Analysis of North American Goniopholidid Crocodyliforms in a Phylogenetic Context

Analysis of North American goniopholidid crocodyliforms in a phylogenetic context

Allen, Eric Randall https://iro.uiowa.edu/discovery/delivery/01IOWA_INST:ResearchRepository/12730561560002771?l#13730723240002771

Allen, E. R. (2012). Analysis of North American goniopholidid crocodyliforms in a phylogenetic context [University of Iowa]. https://doi.org/10.17077/etd.317zy27t

ANALYSIS OF NORTH AMERICAN GONIOPHOLIDID CROCODYLIFORMS IN A PHYLOGENETIC CONTEXT

Eric Randall Allen

A thesis submitted in partial fulfillment of the requirements for the Master of Science degree in Geoscience in the Graduate College of The University of Iowa

July 2012

Thesis Supervisor: Associate Professor Christopher Brochu

Graduate College The University of Iowa Iowa City, Iowa

CERTIFICATE OF APPROVAL ______

MASTER’S THESIS ______

This is to certify that the Master’s thesis of

Eric Randall Allen has been approved by the Examining Committee for the thesis requirement for the Master of Science degree in Geoscience at the July 2012 graduation.

Thesis Committee: ______Christopher Brochu, Thesis Supervisor

______Jonathan Adrain

______Llewellyn D. Densmore

To my wife

ACKNOWLEDGMENTS I wish to thank my advisor, C. Brochu, and my Comprehensive Exam committee, J. Adrain, A. Budd, J. Logsdon, and H. Sims. For discussion and help preparing this manuscript I would like to thank E. Allen, S. Salisbury, E. Wilberg, and the University of Iowa VertPaleo discussion group: S. Drumheller, M.E. Gold, A. Grass, J. McHugh, J. Miller-Camp, J. Nestler, and M. Spencer. For access to collections and specimens, I wish to thank J. Conrad and C. Mehling (American Museum of Natural History), A. Henrici (Carnegie Museum of Natural History), W. Langston Jr. (University of Texas at Austin), D. Main (University of Texas at Arlington), L. Steel (British Natural History Museum), and J. Person (Sam Noble Museum of Natural History). Finally, I wish to thank A. Pritchard for allowing access to unpublished data. This research was funded in part by the Max and Lorraine Littlefield Fund.

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TABLE OF CONTENTS

LIST OF TABLES vi

LIST OF FIGURES vii

LIST OF ABBREVIATIONS viii

CHAPTER

1 OSTEOLOGY OF AMPHICOTYLUS STOVALLI AND COMMENTS ON GONIOPHOLIDIDS OF THE MORRISON FORMATION 1

Introduction 1 Systematic Paleontology 2 Description of Material 3 Form and condition 3 Skull Openings 3 Cranial Elements 5 Discussion 10 Comparison to other taxa 10 Status of Morrison Formation goniopholidids 12

2 NEW MATERIAL OF THE MANDIBLE OF DENAZINOSUCHUS KIRTLANDICUS 18

Introduction 18 Systematic Paleontology 19 Description of Material 20 Observations from the holotype 20 Description of new material 20 Discussion 23

3 PHYLOGENETIC ANALYSIS OF GONIOPHOLIDIDAE 27

Introduction 27 Status of goniopholidid phylogenetics 28 Phylogenetic Analysis 29 Taxon sampling and dataset 29 Results 30 General results and North American goniopholidids 30 Denazinosuchus and Vectisuchus 32 Discussion 33 Biostratigraphy 34 Conclusions 35

APPENDIX A LIST OF CHARACTERS 42

APPENDIX B DATA MATRIX USED IN PHYLOGENETIC ANALYSIS 62

APPENDIX C CHARACTER TRANSITIONS 73

REFERENCES 79

LIST OF TABLES Table

1. Compilation of Morrison Formation goniopholidids 14

LIST OF FIGURES

Figure

1. Example of an extant crocodyliform 15

2. OMNH 2392 Amphicotylus stovalli, dorsal aspect 16

3. OMNH 2392 Amphicotylus stovalli, ventral aspect 17

4. PMU.R 232 Denazinosuchus kirtlandicus holotype 25

5. New material referred to Denazinosuchus kirtlandicus 26

6. Summary of phylogenetic placements of Goniopholididae 37

7. Phylogenetic results of entire dataset 38

8. Detail of cladograms investigating relationships within Goniopholididae 39

9. Detail of cladogram if Arlington Form excluded from analysis 40

10. Stratigraphically calibrated cladogram of selected taxa 41

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LIST OF ABBREVIATIONS Institutions: AMNH, American Museum of Natural History, New York, New York; BYU, Brigham Young University, Provo, Utah; CM, Carnegie Museum of Natural History, Pittsburgh, Pennsylvania; CMNH, Cleveland Museum of Natural History, Cleveland, Ohio; NMMNH, New Mexico Museum of Natural History and Science, Albuquerque, New Mexico;

OMNH, Sam Noble Museum of Natural History, Norman, Oklahoma; PMU.R, Paleontological Museum, University of Uppsala, Uppsala, Sweeden. TMM, Texas Natural Science Center, University of Texas at Austin; YPM, Yale Peabody Museum, New Haven, Connecticut. Anatomical Terms: Fr: frontal Jg: jugal La: lacrimal Mx: maxilla N: nasal Pa parietal Pal: palatine Po: postorbital Pf: prefrontal Pm: premaxilla Q: quadrate Qj: quadratojugal Sq: squamosal

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CHAPTER 1 OSTEOLOGY OF AMPHICOTYLUS STOVALLI AND COMMENTS ON GONIOPHOLIDIDS OF THE MORRISON FORMATION Introduction Crocodyliformes is a large, diverse group of archosaurs including extant crocodylians their fossil relatives. Though all living crocodyliforms are of relatively conserved morphology (Fig. 1), historically the clade was considerably more disparate, including even fully terrestrial and secondarily marine taxa. Goniopholididae constitutes some of the most common, widespread, and distinctive crocodyliforms of the Late

Jurassic and Early Cretaceous. These neosuchians are among the first crocodyliform taxa to superficially resemble modern crocodylians, and are presumed to have occupied a similar ecological role (Schwarz, 2002; Buffetaut, 1982). Goniopholidids were widespread throughout Laurasia during the Late Jurassic and Early Cretaceous, and are frequently found in freshwater or estuarine deposits (Hups et al., 2006). The majority of Goniopholis species are known from the United Kingdom, but also occur in continental Europe, particularly in the north and west (Owen 1842, 1849-1884, 1878, 1879; Hulke, 1878; Dollo, 1883; Koken 1883, 1886, 1887, 1896; Hooley,1907; Edinger, 1938; Jonet, 1981; Huckriede, 1982; von Oekentorp, 1984; Buffetaut, 1986; Norman, 1987; Buscalioni & Sanz, 1987a, b; Buffetaut et al., 1989; Cuny et al., 1991; Ortega et al., 1996; Salisbury et al., 1999; Salisbury, 2002; Schwarz, 2002; Schwarz-Wings et al., 2009; Salisbury & Naish, 2011). Goniopholidids have also been reported from Asia (Young, 1948; Buffetaut & Ingavat, 1980, 1983; Wu et al, 1996; Maisch et al., 2003) and North America (Marsh, 1877; Cope, 1878; Holland, 1905; Wiman, 1932; Mook, 1964, 1967; Tykoski et al., 2002). They are united in general by (1) nasals which do not participate in the narial border; (2) amphicoelous vertebrae; (3) two rows of rectangular, imbricated paravertebral osteoderms with rostrally-directed articular spine on the rostrolateral margin; (4) choanae bound caudally by the pterygoid and anteriorly by the

palatine; (5) the presence of a maxillary fossa on the caudolateral margin of the maxilla. This maxillary fossa is enigmatic, but it does lie in line with the line of neurovascular foramina on the lateral surface of the maxilla, and may thus represent a region of hyper- enlarged foramina (De Andrade, 2009). The Upper Jurassic Morrison Formation accounts for the most specimens and largest number of reported species in North America with five taxa currently recognized. Furthermore, all North American taxa attributed to Goniopholis Owen, 1841 are known from the Morrison (Table 1). This provides a unique opportunity to investigate Goniopholididae in North America and the affinity of Goniopholis identified on that continent and Europe. Amphicotylus stovalli is a comparatively well-represented Morrison Formation goniopholidid with two well-preserved skulls and considerable attributed postcrania. The original material for A. stovalli was excavated by the Works Progress Administration in the 1930s in Cimarron County, Oklahoma and initially investigated by J. W. Stovall, who noted overall similarity to A. gilmorei (Mook, 1964). The material was first described by Mook (1964) who ascribed it to Goniopholis Owen, 1841. Unfortunately, the initial, and preliminary, description lacked considerable detail. This study redescribes the type material of Amphicotylus stovalli and investigates Abbreviations—AMNH, American Museum of Natural History, New York, New York; BYU, Brigham Young University, Provo, Utah; CM, Carnegie Museum of Natural History, Pittsburgh, Pennsylvania; CMNH, Cleveland Museum of Natural History, Cleveland, Ohio; OMNH, Sam Noble Museum of Natural History, Norman, Oklahoma; TMM, Texas Natural Science Center, University of Texas at Austin; YPM, Yale Peabody Museum, New Haven, Connecticut.

