.1 2.6 * * * * t lmn.Almaueet ncentimeters. in measurements All element. gana from measurements Select measurements Skeletal E: Appendix nerve hypoglossal nerve glossopharyngeal XII nerve vestibulocochlear IX nerve facial nerve trigeminal VIII nerve optic VII V phalanx ungual ulnare II the for facet upx ulnare humerus of ulnf hemicondyle ulnar ulna for uln facet ulna ulh valve ear upper ulf for groove ul uef * * G Comments natrs eoe esrmn fapartial a of measurement denotes asterisk An . getrtrochanter” ”greater New Araripesuchus rrpscu tsangatsan- Araripesuchus rmMadagascar from 361 Downloaded By: [Columbia University] At: 05:34 17 September 2007 Skulls E Appendix MHP 2298 PR FMNH MHP 2299 PR FMNH A8775 UA 362 osvnrldph otria ovnrlms xeto ua 1.57 occipital of extent ventral jugal to supraoccipital of depth, extent dorsoventral ventral-most to postorbital depth, dorsoventral rnoaitlstr obc focpt? ? ? ? ? ? of extent ventral to jugal supraoccipital of depth, extent dorsoventral ventral-most to postorbital orbits depth, of dorsoventral front in snout ? depth, occiput dorsoventral of back to suture suture frontoparietal frontoparietal to premaxilla ? of orbits ip to ? anterior just snout contact width, pmx-max at snout of ? width orbits quadrate b/w left frontal to of quadrate width right of 1.46 edge lateral width, STF b/w width jugals STF b/w of 1.60 skull margin of posterior width along table skull of width fenestra antorbital width, mediolateral fenestra antorbital length, anteroposterior fenestra supratemporal width, mediolateral fenestra supratemporal length, anteroposterior rnoaitlstr obc focpt2.36 ? ? ? ? ? of extent ventral to jugal supraoccipital of depth, extent dorsoventral ventral-most to postorbital orbits depth, of dorsoventral front in snout ? depth, occiput dorsoventral of back to suture suture frontoparietal frontoparietal to premaxilla ? of orbits tip to ? anterior just snout contact width, pmx-max at snout of ? width orbits quadrate b/w left frontal to of quadrate width right of ? edge lateral width, STF b/w width jugals STF b/w of ? skull margin of posterior width along table skull of width fenestra antorbital width, mediolateral fenestra antorbital length, anteroposterior fenestra supratemporal width, mediolateral fenestra supratemporal length, anteroposterior rnoaitlstr obc focpt1.76 ? ? ? ? ? occipital of extent ventral jugal to supraoccipital of depth, extent dorsoventral ventral-most to postorbital orbits depth, of dorsoventral front in snout 3.82 depth, occiput dorsoventral of back to suture suture frontoparietal frontoparietal to premaxilla of orbits tip to anterior just snout contact width, pmx-max at snout of width orbits quadrate b/w left frontal to of quadrate width right of edge lateral width, STF b/w width jugals b/w skull of width eiltrlwdh nobtlfnsr ? ? 1.37 STF 1.39 of posterior margin along table skull of width fenestra antorbital width, mediolateral fenestra antorbital length, anteroposterior fenestra supratemporal width, mediolateral fenestra supratemporal length, anteroposterior eiltrlwdh nobtlfnsr ? ? 1.06 ? fenestra antorbital width, mediolateral fenestra antorbital length, anteroposterior fenestra supratemporal width, mediolateral fenestra supratemporal length, anteroposterior condyle ciia condyle occipital ciia condyle occipital condyle .H Turner H. A. (Continued) 0.70 ? 0.92 ? 0.78 ? 0.89 ? ? 1.74 ? 