Systematic Paleontology CROCODYLIFORMES Hay, 1930 MESOEUCROCODYLIA Whetstone and Whybrow, 1983

GONIOPHOLIDIDAE Cope, 1875 AMPHICOTYLUS Cope, 1878 AMPHICOTYLUS STOVALLI (Mook, 1964) (Figs. 2-3) Holotype—OMNH 2392; near-complete articulated skull. Paratype—OMNH 2322; near-complete articulated skull. Horizon and Locality—Jurassic (?Kimmeridgian), Morrison Formation, Locality V97 (Kenton 8), Cimarron County, Oklahoma. Revised Diagnosis—A goniopholidid crocodyliform with constricted “figure- eight” external naris, subtriangular lacrimal, frontal which participates in the orbit, and a narrow, channel-like internal choanae completely separating the pterygoids and bounded anteriorly by the maxilla, caudally by the pterygoid, and not extending caudal to the orbits. Description of Material Form and condition All observations are from OMNH 2392, a near-complete, articulated skull, with supplemental observations from OMNH 2322. Quality of preservation is very good on both specimens. OMNH 2392 is slightly crushed dorsoventrally, particularly the caudal portions, and slightly skewed to the left. The snout region is in very good condition with some slight damage along the lateral margins. The occipital condyle is broken off. In ventral aspect, the braincase caudal to the orbits is badly crushed, especially on the left. Skull openings The external nares are confluent along the midline of the skull. In dorsal aspect, the aperture is shaped like a broad “figure eight” due to midline protrusions of the premaxille into the narial border from both the rostral and caudal margins. The nares are directed dorsally and entirely contained within the premaxillae. Anterior and posteriolateral margins are slightly vaulted.

This posterior palatine vacuity is large and ovoid in shape, acute rostrally. The rostral and lateral margins are bound by the maxilla, and medially by the palatine. The opening extends rostrally to beneath the maxillary fossae. In dorsal aspect, the orbit is a subrectangular ellipse elongated rostrocaudally. Due to distortion of the skull, the left orbit appears triangular in shape. It is bound by the lacrimal , prefrontal, and frontal(?) medially, the postorbital caudally, and the jugal laterally. The supratemporal fenestra is nearly circular. The rostral and caudal margins are slightly elevated. The margins only slightly overhang the recessus supratemporales, particularly along the anterior margin. The foramen is bounded by the squamosal caudolaterally, the postorbital rostrolaterally, the frontal rostromedially, and the parietal caudomedially. The infratemporal fenestra is oblong, over twice as long as it is high. A broad processus infratemporalis of the quadratojugal, inserting approximately halfway along the occipital margin bifurcates that margin. The fenestra is bound by the jugal rostroventrally, the postorbital rostrodorsally, the squamosal dorsally, and the quadratojugal caudoventrally. The internal choanae form a deep, narrow channel 55mm long and 15mm wide along the midline of the palate, completely separating the palatines. The rostral margin is formed by the posterior margin of the maxillae and the caudal and caudolateral margins are formed by the pterygoids. A thin septum is partially preserved running the length of the opening. The posttemporal fenestra has been destroyed on both specimens due to dorsoventral crushing, though its position may still be noted. This foramen would be a very long but not very high slit-like opening bounded dorsally by the parietal and squamosal and ventrally by the supraoccipital and otoccipital ventrally. It is visible only is occipital aspect.

The medial Eustachian opening is a small, circular opening on the ventral surface of the braincase directly between the basioccipital and basisphenoid. It appears to have possibly been artificially widened during preparation. Cranial elements Premaxilla—In dorsal aspect, the premaxillae are as wide as the rostral portions of the maxillae. The rostral portion is spatulate in shape and the external nares are located 20mm from the rostral-most margin. They are convex laterally and indented caudally be a deep notch for reception of a large dentary pseudocanine. Caudal to this notch, they extend caudally medial to the macillae to contact the nasals about even with the lateral swelling of the maxilla over the maxillary pseudocanines. Dorsoventral crushing of the snout has slightly separated the premaxillae from the maxillae along this suture. The dorsal, medial, and lateral portions are sculptured with indistinct rugosities and shallow circular pits 2-3mm in diameter. The rostral margin is not sculptured, but is perforated by two 4mm wide holes which are inferred as wear-holes to accept the rostral-most dentary teeth. Ventrally, these pits lie in the rostral portion of the nasal vestibule and are 18mm apart. In vertral aspect, the caudal margin of the premaxillae forms a broad U-shape, convex rostrally, from the caudal notch to within 7mm of the caudal margin of the nasal vestibule. Five alveoli are present. A partially-erupted tooth crown is visible in the 4th alveolus on both the left and right elements. A small, 4mm long and 5mm wide triangular anterior palatine vacuity lies at the caudal midline, at the juncture with the midline of the maxillae. Two small 4mm wide pits lie along the premaxilla/maxilla suture lateral to this vacuity. These pits can be viewed in the occipital portion of the external nares. Their nature is unclear, as they are too medial to be considered tooth-reception pits. Maxilla—The Maxillae form the majority of the snout and secondary palate. The

lateral margin is convex over the 4th and 5th alveoli, becoming broadly concave caudally.

The dorsal and lateral surfaces are sculptured by indistinct pits and rugosities oriented roughly rostrocaudally. A shallow, oblong depression on the caudolateral margin marks the mandibular fossa. The rostral margin of the depression is marked by a broad, low protuberance on the dorsolateral margin of the snout measuring 25mm long, 15mm wide, and raising ~5mm from the surrounding snout. Neither the depression of the fossa nor the associated protuberance are sculptured, in contrast with the surrounding bone. The interior of the fossa is not smooth, but interior detail has been destroyed. In ventral aspect, the maxillae contact medially to form the majority of the secondary palate. 19 alveoli are preserved on the tooth row, with seven in-situ tooth crowns on the right and eight on the left. The suture with the palatines originates at the rostral apex of the posterior palatine vacuity and is broadly infolded, terminating medially at the choanae. The maxillae form the rostral margin of the choanae. Laterally, the maxilla contacts the jugal caudal to the posterior palatine vacuity and is overlain dorsally by the jugal. Nasal—The margins of the nasals are subparallel between the maxillae, swelling laterally to a maximum width even with the rostral margin of the maxillary fossa. At the caudal margin, the nasal contacts the frontal and lies lateral to the frontal, wedging between the frontal and prefrontal. The dorsal surface is sculptured by indistinct rugosities and pits 2-4mm in diameter which obscure the caudal margins and sutures with the lacrimal, prefrontal, and frontal. The midline suture between the nasals is straight and forms a distinct groove along the sagittal plane of the snout. Rostrally, the nasals wedge medially between the caudal processes of the premaxillae and slightly overlap laterally the occipital-most point of the premaxillae by <8mm, resulting in a sharp W-shape. This differs from Goniopholis simus (Salisbury et al., 1999) and G. baryglyphaeus (Schwarz, 2002). This arrangement can be seen on OMNH 2322 and is more easily seen on that specimen.

Lacrimal—in dorsal aspect, the lacrimal is subtriangular with the acute apex directed rostrally wedging between the maxilla and nasal. The medial contact with the nasal is complex with V-shaped rostrocaudally directed interfingerings. The lacrimals extend rostrally to even with the rostral margin of the maxillary fossa. The element reaches its widest point approximately 1/3 the distance from the caudal margin of the maxillary fossa and narrows caudally to the contact with the orbit. The lacrimal forms part of the rostral margin of the orbit. The dorsal surface is sculptured by fairly distinct shallow circular to subcircular pits ~3mm in size. The jugal is contacted laterally along a suture extending rostrally from the rostrolateral apex of the orbit. Prefrontal—The prefrontal is narrow, 45mm long and 22mm wide. It is roughly triangular in shape, with the acute apex oriented rostrally between the maxilla and lacrimal. The caudal margin forms part of the rostral margin of the orbit. The dorsal surface is sculptured by indistinct shallow rugosities. The exact contact with the nasal is mostly obscured by sculpturing. Frontal—In dorsal aspect, the frontals are spanned by a transverse interorbital crest. This crest forms a distinct “step” neatly separating the snout and skull table. Rostral of the crest, sculpturing is by indistinct rostrocaudal directed rugosities and caudal by distinct shallow circular pits to 5mm in size. Rostrolateral contact with the prefrontal is largely obscured by sculpturing. Laterally, the frontal forms part of the dorsomedial margin of the orbit and contacts the rostromedial process of the postorbital. The caudolateral margin forms the rostromedial margin of the supratemporal fenestra which is slightly raised. Caudally, the frontal narrows considerably medial to the supratemporal fenestra to contact with the parietal. The caudal process features a deep groove corresponding to the midline suture flanked by the raised medial margins of the supratemporal fenestrae. Dorsolaterally within the fenestra and in ventral aspect the frontal is unsculptured.

Postorbital—In dorsal aspect, the rostrolateral process of the postorbital forms the dorsocaudal margin of the orbit and the rostral margin of the infratemporal fenestra. This process is small, smooth, and rounded and extends ventrolaterally to contact the postorbital process of the jugal. The rostromedial process separating the orbit from the supratemporal fenestra is sculptured on the dorsal surface by shallow distinct rounded pits to 5mm in diameter and contacts the frontals medially at the rostral apex of the supratemporal fenestra in an interfingered suture. The caudal process is contiguous with the rostromedial process and contacts the squamosal caudally, slightly inserting into the squamosal in an offset V-shape. The ventromedial margin is obscured by damage to the skull, but is presumed to contact the laterosphenoid medially. There is a distinct groove along the ventral surface of the rostromedial process. Parietal—The parietal forms the caudomedial margin of the supratemporal fenestra, and this margin is slightly raised. The parietal contacts the caudal process of the frontal rostrally and the squamosal laterally. Sculpturing on the dorsal surface is consistent with surrounding elements. The occipital margin is rough and slightly concave with slight lateocaudal protuberances overhanging the posttemporal fenestrae. Squamosal—The occipital margin of the squamosal is concave, as is contact with the parietal. The rostrally-directed postorbital process contacts the postorbital, broadly overlapping it laterally and slightly medially. The squamosal is roughly subtriangular in dorsal aspect and heavily sculptured as the rest of the skull table. The dorsal surface of the caudolateral process has a distinct, heavily sculptured, protuberance 40mm long and 20mm wide and elliptical in shape. A deep furrow separates it from the rest of the squamosal. Supraoccipital—The supraoccipital is only visible in caudal aspect and appears as a ventrally-directed triangle inclined rostroventrally. It is bound forsally by the parietal and laterally by the otoccipital. It consists of a 5mm wide dorsoventrally directed medial ridge flanked laterally by deep 7mm wide depressions.

Otoccipital—The otoccipital is visible primarily in caudal aspect and does not extend laterally past the lateral margin of the squamosal. The paraoccipital process forms a broad, ovoid plate directed caudolaterally, expanding laterally and inclined rostroventrally. The occipital surface is slightly concave. In ventral aspect, the otoccipital appears to lie adjacent to the quadrate at the lateral end, although a deep groove separates the ventral margin from the rest of the skull. Laterally, the paraoccipital process broadly contacts the squamosal dorsally. Basioccipital and Basisphenoid—The occipital condyle is formed entirely by the basioccipital. In caudal aspect, the basioccipital is bounded dorsally by the otoccipitals and is oriented dorsoventrally. Ventral to the basioccipital, the basisphenoid expands laterally and features a median ridge flanked by shallow depressions. The caudolateral margin is extended to form a dorsoventrally directed flange. Rostral to this flange, the braincase is severly smashed in both OMNH 2392 and OMNH 2322. Exoccipital—In ventral aspect, the exoccipital is exposed as a rectangular flange 45mm long, 25mm wide, and extending caudolaterally away from the basisphenoid. It contacts the quadrate rostrolaterally and the otoccipital caudally. The caudal margin forms a distinct ridge. Jugal—The jugal is long and narrow, forming the ventrolateral margin of the infratemporal fenestra and orbit. It rostrally contacts the maxilla in a broad lap joint from just rostral of the postorbital bar to the maxillary fossa. Caudally, it contacts the quadratojugal in a lap joint at the caudal-most apex of the infratemporal fenestra. The postorbital process is narrow, originating on the ventromedial surface and bowed slightly medially and dorsally to contact the jugal process of the postorbital in a butt joint to form the postorbital bar. In dorsal aspect, the dorsal and lateral surfaces are sculptured by distinct shallow pits to 5mm across, identical to the sculpturing of the skull table. In ventral aspect, the jugal is not sculptured.

Quadratojugal—In dorsal aspect, the quadratojugal is divided into two parts: a lateral rugose surface sculptured with distinct pits consistent with surrounding elements and a medial unsculptured surface. A rostrally-directed process penetrates the caudal margin of the infratemporal fenestra just ventral to contact with the squamosal. The quadratougal is overlain mediodorsally by the squamosal and contacts the jugal rostrally in a butt joint running mediolaterally originating from the caudal tip of the infratemporal fenestra. In ventral aspect, the quadratojugal is smooth and subrectangular in outline. Quadrate—The quadrate is unsculptured and broad laterally. It is overlapped by the squamosal dorsomedially and the quadratojugal rostrolaterally. The caudomedial margin curves ventrally, becoming more sharply downturned caudally so that the medial process of the mandibular condyle is directed dorsolaterally at nearly 45o. The ventral surface exhibits a midline ridge along the long axis. Palatine—The rostral portion of the palatine is flat and triangular, with the rostrolateral portion inserting into the maxilla. The palatine contacts the maxilla rostrally from the rostral-most apex of the posterior palatine vacuity medially to the choanae. The caudal portion of the palatine becomes very narrow, forming a slender cylinder along the lateral margin. The caudal margin contacts the pterygoid. Pterygoid—The pterygoids are not well preserved. A 20mm portion of the right element at the contact with the palatine is present. It is thicker medially, expanding into a broad plate facing rostrally and slightly medially and inclined caudolaterally. This plate lies lateral to the braincase. Discussion Comparison to other taxa Mook (1964) noted similarity of Amphicotylus stovalli to “Goniiopholis” felix and particularly A gilmorei in overall shape if not specific detail. The external naris of A. stovalli is wider than it is long, a characteristic shared with A. gilmorei and A. lucasii. Also, the premaxillas of these taxa are broad and deep while the distance anterior to the

premaxillary notch is relatively short (Mook, 1964). However, the naris of A. stovalli incorporates a rostrally-directed projection of the premaxilla on the caudal margin resulting in a constricted figure-eight naris unique to that taxon. The lacrimal of A. stovalli is also subtriangular in dorsal aspect, as opposed to the broader rostral region and more blunted termination observed in other goniopholidids. Common to all currently known Morrison Formation goniopholidids is a very distinct termination of the nasal passages. In all except “G.” Felix—for which the palate is not known—the choanae form a long, slender channel completely bisecting the palatines. This channel is bound rostrally by the maxilla and caudally by the pterygoid. It is bisected along its length by a narrow septum consisting of the vomer (Pritchard, 2011). In Eutretauranosuchus delfsi, A. lucasii, and A. gilmorei, the choanae is exceedingly long at >1/3 the total length of the skull and constricted by medial expansion of the palatines lending an hourglass shape. This constriction is as extreme in some specimens as to almost separate the channel in two. Indeed, YPM 517, the E. delfsi holotype, shows such extreme constriction the anterior portion of the choanae was initially incorrectly construed as a separate opening (Mook, 1967; Smith et al., 2010). This morphology is also observed in Calsoyasuchus valliceps of the Lower Jurassic Kayenta Formation (Tykoski et al., 2002). However, not all North American taxa show this condition as evidenced by an undescribed skull from the Cretaceous of Arkansas (TMM 41881-1, pers. obs.; Langston, pers. com. 2007). It should be noted that though a rostrocaudally bisected choanae is not found in North American forms, the morphology is reported in other goniopholidids. Buffetaut and Ingavat (1980) report anterior expressions of the nasal passages in Sunosuchus maioi from the Upper Jurassic Hokou series of China. However, their reconstruction involves anterior openings of the choanae entirely contained within the maxilla and extensive contact of the palatines rostrally. This contrasts with the condition originally