2.97 1.80 1.64 * Comments Downloaded By: [Columbia University] At: 05:34 17 September 2007 eiltrlwdhof: width mediolateral MHP 2299 PR FMNH MHP 2297 PR FMNH COLUMN VERTEBRAL A8767 UA SACRUM variation of range the exemplifying sample small a Therefore, column. preserved any noted. along be numbered exactly will be can none Vertebrae: Dorsal MHP 34 etscapula left 2334, PR FMNH MHP 2301 PR FMNH MHP 27 etscapula left 2297, PR FMNH MHP 27 etramus left 2297, PR FMNH ramus right 2297, PR FMNH MANDIBLE Skulls E Appendix HUDRGIRDLE SHOULDER egh i fbaet lni os 3.60 3.20 tip posterior to tip anterior fossa width, glenoid blade to blade of tip length, xscnrmatr-otro egh0.70 of: length anteroposterior length antero-posterior centrum axis xscnrmatr-otro egh0.45 ? ? of: length anteroposteriorl length antero-posterior centrum axis height dorsoventral axis width mediolateral intercentrum atlas length anteroposterior intercentrum atlas ta necnrmmdoaea it 0.65 0.46 height dorsoventral axis width mediolateral intercentrum atlas length anteroposterior intercentrum atlas armlength sacrum width mid-sacrum 2306) PR FMNH 2305, PR 2306) FMNH PR 8722–8727, FMNH (UA 2305, 0.40 PR from FMNH ranges 8722—8727, process (UA transverse 0.80 to to height (#3) 0.50 from ranges length egh i fbaet lni os 4.72 0.58 ? surface glenoid tip of length posterior anteroposterior to tip anterior fossa width, glenoid blade to blade of tip length, egho au aiu osvnrl9.72 2.09 mandibular of fenestra depth dorsoventral ramus of height dorsoventral maximum ramus of length eiltrlwdhof: width mediolateral armlength sacrum width mid-sacrum egh i fbaet lni os 2.58 0.35 ? surface glenoid tip of length posterior anteroposterior to tip anterior fossa width, glenoid blade to blade of tip length, A87,rgtscapula right 8779, UA Scapulae egho au aiu osvnrl9.30 ? mandibular of fenestra depth dorsoventral ramus of height dorsoventral maximum ramus of length c6 c5 c4 c3 c6 c5 c4 c3 c4 c3 c3 (Continued) , 0.28 0.56 0.52 0.86 1.09 1.71 0.62 0.61 0.71 0.74 0.79 0.79 0.86 0.75 ? 0.40 0.46 0.45 1.0 * * * * * * * * lni sbdydamaged badly is glenoid eytpo ld sbroken is blade of tip very neirms oto frmsdamaged ramus of portion anterior-most damaged ramus of anteriorportions and posterior ld shaiydmgdcnatsraefrtecoracoid the for surface contact damaged heavily is blade ifiutt esr eas fmatrix of because measure to difficult Comments sfragmentary is New Araripesuchus rmMadagascar from 363 Downloaded By: [Columbia University] At: 05:34 17 September 2007 MHP 32 ethumerus left 2302, PR FMNH MHP 28 ih humerus right 2298, PR FMNH 364 MHP 27 ih humerus right 2297, PR FMNH A82,rgthumerus right 8721, UA MANUS AND FORELIMB MHP 33 etcoracoid left 2313, PR FMNH MHP 27 ih coracoid right 2297, PR FMNH A84,rgtcoracoid right 8741, UA A84,lf coracoid left 8740, UA Coracoids MHP 33 etscapula left 2313, PR FMNH Skulls E Appendix egho etpcoa rs 1.04 0.63 ? ? crest deltopectoral of length condyles distal across width width proximal width midshaft length crest deltopectoral of length condyles distal across width egho etpcoa rs 1.84 ? crest deltopectoral of length condyles distal across width width proximal isatwidth midshaft rxmlwidth proximal width midshaft length egho etpcoa rs ? 0.75 crest deltopectoral of length condyles distal across width width proximal width midshaft Humeri length length surface glenoid length, expansion distal of width width midshaft rxmlwidth proximal length egh lni surface glenoid length, expansion distal of width width midshaft width proximal egh lni surface glenoid length, expansion distal of width width midshaft width proximal length length it fdsa expansion distal of width width midshaft egh lni surface glenoid length, rxmlwidth proximal egh i fbaet lni os 2.80 0.39 2.07 surface glenoid tip of length posterior anteroposterior to tip anterior fossa width, glenoid blade to blade of tip length, length neootro egho lni ufc ? surface glenoid of length anteroposterior .H Turner H. A. (Continued) 0.71 1.00 0.82 1.46 ? 