reconstructed for Eutretauranosuchus where the anterior openings are bound caudally by the palatines and thus the palatines do not contact rostrally. The choanae of A. stovalli differ significantly from that of other Morrison Formation goniopholidids. Whereas in A. gilmorei, A. lucasii, and E. delfsi the choanae extend far into the pterygoid so that the caudal margin is well behind the orbits and very near the back of the skull, in A. stovalli it terminates approximately even to the rostral margin of the orbits and the pterygoids meet broadly caudal to this margin. As the caudal regions of both A. stovalli specimens are damaged and largely missing, an alternative morphology cannot be ruled out. It is possible the preserved choanae represents only the rostral portion and the palatines bisect the nasal passages in the manner similar to that originally reconstructed for Eutretauranosuchus by Mook (1967) and Sunosuchus maioi (Buffetaut & Ingavat, 1980), and the “true” choana is located farther caudal. However, this reconstruction does differ significantly in that in A. stovalli, the palatines are entirely separated and the constriction occurs farther caudal than in Eutretauranosuchus or S. maioi. In either scenario, this morphology is unique to A. stovalli, and is not observed in other crocodyliforms. Status of Morrison Formation goniopholidids North American goniopholidids have long been recognized as Goniopholis. However, the possibility these taxa are generically distinct from their European counterparts has been suggested (Clark, 1986; Salisbury et al., 1999; Lauprasert et al., 2007; Allen, 2007, 2010). Morrison Formation goniopholidids are united by their distinctive palate morphology, with the choanae entirely separating the palatines. By contrast, in European Goniopholis the choana lies between the palatine and pterygoid and the former element meets broadly anterior to the opening in a classic “mesosuchian” condition. Additionally, Morrison forms are united by a subtriangular prefrontal in dorsal aspect which extend rostrally farther than the lacrimal and excludes the lacrimal from contact with the maxilla, while in European forms the lacrimal extends farther rostrally

than the prefrontal and contacts the maxilla medially. The transverse intraorbital crest, though often present, is generally less pronounced in North American forms—though this may be variable. Therefore, Goniopholis, under current understanding, should only refer to those European forms. North American “Goniopholis” can be considered a distinct taxonomic unit from European Goniopholis, and those taxa should be considered constituents of a single genus to which Amphicotylus Cope, 1878 would apply, of which Amphicotylus lucasii Cope, 1878 is type (Allen, 2010; 2011; contra Smith et al., 2010). It should be noted that though Diplosaurus Marsh , 1877 technically holds precedent, the incomplete nature of YPM 986, the only known specimen, prevents more precise attribution pending further discoveries. Whereas A. stovalli is unique among Amphicotylus species, A. lucasii and A. gilmorei are exceedingly similar as to be nearly indistinguishable in detail. Cursory investigation indicates that while the rostrum of A. lucasii grades smoothly into the infraorbital region, A. gilmorei shows a marked lateral expansion of the skull at the level of the orbits. However, a more in-depth analysis of CM 2312—the only known specimen of A. gilmorei—reveals that this expansion is largely artificial due to breakage and lateral splaying of the jugal, particularly on the right side of the skull. It is thus likely that A. gilmorei can be considered a junior synonym of A. lucasii (Allen, 2010). This reduces the number of taxa known from the Morrison Formation to four: “G.” felix, E. delfsi, A. stovalli, and A. lucasii.

Referred Specimens OMNH 2322

BYU 17628; AMNH FR 570 FR AMNH 17628; BYU AMNH 5782; AMNH 652 AMNH 5782; AMNH

City, CO

OK Co., arron

Cim Cañon Cañon City, CO Locality Cañon City, CO WY Mtns., Freezeout

skull postcrania skull Material roof skull dorsal skull

Holotype OMNH 2392 AMNH 5766 AMNH CMNH 8028 CMNH YPM 986 CM 2312 CM

Mook, 1967 Mook,

(Holland, 1905) (Holland,

(Mook, 1964) (Mook, (Cope, 1878) (Cope,

(Marsh, 1877) (Marsh,

felix ” Compilation Formation Morrison goniopholidids of

Goniopholis “ Amphicotylus stovalli Amphicotylus lucasii Amphicotylus Eutretauranosuchus delfsi Eutretauranosuchus Table 1. Taxon Amphicotylus gilmorei Amphicotylus

Figure 1. Example of an extant crocodyliform. Alligator mississippiensis Photo by the author.

Figure 2. OMNH 2392 Amphicotylus stovalli, dorsal aspect photograph (top) and line interpretation (bottom). Known sutures and elements indicated on left side in red. Note the constricted “figure-eight” and lacrimal that is excluded from contact with the nasal. Position of maxillary fossa on lateral surface indicated by arrow.

Figure 3. OMNH 2392 Amphicotylus stovalli, ventral aspect photograph (top) and line interpretation (bottom). Palate elements indicated in red. Note the long, channel-like choana bound rostrally by the maxilla indicated by the arrow.

CHAPTER 2 NEW MATERIAL OF THE MANDIBLE OF DENAZINOSUCHUS KIRTLANDICUS

Introduction The Upper Cretaceous Fruitland and Kirtland Formations in the San Juan Basin, New Mexico have yielded a diverse vertebrate assemblage including turtles, lizards, dinosaurs, and mammals (Lucas, 1981; Hunt & Lucas, 1992). Crocodyliforms from the San Juan Basin are less well known, with four species reported: the alligatoroids Leidyosuchus sp. and Brachychampsa montana based off of isolated postranial elements and a nearly-complete skull respectively, the large alligotoroid Deinosuchus, and the enigmatic but more common Denazinosuchus kirtlandicus (Gilmore, 1916; Reeside, 1924; Powell, 1972; 1973; Kues et al., 1977; Wolberg & LeMone, 1980; Lucas, 1992; Sullivan & Lucas, 2003; Lucas et al., 2006). Wiman (1932) originally described D. kirtlandicus as a new species of Goniopholis Owen, 1841 based on its general morphology, including overall skull shape, dermal sculpturing, lacrimal separating the maxilla from the prefrontal, presence of a skull table, and rectangular paravertebral osteoderms with an anterior-directed articulating spine. Despite these being largely plesiomorphic characteristics common in crocodyliforms, subsequent workers found little reason to question this assignment (e.g. Wolberg, 1980; Mateer, 1980; Buffetaut, 1982; Lucas, 1992). Lucas and Sullivan (2003) reassessed the holotype skull and erected it as a new genus distinguished primarily by the lack of a definite maxillary fossa on the lateral surface of the snout and enlarged supratemporal fenestrae. However, Lucas and Sullivan (2003) did note considerable similarity to goniopholidids, in particular Amphicotylus lucasii, and as such concluded that Denazinosuchus is closely related to Goniopholis. Although Denazinosuchus is the most common and distinctive crocodyliform in the San Juan Basin (Lucas et al., 2006), material is highly fragmentary. In 2001, elements

of a partial but articulated left mandible (NMMNH P-33828) were recovered in association with some fragmentary cranial elements within the Late Cretaceous (Campanian) De-Na-Zin Member of the Kirtland Formation. A nearly complete splenial is present in articulation with a nearly complete angular and in association with a fragmentary surangular, palatine, and nasal. The new material described here provides new insights into the morphology of Denazinosuchus and allows for further investigation of its placement among crocodyliformes. Abbreviations—AMNH, American Museum of Natural History, New York, New York; NMMNH, New Mexico Museum of Natural History and Science, Albuquerque, New Mexico; PMU.R, Paleontological Museum, University of Uppsala, Uppsala, Sweeden.

Systematic Paleontology CROCODYLIFORMES Hay, 1930 MESOEUCROCODYLIA Whetstone and Whybrow, 1983 DENAZINOSUCHUS Lucas and Sullivan, 2003 DENAZINOSUCHUS KIRTLANDICUS (Wiman, 1932) (Figs. 4-5) Holotype—PMU.R 232; partial articulated skull (Fig. 1). Horizon and Locality—Late Cretaceous (Campanian), De-Na-Zin Member, Kirtland Formation, San Juan County, New Mexico (Wolberg, 1980; Hunt et al., 1992). Referred Material—NMMNH P-33828; articulated partial left mandibular ramus and associated partial right nasal and palatine. Horizon and Locality—Late Cretaceous (Campanian), De-Na-Zin Member, Kirtland Formation, Locality L-4714, South Mesa, San Juan County, New Mexico. Revised Diagnosis—A mesoeucrocodylian crocodyliform distinguished by supratemporal fenestrae that are much larger than the orbits, extensive participation by the splenial in a short mandibular symphasis, a ventral expansion of the splenial at the

symphasis, a rectangular caudal process of the palatine ventrally overlying the pterygoid, and a small neurovascular groove on the lateral surface of the maxilla.

Description of Materail Observations from the holotype Reinvestigation of NMMNH C-2438, a high-quality cast of the holotype skull, broadly conforms to the reconstruction of Lucas and Sullivan (2003). An isolated element interpreted by Lucas and Sullivan (2003) as a fragmentary parietal and articulated fragment of squamosal bears special mention, however. Examination of this element reveals that the lateral curvatures identified as the mediocaudal margins of the supratemporal fenestrae are of drastically different curvature, with the left side being of considerably smaller radius resulting in asymmetry unexpected in a midline element and necessitating reconstructed supratemporal fenestrae of radically different size. Furthermore, the anterior surface of the reconstructed squamosal fragment is confluent with the lateral margin of the parietal and shows no sign of breakage as implied by Lucas and Sullivan (2003) to account for the difference in curvature. Therefore, we reject the interpretation of this element as a parietal with attached squamosal fragment and reaffirm Wiman’s (1932) original interpretation of the fragment as the posterior corner of the right squamosal with a small fragment of the posterior process of the postorbital. The lateral surface of the snout shows a series of small neurovascular pits arranged linearly above the toothrow. Whereas the maxillary depression found in Goniopholis and related taxa (Buffetaut, 1982) is indeed not present (Lucas & Sullivan, 2003), at the same position along the snout, the neurovascular foramina are confluent forming a short, slit-like groove. Description of new material New material accessioned under NMMNH P-33828 consists of a partial left mandibular ramus in two portions, anterior and posterior with associated cranial elements. Quality of preservation is very good with no appreciable distortion or crushing.

This material represents the first known mandible from Denazinosuchus and expands on the known palate. Nasal—A partial right nasal is preserved consisting of the caudal portion at the widest point of the element with the anterior process and caudal-most process broken away. The mediocaudal margin shows the suture with the frontal consisting of a series of longitudinal rugosities and overlain slightly by the frontal. The lateral margin exposes a tongue-and-groove articulation with the maxilla with a long, rectangular longitudinal ridge approximately half the thickness of the nasal running the length of the preserved suture. The dorsal surface of the nasal is sculptured while the ventral surface is smooth and concave longitudinally. Palatine—The partial right palatine preserves the posterior end and is broken rostrally and caudomedially. The dorsal surface is very rugose while the ventral surface is smooth and has a shallow, broad longitudinal depression just lateral to the medial suture with the left palatine. Caudally, the palatine abuts and passes beneath the pterygoids. The posterior process at the contact is very broad and blunt, lending a near rectangular profile to the contact. The rostrolateral margin shows abutment with the very posterior-most projection of the maxilla, which terminates less than 1/5 the length along the preserved lateral margin. The remaining caudal half of the margin is smooth, delineating the medial boundary of the palatine vacuity. This participation of the palatine in the medial margin of the palatine vacuity differs from previous reconstructions of the palate (Wiman, 1932; Lucas & Sullivan, 2003), and also notes the first direct observation of the palatine in this taxon. The medial and caudal margin do not appear to preserve the margins of the choanae, but the mediocaudal portion of the element is severely broken. As such, the position of the choanae cannot be confidently reconstructed at this time. Angular—The angular is long and dorsoventrally very broad, rounded anteroventrally, and flattened caudoventrally. The dorsal margin of the medial surface is smooth outlining the vental margin of an extensive interior mandibular fenestra.

Extending caudally, the medial surface almost inverts to face ventrolaterally as it approaches the articular. The caudal margin is broken off just rostral to contact with the articular. The lateral surface is heavily sculptured with shallow circular pits in the same manner as the cranium of the holotype. Ventral and medial surfaces are smooth. The angular overlaps the splenial laterally and ventrally and the coronoid laterally, though the latter element is not preserved. The dorsal margin is badly broken, obliterating contact with the surangular. The rostral ~1/3 of the preserved length of the rostrodorsal margin is largely intact and is smooth both dorsally and laterally forming a smooth lamina where the element was overlain laterally by the dentary. A damaged apparently smooth margin immediately caudal to this contact may represent the ventral margin of an external mandibular fenestra or may show contact with the surangular. If indeed indicative of an external mandibular fenestra, its size and extent cannot be determined with current material. Splenial—The splenial is preserved in two pieces, rostral and caudal. The caudal fragment is in articulation with the angular which overlays it laterally while the rostral fragment is in isolation. The splenial is long and slender and sinusoidal in dorsal aspect. The caudal process is narrow and rectangular in cross-section while the rostral fragment widens considerably along its length. The lateral surface is smooth while the dorsal surface preserves a worn groove along the length immediately ventral to the expected position of the toothrow, though no distinct alveoli can be determined. The rostral-most tip is absent, but the rostral end does expand dramatically ventrally at the level of the symphasis. The splenial participates extensively in the mandibular symphasis. The anteromedial surface preserves a radial series of rugosities forming an articular surface ~45mm long as preserved. Surangular—An isolated fragment identified as a partial right surangular was found in association with the mandible. The fragment is small, but shows a near

rectangular cross-section and a very narrow (~40mm) dorsoventral height. The ventral margin shows a rugose contact surface.

Discussion NMMNH P-33828 is tentatively referred to Denazinosuchus kirtlandicus pending further discoveries based on general form and suture pattern, expected size, bone sculpturing, and the overlap of associated elements with the holotype, specifically the nasal. Comparison to the same element of the holotype implies a reconstructed skull size for NMMNH P-33828 comparable to that of the holotype PMU.R 232. Denazinosuchus has long been associated with Goniopholididae (Wiman, 1932; Kälin, 1955; Wolberg, 1980; Mateer, 1980; Buffetaut, 1982; Lucas, 1992; Williamson, 2000; Lucas & Sullivan, 2003). In general form and antorbital suture pattern it does indeed resemble Amphicotylus and Goniopholis. However, it lacks the distinctive maxillary fossa on the lateral margin of the snout characteristic of goniopholidids. It also lacks an interorbital transverse crest on the frontal as observed in Goniopholis (Salisbury et al. 1999; De Andrade & Hornung, 2011) and to a lesser extent in Amphicotylus stovalli (Allen, 2007), and A. lucasii (pers. obs.). Indeed, those characteristics used to unite Denazinosuchus with Goniopholis are not unique to that clade and those characteristics by which it differs are largely congruent a basal mesoeucrocodylian from the Cenomanian Woodbine Formation of Texas (Allen et al., 2011), herein termed the “Arlington Form”. Both taxa share, in addition to general form, “goniopholidid-style” paravertebral osteoderms with a pitted dorsal surface and anteriorly-directed articular spine; enlarged supratemporal fenestrae which are larger than the orbits; nasals which do not contact the external naris; a T-shaped frontal; blunt, nearly rectangular caudal process of the palatine overlapping ventrally the pterygoid; and extensive participation by the splenial in a short mandibular symphasis. However, Denazinosuchus possesses a broader distal end to the splenial and lacks the enlarged paired pseudocanines found in the “Arlington Form”.