0.94 0.83 0.31 ? ? 0.28 0.64 0.43 0.42 ? 0.30 1.09 3.88 0.31 1.27 ? 0.79 5.1 3.86 0.39 0.87 ? 0.23 3.07 ? 3.59 2.24 * * * * * * * * * * * * * itledmsig hf sfatrdadpoia n is end proximal and fractured is shaft missing, end distal itledmsig hf sfatrdadpoia end proximal and fractured is shaft missing, end distal opeewt ml eardfatr htsotn the shortens that fracture repaired small a with complete rxmlha smissing is head proximal h rxmlms oto ssihl damaged slightly is portion proximal-most the neirprino h ld sdamaged is blade the of portion anterior oto h hf smissing is shaft the of most h rxmlms oto ssihl aae si the is as damaged slightly is portion proximal-most the lgtydamaged slightly ssihl damaged slightly is measurement Comments itlexpansion distal Downloaded By: [Columbia University] At: 05:34 17 September 2007 MHP 30 etradiale left 2310, PR FMNH MHP 34 ih ulna right 2324, PR FMNH A83,lf radiale left 8737, UA Radialia MHP 37 ih ulna right 2327, PR FMNH ulna left 2326, PR FMNH MHP 28 etulna left 2298, PR FMNH Ulnae MHP 39 etradius left 2329, PR FMNH MHP 37 ih radius right 2327, PR FMNH MHP 36 etradius left 2326, PR FMNH MHP 34 ih radius right 2324, PR FMNH radius left 2298, PR FMNH Radii left and right 2326/2327, PR FMNH MHP 35 ih humerus right 2325, PR FMNH Skulls E Appendix isatwidth midshaft itlwidth distal width proximal width midshaft length itlwidth distal width proximal width midshaft length isatwidth midshaft length width distal width proximal width midshaft length radiale left 8736, UA isatwidth midshaft length isatwidth midshaft length length width midshaft length ulna right 8758, UA isatwidth midshaft length isatwidth midshaft length isatwidth midshaft length isatwidth midshaft length egho etpcoa rs ? ? 1.15 ? crest deltopectoral of length condyles distal across width width proximal width midshaft length humerus crest deltopectoral of length condyles distal across width width proximal width midshaft isatwidth midshaft length length (Continued) , 0.57 ? 0.44 0.44 0.19 0.44 0.16 ? 0.31 0.10 ? ? 0.20 1.83 3.97 1.56 1.36 0.16 ? 4.07 3.58 0.14 2.76 3.91 ? 0.21 ? 0.97 3.44 ? ? 0.30 2.92 ? ? ? 3.58 0.14 * * * * * * * rxmledsihl damaged slightly end proximal opeebtprin tl oee ymatrix by covered still portions but complete o fragmentary too complete hf aae nplaces in damaged shaft itledmsig hf atal edi matrix in held partially shaft missing, end distal erycmlt u rcue n oae nsprt slabs separate on located and fractured but complete nearly nytepoia w-hrsi preserved is two-thirds proximal the only rxmledmissing end proximal o fragmentary too rxmledmissing end proximal itledadcnye missing condyles and end distal o fragmentary too opeeecp ml oto ftedsa surface distal the of portion small a except complete Comments o fragmentary too New Araripesuchus rmMadagascar from 365 Downloaded By: [Columbia University] At: 05:34 17 September 2007 MHP 30 etischium left 2330, PR FMNH Ischia MHP 30 etilium left 2330, PR FMNH ilium right 2303, PR FMNH MHP 31 ih ilium right 2301, PR FMNH MHP 28 ih ilium right 2298, PR FMNH 366 MHP 27 etilium left 2297, PR FMNH Ilia HIP MHP 34 ih ulnare right 2324, PR FMNH MHP 32 ih phalanx right 2312, PR FMNH Phalanges MHP 38 ih CII MC right 2328, PR FMNH MHP 32 ih CII MC right 2312, PR FMNH Metacarpals A85,rgtulnare right 8759, UA A83,lf ulnare left 8736, UA Ulnaria MHP 34 ih radiale right 2324, PR FMNH Skulls E Appendix neootro egho ctblm0.7 0.4 acetabulum of length anteroposterior process posterior of height dorsoventral eghfo la rcse odsa i 1.6 tip distal to processes iliac from length neootro egho ctblm0.8 0.4 ? 