The potential lack of an external mandibular fenestra is of interest. Absence of this opening has been considered characteristic of goniopholidids (Clark, 1994; Foster, 2006), and indeed is absent in Anetophthalmosuchus (Salisbury & Naish, 2011) and Amphicotylus stovalli (pers. obs.). However, external mandibular fenestrae have been reported in other goniopholidid taxa including Goniopholis simus (Salisbury et al., 1999; Salisbury, 2002), Eutretauranosuchus delfsi (Foster, 2006; Smith et al. 2010), and ?Amphicotylus lucasii ( AMNH DVP 652, pers. obs.). As such, the presence or absence of this opening cannot be considered characteristic of Goniopholididae. Reinvestigation of Denazinosuchus in light of this new material and a more recent understanding of crocodyliform phylogeny forces a reassessment of its classification. Denazinosuchus is not a Cretaceous North American goniopholidid as has been long assumed, but instead represents a more plesiomorphic and ambiguous position amongst neosuchians.

Figure 4. PMU.R 232 Denazinosuchus kirtlandicus holotype. Skull in dorsal aspect with line reconstruction. Plate IV of Wiman (1932).

Denazinosuchus kirtlandicus

(A) partial right nasal, dorsal aspect. dorsal nasal, right partial (A) aspect. lateral angular, articulated and splenial mandible; left (D) partial (B) partial right palatine, ventral aspect. ventral palatine, right partial (B) (C) partial surangular, lateral aspect; Rostral

Figure 5.New material referred to

CHAPTER 3 PHYLOGENETIC ANALYSIS OF GONIOPHOLIDIDAE Introduction Crocodyliform systematics has been highly influenced by the work of Benton and Clark (1988) and especially Clark (1994) whose character set has formed the basis of nearly every other crocodyliform study (e.g. Wu et al., 1994a; Wu et al., 1994b; Gomani, 1997; Wu et al., 1997; 2001; Buckley & Brochu, 1999; Buckley et al., 2000; Clark et al., 2000; Larsson & Gado, 2000; Pol et al., 2001; Sereno et al., 2001; Brochu et al., 2002; Clark & Sues, 2002; Ji et al., 2002; Pol, 2002; 2003; Tykoski et al., 2002; Rogers, 2003; Pol et al; 2004; Pol & Norell, 2004a; b; Turner & Calvo, 2005; Gasparini et al., 2006; Turner et al., 2005; Turner & Buckley, 2008; Smith et al., 2010; but see Ortega et al., 2000). However, understanding of the interrelationships within Crocodyliformes is in a constant state of flux. Phylogeny reconstruction in Crocodyliformes is highly dependent on taxon sampling, especially of basal mesoeucrocodylians and neosuchians (Turner & Buckley, 2008). Goniopholididae was established as a family by Cope (1875). Steel (1973) included the family in Mesosuchia and ascribed the genera Baharijodon, Coelosuchus, Dakotasuchus, Doratodon, Eutretauranosuchus, Goniopholis, Itasuchus, Microsuchus, Oweniaschus, Pertosuchus, Pinacosuchus, Pliogonodon, Polydectes, Shamosaurus, and Symptosuchus to it. It has subsequently been relegated to a “wastebasket taxon” status, though recent work has served to clarify certain taxonomy, particularly among European forms (e.g. Salisbury et al., 1999; Salisbury, 2002; Schwarz, 2002). Goniopholidid fossils are distributed throughout Western Europe, North America, and eastern Asia—a distinctly Laurasian distribution. Historically, some material has been identified from Gondwannan localities, but this material is either non-diagnostic or subsequent work has proposed a different relationship than Goniopholididae (e.g. Buffetaut, 1985; 1988; Carvalho et al., 2004).

Sereno (2005) suggested a branch-based definition for the group coinciding with the system established for TaxonSearch (Sereno et al., 2005) of the most inclusive clade containing Goniopholis crassidens Owen, 1841, but not Pholidosaurus geoffroyi (Owen, 1884), Alligarotellus beaumonti Gervais, 1871, Peirosaurus torminni Price, 1955, Araripesuchus gomesii Price, 1959, Notosuchus terrestris Woodward, 1896, or Crocodylus niloticus (Laurenti, 1768). Though technically in general use, this definition has not been formally put forward outside of TaxonSearch. For the purposes of this analysis, I affirm Sereno’s (2005) definition as a working phylogenetic definition of Goniopholididae with the modification of Goniopholis simus Owen, 1878 as the internal specifier. The reason for this change is threefold: 1) Goniopholis crassidens is known almost exclusively from postcrania making it unsuitable for inclusion in a data matrix consisting predominately of cranial characters, 2) material of Goniopholis simus is comparatively more abundant making it one of the most well- represented goniopholidid taxa, and 3) Goniopholis simus may be a junior synonym of G. crassidens (Salisbury et al., 1999; Salisbury, 2002; Salisbury & Naish, 2011). Status of goniopholidid phylogenetics Goniopholididae has appeared in several phylogenetic analyses, though typically as only a single exemplar taxon. Though Goniopholididae has been subjected to phylogenetic analysis in recent years (e.g. Buckley et al., 2000; Tykoski et al., 2002), these have been limited in scope. As such, no coordinated attempt to revise its taxonomy has been made, and attempts have focused primarily on Europe and have not been phylogenetic in nature (e.g. Salisbury et al., 1999). In most previous work, Goniopholididae includes Goniopholis, Eutretauranosuchus, Sunosuchus, and Calsoyasuchus, though mostly not by phylogenetic testing (Mook, 1967; Wu et al., 1996; Salisbury et al. 1999; Schwarz, 2002; Tykoski et al., 2002). In the majority of phylogenetic analyses to include goniopholidids, they are resolved as sister to Bernissartia + Eusuchia to the exclusion of pholidosaurs,

thalattosuchians, and dyrosaurs (e.g. Tykoski et al, 2002; Smith et al., 2010). Though less common in the literature, alternate topologies have been proposed (Fig. 6) for Goniopholididae more closely related to pholidosaurs and thalattosuchians than Eusuchia (e.g. Sereno et al., 2001; Jouve et al., 2006). One analysis, Jouve et al. (2006), recovered Goniopholididae as paraphyletic with Eutretauranosuchus, Sunosuchus, and Calsoyasuchus forming a clade and Vectisuchus and Goniopholis as successive sister taxa to Pholidosauridae + Thalattosuchia. This topology was not well supported and was dependent on the inclusion of Vectisuchus. Jouve et al. (2006) concluded this result may be due to the effect of convergence in long, tube-snouted (“longirostrine”) anatomy in Vectisuchus, dyrosaurs, and thalattosuchians.

Phylogenetic Analysis The comprehensive description of G. stovalli as well as new observations of other Morrison Formation goniopholidids (Smith et al., 2010; Pritchard, 2011) and the description of new specimens from Europe (Salisbury & Naish, 2011) allow for testing of the phylogenetic relationships of Goniopholididae. As Denazinosuchus and Vectisuchus have often been attributed to the goniopholidids (Wiman, 1932; Buffetaut & Hutt, 1980; Mateer, 1980; Wolberg, 1980; Buffetaut, 1982; Lucas, 1992; Lucas & Sullivan, 2003; Jouve et al., 2006; Salisbury & Naish, 2011), this also provides an opportunity to test the phylogenetic relationships of those taxa. Taxon sampling and dataset This analysis is based on modification of previous datasets (Clark, 1994; Wu & Sues, 1996; Pol, 1999; Ortega et al., 2000; Pol & Norell, 2004a; Gasparini et al., 2006; Turner et al., 2005; Allen, 2007; 2010), and particularly that of Turner and Buckley (2008). Modifications include character definitions and scorings. One new character was added to further account for variation among goniopholidids describing the morphology of the infraorbital crest. Due to the potential affinity of Goniopholididae with Pholidosauridae and Thalattosuchia, taxon sampling was increased for these clades.