0.7 acetabulum of length anteroposterior process posterior of height dorsoventral length anteroposterior acetabulum of length anteroposterior process posterior of height dorsoventral neootro length anteroposterior neootro egho ctblm0.8 0.5 acetabulum of length anteroposterior process posterior of height dorsoventral neootro length anteroposterior neootro egho ctblm? 0.5 acetabulum of length anteroposterior process posterior of height dorsoventral neootro length anteroposterior neootro length anteroposterior neootro egho ctblm0.8 0.4 acetabulum of length anteroposterior process posterior of height dorsoventral length anteroposterior ilium left 8768, UA width length isatwidth midshaft length width midshaft isatwidth midshaft length isatwidth midshaft length length itlwidth distal width proximal width midshaft length isatwidth midshaft length .H Turner H. A. (Continued) 1.6 1.5 2.0 1.6 2.1 1.6 0.10 0.19 0.15 0.45 0.19 0.16 0.95 1.17 0.81 0.37 ? 1.13 1.90 0.09 1.38 1.08 * * * * * * * * * * neir13o shu sdmgdado bcrdb matrix by obscured and/or damaged is ischium of 1/3 anterior isn neirprocess anterior missing neirhl fiicbaei aae einn halfway beginning damaged is blade iliac of half anterior opeebtprilyecsdi matrix in encased partially but complete h eydsa ato h otro rcs sdamaged, is process posterior the of part distal very the erycmlt u nymda surface medial only but complete nearly tl atal nmti itlwdhmyb overestimate be may width distal matrix in partially still rcue tmdhf;poia n damaged end proximal midshaft; at fractured hog uraeaua crest supraacetabular through swl stetpo h neirprocess anterior the of tip the as well as rxmladdsa nssihl damaged slightly ends distal and proximal sexposed is Comments Downloaded By: [Columbia University] At: 05:34 17 September 2007 MHP 28 ih tibia right 2298, PR FMNH Tibiae MHP 37 ih femur right 2337, PR FMNH MHP 30 ih femur right 2330, PR FMNH MHP 30 etfemur left 2300, PR FMNH MHP 28 ih femur right 2298, PR FMNH MHP 27 ih femur right 2297, PR FMNH A83,rgtfemur right 8734, UA Femora HINDLIMB MHP 27 pubis 2297, PR FMNH MHP 31 etpubis left 2331, PR FMNH MHP 30 ih pubis right 2330, PR FMNH pubis left 2330, PR FMNH pubis left 8778, UA A87,rgtpubis right 8777, UA A87,rgtpubis right 8776, UA Pubes Skulls E Appendix neootro width anteroposterior length width midshaft length isatwidth midshaft length isatwidth midshaft length width midshaft length isatwidth midshaft length isatwidth midshaft length isatwidth midshaft neck of width mediolateral neck of width mediolateral length proximodistal isatwidth midshaft neck of width mediolateral neck of width mediolateral length proximodistal isatwidth midshaft neck of width mediolateral neck of width mediolateral length proximodistal isatwidth midshaft neck of width mediolateral neck of width mediolateral length proximodistal width midshaft neck of width mediolateral neck of width mediolateral length proximodistal width midshaft neck of width mediolateral neck of width mediolateral length proximodistal isatwidth midshaft neck of width mediolateral eiltrlwdho neck of width mediolateral length proximodistal (Continued) ? 0.25 ? 