Goniopholidid taxa were selected based on the presence of good quality, diagnostic cranial material. Putative taxa not possessing these (e.g. Goniopholis hartti, known only from teeth) were not considered. This includes a full sampling of all currently recognized goniopholidids of the Morrison Formation, Calsoyasuchus valliceps from the Lower Jurassic Kayenta Formation, two Asian, and four European goniopholidids. This is the largest and most inclusive sampling of goniopholidids to date. The total dataset is based on 64 taxa and 216 characters. Taxa were coded from the literature with the exception of the Morrison Formation goniopholidids excluding Eutretauranosuchus, G. simus, Denazinosuchus, and an undescribed specimen from the Cretaceous Woodbine Formation of Texas—the Arlington Form—which were coded from direct observation. Vectisuchus was included and recoded to test the affinity of Goniopholididae and Pholidosauridae following Jouve et al. (2006). Previously, only two European taxa, Goniopholis simus and G. baryglyphaeus, have been considered phylogenetically. Recently, two new taxa, Goniopholis willetti and Anetophthalmosuchus hooleyi, have been described from the Lower Cretaceous Wealden Supergroup of southern England (Salisbury & Naish, 2011). Though they avoid statements of relationships, Salisbury and Naish (2011) noted considerably similarity with G. simus, G. baryglyphaeus, and G. gracilidens. Remains of these two taxa include well-preserved and mostly complete cranial material allowing them to be incorporated into a phylogenetic matrix for the first time.

Results General results and North American goniopholidids The dataset was analyzed using TNT v. 1.1 (Goloboff et al., 2003). A heuristic tree search strategy was implemented using random addition sequences followed by Tree Bisection and Reconnection (TBR) branch swapping holding 10 trees per replicate. Character states were unordered. Zero-length branches were collapsed if they lack support in any most parsimonious reconstruction (i.e. Rule 1 of Coddington & Scharff,

1994). Nodal support is expressed by decay indices (Bremer, 1994) and statistical support calculated by jackknife resampling (Farris et al., 1996). This results in 280 most parsimonious trees (MPTs) of length 782 (CI = 0.355, RI = 0.681). All most parsimonious trees (MPTs) recovered in this analysis (Fig. 7) resolve an overall topology broadly consistent with previous analyses, though it does differ in the placement of Atoposauridae basal to Eusuchia + Pholidosauridae + Thalattosuchia as opposed to sister to Eusuchia + Goniopholididae (e.g. Turner, 2004; Turner & Buckley, 2008). Goniopholididae is monophyletic and sister to Bernissartia + Eusuchia and consists of Goniopholis, Sunosuchus, Siamosuchus, Anetophthalmosuchus, Eutretauranosuchus, and Amphicotylus as well as “G.” felix. North American forms resolve as monophyletic to the exclusion of all other goniopholidids. The North American clade is defined by their distinctive palate morphology with a long, channel-like choanae entirely separating the palatines (character 29.1) and the Morrison Formation clade further by triangular prefrontals which extend rostrally beyond the lacrimals and completely exclude the lacrimals from contact with the nasals (characters 6.1; 7.2). Within the North American group, the Lower Jurassic Calsoyasuchus is the basal-most member. Unfortunately, outside of this North American clade, goniopholidids resolve as a basal polytomy. Investigation reveals this is almost exclusively due to the influence of Goniopholis willetti. The derived long, narrow rostrum of this taxon renders it highly labile, falling everywhere from the basal-most member of the group to sister to the North American clade. Removal results in 39 most parsimonious trees of length 777 (CI = 0.358, RI = 0.684) and increased precision among non-North American goniopholidids. Under these conditions (Fig. 8a), the Lower Cretaceous Anetophthalmosuchus is resolved as sister taxon to the Upper Jurassic Goniopholis baryglyphaeus with which it shares extensive and homogenous sculpturing of rostral dermal elements (Salisbury & Naish, 2011). Additionally, the two Asian taxa, Sunosuchus junggarensis and

Siamosuchus, resolve as sister taxa despite the latter’s incompleteness based predominantly on their reduced infraorbital crest morphology restricted to the frontals. Finally, Goniopholis simus is sister taxon to all other goniopholidids. Alternatively, ambiguity in non-North American goniopholidids is a result of the highly homoplastic nature of the infraorbital crest. Exclusion of this character (character 216) results in 220 trees of length 780 (CI = 0.355, RI = 0.681). Goniopholis willetti is still labile in this analysis, but is more restricted, falling either basal to all other goniopholidids, sister to G. simus, or sister to the North American clade, or sister to a Siamosuchus + Sunosuchus + Anetophthalmosuchus + G. baryglyphaeus clade (Fig. 8b). Denazinosuchus and Vectisuchus The Upper Cretaceous North American taxon Denazinosuchus kirtlandicus does not resolve as a goniopholidid. It is found as sister taxon to the undescribed Arlington Form in a small, weakly supported clade with the enigmatic Vectisuchus united by the shape of the postorbital (character 24.1) and slight evagination around the orbits (character 188.1). This group is sister to Pholidosauria + Thalattosuchia, and this relationship is supported by a flattened postorbital bar (character 21.0), elongated retroarticular process (character 55.3), and extensive involvement of the splenial in the mandibular symphasis (character 61.2). The position of Denazinosuchus basal to pholidosaurs is dependent on the inclusion of the Arlington Form. Removal of this taxon results in Denazinosuchus placed within Pholidosauridae, alternatively sister to Oceanosuchus or Oceanosuchus + Terminonaris (Fig. 9). Topology aside from this change is unaffected. Consistent with the relationship found by Jouve et al. (2006), Vectisuchus falls outside of Pholidosauria + Thalattosuchia. Exclusion of Denazinosuchus or the Arlington Form does not affect this result. However, if the Arlington Form and Denazinosuchus are both removed from the analysis, Vectisuchus and Pholidosaurus collapse to a polytomy at the base of Pholidosauridae + Thalattosuchia.

Discussion North American goniopholidids—North American “Goniopholis” cannot be ascribed to Goniopholis sensu stricto. This phylogenetic analysis corroborates anatomical observation that Goniopholis should only refer to European forms and North American forms should be ascribed to Amphicotylus Cope, 1878. The missing palate of “G.” felix renders it labile within the Morrison Formation clade, so the status of Diplosaurus Marsh, 1877 cannot be determined at this time. In this dataset, Amphicotylus lucasii and A. gilmorei are almost identical in coding, differing only where a character is obscured in one or the other due to non-preservation (e.g., A. gilmorei does not preserve a lower jaw or postcrania). As expected, these two are resolved as sister taxa, lending further support to the conclusion A. gilmorei is a junior synonym of A. lucasii. The monophyly of Morrison Formation goniopholidids is of interest. Amphicotylus and Eutretauranosuchus could represent an endemic radiation of goniopholidids in the American West. The persistence of the incomplete palate, shared with Calsoyasuchus, while contemporary taxa elsewhere (e.g. G. baryglyphaeus) posess a more complete palate may represent geographic isolation of these taxa during the Jurassic. Though the incomplete palate can be considered derived in this dataset, it is possible this condition should represent the basal condition in goniopholididae, as it is present in the basal-most member. New data and new discoveries, particularly of basal Early Jurassic goniopholidids, are anxiously awaited. Denazinosuchus—This analysis reveals a somewhat ambiguous taxonomic relationship for Denazinosuchus. However it cannot be considered a constituent of Goniopholididae. Instead, it is more closely associated with Phlidosauridae, though how closely depends on the inclusion of the Arlington Form. Current material referred to the Arlington Form is fragmentary, but the similarity between these two taxa is striking. It is interesting to note that, though aligned with longirostirine taxa, neither of these two forms deviate strongly from a classic strongly triangular rostrum. Furthermore, it is