3.76 2.90 ? 0.59 ? 8.82 4.59 0.49 0.51 0.32 0.78 0.32 ? ? 0.8 0.4 1.04 ? 0.70 0.25 ? 0.39 1.73 1.72 2.04 1.7 1.99 1.84 6.18 ? 2.69 2.11 0.13 0.16 ? 0.3 ? 0.11 0.16 * * * * * * * * * nypoia n spreserved is end proximal only o aae omeasure to damaged too eghi lgtyofdet rcuernigtruhtebone the through running fracture a to due off slightly is length hf scmrse n damaged and compressed is shaft erycmlt u ak eoa head femoral lacks but complete nearly nydsa n spreserved is end distal only hseeeti al rknadisiett sete right either as identity its and broken badly is element this opeebtdsa ld spol preserved poorly is blade distal but complete mle hn#1 lotcmlt u ak rxmlend proximal lacks but complete almost #91, than smaller itlbaefatrdadpoia tip proximal and fractured blade distal eyfamnay nytebaei preserved is blade the only fragmentary, very Comments rlf ol o edetermined be not could left or broken New Araripesuchus rmMadagascar from 367 Downloaded By: [Columbia University] At: 05:34 17 September 2007 MHP 30 ndnie MT unidentified 2330, PR FMNH MHP 39 ih T(II?) MT right 2319, PR FMNH MHP 39 TIV MT 2319, PR FMNH 368 MHP 27 etcalcaneum left 2297, PR FMNH Calcanea MHP 39 TIII MT 2319, PR FMNH MHP 30 ih astragalus right 2330, PR FMNH Astragali MHP 39 TII MT 2319, PR FMNH MHP 28 ih calcaneum right 2298, PR FMNH A83,uietfidMT unidentified 8736, UA Metatarsals MHP 35 ih fibula right 2335, PR FMNH MHP 30 etfibula left 2330, PR FMNH fibula right 2300, PR FMNH MHP 28 ih fibula right 2298, PR FMNH fibula left 8781, UA Fibulae MHP 37 ih tibia right 2337, PR FMNH MHP 35 ih tibia right 2335, PR FMNH MHP 30 ih tibia right 2330, PR FMNH MHP 30 ih tibia right 2300, PR FMNH Skulls E Appendix isatwidth midshaft length width midshaft length isatwidth midshaft length width midshaft it ftuber of width tuber of height width anteroposterior ball of height it fatclrsurface articular of width length width midshaft neootro it mdhf)0.13 surface articular of width astragalus left 2297, PR FMNH (midshaft) width anteroposterior it ftuber of width tuber of height length isatwidth midshaft neootro width anteroposterior ball of height length length neootro it mdhf)0.20 (midshaft) width anteroposterior neootro it mdhf)0.27 0.13 (midshaft) width anteroposterior length (midshaft) width anteroposterior length length neootro it mdhf)0.16 (midshaft) width anteroposterior length neootro width anteroposterior length width anteroposterior length neootro width anteroposterior length width anteroposterior length .H Turner H. A. (Continued) 0.61 0.92 0.48 0.28 0.76 ? 0.19 0.09 0.44 ? 0.41 0.50 ? 3.09 0.09 0.32 0.74 ? ? 1.13 0.22 1.27 ? 0.21 3.64 0.40 0.8 6.60 3.61 3.04 6.07 3.09 ? 3.92 7.24 * * * * * * * * * * o fragmentary too atal bcrdb matrix by obscured partially rxmledi missing is end proximal rxmledi missing is end proximal damaged rxmledi missing is end proximal ak rxmlend proximal lacks rxmledi missing is end proximal damaged damaged o aae omeasure to damaged too nypoia n spreserved is end proximal only Comments Downloaded By: [Columbia University] At: 05:34 17 September 2007 MHP 37 eodpaaxlnt 0.41 0.64 measured. be to matrix by obscured or fragmentary too are elements other All length phalanx second 2337, PR length FMNH phalanx proximal 2337, PR FMNH Phalanges Pedal MHP 38 etM (III?) MT left 2338, PR FMNH MHP 37 ndnie MT unidentified 2337, PR FMNH Skulls E Appendix isatwidth midshaft length isatwidth midshaft length (Continued) 0.11 ? 2.29 2.02 * rxmleddamaged end proximal Comments New Araripesuchus rmMadagascar from 369