possible Denazinosuchus, and the Arlington Form represent the vanguard of a heretofore unidentified clade of neosuchians basal to pholidosaurs and thalattosuchians. This new clade would likely be distinguished by enlarged supratemporal fenestra on a broad skull table and extensive involvement of the splenial in the mandibular symphasis regardless of symphasis length. New discoveries and a more thorough sampling of basal neosuchians may help illuminate these issues. A potentially lucrative avenue of investigation would be crocodyliforms from the Cretaceous of Texas, for example Woodbinesuchus from the Woodbine Formation (Lee, 1997). Vectisuchus—The weakly tubular-snouted Vectisuchus is not a goniopholidid under this analysis, as is corroborated by the apparent lack of a maxillary fossa. Contrary to the results of Jouve et al. (2006), the inclusion of Vectisuchus in this analysis does not affect Goniopholididae resolving as closer to Pholidosauridae + Thalattosuchia than Bernissartia + Eusuchia. In fact, that topology is recovered only in a minority of MPTs with no unambiguous support only if Vectisuchus is excluded from the analysis but Denazinosuchus and the Arlington Form are retained. In all results, Goniopholididae is monophyletic, and in no result is Vectisuchus in any other topology than basal to Dyrosauridae + Thalattosuchia. Jouve et al. (2006) suggest Vectisuchus’s affinity with the pholidosaurs, dyrosaurs, and thalattosuchians is due to convergent evolution of a long, tubular rostrum in these taxa. However, Denazinosuchus and the Arlington Form, with which Vectisuchus is closely related in this analysis, are by no means longirostrine, and in the absence of the Arlington Form, Denazinosuchus falls out with Sarcosuchus and Oceanosuchus, two pholidosaurs notable for possessing a less tubular rostrum. This implies that certain characters assumed to be related to longirostry (e.g. extensive splenial involvement in the symphasis) may not be entirely correlated with rostrum morphology. Biostratigraphy Calibrating the recovered phylogeny with stratigraphy reveals substantial unreported diversity in neosuchians (Fig. 10). The position of Calsoyasuchus as the

basal-most member of North American goniopholidids necessitates a ghost lineage for Amphicotylus + Eutretauranosuchus extending to the Early Jurassic. The earliest-reported goniopholidid from Europe is G. baryglyphaeus from the Upper Jurassic (Kimmeridgian) of Portugal (Schwarz, 2002), approximately contemporaneous with the Morrison Formation (Kowallis et al., 1998; Mateus, 2006). As Calsoyasuchus is nested within Goniopholididae, this necessitates the invocation of ghost lineages for European and Asian forms to the Early Jurassic as well. Furthermore, the potential topology of Goniopholis simus as sister taxon to all other goniopholidids indicates substantial missing diversity for that lineage. The position of Vectisuchus and Denazinosuchus as sister to the Pholidosauridae + Thalattosuchidae clade requires an extensive ghost lineage to the Early Jurassic, representing much unrepresented diversity in that lineage. The alternate topology for Denazinosuchus as nested within Pholidosauridae as recovered by exclusion of the Arlington Form from the dataset is potentially more stratigraphically congruent for that taxon, but as the Cretaceous Vectisuchus remains outside the clade, this does not alleviate. If Vectisuchus, Denazinosuchus, and the Arlington Form represent a previously unidentified major division of neosuchian phylogeny, attention should be focused on Early Cretaceous formations of North America and Europe. Further investigation and discoveries are required to resolve these taxonomic issues.

Conclusions Goniopholidid systematic taxonomy is far from resolved. The most inclusive phylogenetic dataset of goniopholidids to date reveals a nested North American clade characterized by a distinctive channel-like choana morphology. However, relationships among Goniopholididae as a whole, and particularly European and Asian taxa, remain ambiguous. North American goniopholidids of the Jurassic are well represented. By contrast, goniopholidids from the Cretaceous of North America are notably under- represented, with identified taxa either being found not to be goniopholidid at all (e.g.

Denazinosuchus kirtlandicus) or currently undescribed. Future work will necessarily focus on taxon sampling, particularly of basal neosuchians, to further elucidate taxonomy.

A B

Figure 6. Summary of phylogenetic placements of Goniopholididae. (A) monophyletic, closer to Eusuchia (modified from Tykoski et al., 2002) (B) monophyletic, closer to pholidosaurs (modified from Sereno et al., 2001) (C) paraphyletic, closer to pholidosaurs (modified from Jouve et al., 2006)

Figure 7. Phylogenetic results of entire dataset.Strict consensus of 280 most parsimonious trees of length 782 (CI = 0.355, RI = 0.681). Nodal support indicated by Jackknife (>50) and Bremer (>1) values. Character state changes at lettered nodes are tabulated in Appendix C.

Figure 8. Detail of cladograms investigating relationships within Goniopholididae. (A) If Goniopholis willetti is excluded. Consensus of 39 MPTs (Length = 777, CI = 0.358, RI = 0.684). (B) If infraorbital crest is excluded. Consensus of 220 MPTs (Length = 778, CI = 0.355, RI = 0.681)

Figure 9. Detail of cladogram if Arlington Form excluded from analysis. Consensus of 223 MPTs (Length = 772, CI = 0.360, RI = 0.687).

Figure 10. Stratigraphically calibrated cladogram of selected taxa. Goniopholididae is indicated in red.

APPENDIX A LIST OF CHARACTERS

1. (modified from Clark, 1994: char. 1): External surface of dorsal cranial bones: smooth (0), slightly grooved (1), or heavily ornamented with deep pits and grooves (2).

2. (modified from Clark, 1994: char. 2): Skull expansion at orbits: gradual (0), or abrupt (1).

3. (modified from Clark, 1994: char. 6): External nares facing anterolaterally or anteriorly (0), dorsally not separated by premaxillary bar from anterior edge of

rostrum (1), or dorsally separated by premaxillary bar (2).

4. (Clark, 1994: char. 7): Palatal parts of premaxillae: do not meet posterior to incisive foramen (0), or meet posteriorly along contact with maxillae (1).

5. (modified from Clark, 1994: char. 10): posterior palatal branches of maxillae anterior to palatines: do not meet (0), or meet extensively but posterior most parts fail to meet (1), or meet entirely (2).

6. (Clark, 1994: char. 11): Nasal contacts lacrimal (0), or do not contact (1). 7. (Clark, 1994: char. 12): Lacrimal contacts nasal along medial edge only (0), or medial and anterior edges (1), or does not contact (2).

8. (Clark, 1994: char. 13): Nasal contribution to narial border: yes (0), or no (1). 9. (Clark, 1994: char. 14): Nasal-premaxilla contact: present (0), or absent (1). 10. (modified from Clark, 1994: char. 15): Descending process of prefrontal: does not contact palate (0), or contacts palate(1).

11. (Clark, 1994: char. 16): Postorbital-jugal contact: postorbital anterior to jugal (0), or postorbital medial to jugal (1), or postorbital lateral to jugal (2).

12. (Clark, 1994: char. 17): Anterior part of the jugal with respect to posterior part: as broad (0), or twice as broad (1).

13. (Clark, 1994: char. 18): Jugal bar beneath infratemporal fenestra: flattened (0), or rod-shaped (1).

14. (Clark, 1994: char. 19): Quadratojugal dorsal process: narrow, contacting only a small part of postorbital (0), or broad, extensively contacting the postorbital (1).

15. (Clark, 1994: char. 20): Frontal width between orbits: narrow, as broad as nasals (0), or broad, twice as broad as nasals (1).

16. (Clark, 1994: char. 21): Frontals: paired (0), or unpaired (1). 17. (Clark, 1994: char. 22): dorsal surface of frontal and parietal: flat – table like (0), or with midline ridge (1).

18. (modified from Clark, 1994: char. 23): Parieto-postorbital suture: absent from dorsal surface of skull roof and supratemporal fossa (0), absent from dorsal surface of skull roof but broadly present within supratemporal fossa (1), or present within supratemporal fossa and on dorsal surface of skull roof (2).

19. (Clark, 1994: char. 24): Supratemporal roof dorsal surface complex (0), or dorsally flat “skull table” developed with postorbital and squamosal with flat shelves extending laterally beyond quadrate contact (1).

20. (modified from Clark, 1994: char. 25): postorbital bar sculpted (if skull sculpted) (0), or unsculpted (1).

21. (modified from Clark, 1994: char. 26): Postorbital bar transversely flattened (0), or cylindrical (1).

22. (Clark, 1994: char. 27): Vascular opening in dorsal surface of postorbital bar: absent (0), or present (1).

23. (modified from Clark, 1994: char. 28): Postorbital anterolateral process: absent or poorly developed (0), or well developed, long, and acute (1).

24. (Clark, 1994: char. 29): Dorsal part of the postorbital: with anterior and lateral edges only (0), or with anterolaterally facing edge (1).

25. (Clark, 1994: char. 30): Dorsal end of the postorbital bar broadens dorsally, continuous with dorsal part of postorbital (0), or dorsal part of postorbital bar constricted, distinct from dorsal part of the postorbital (1).

26. (Clark, 1994: char. 31): Bar between orbit and supratemporal fossa broad and solid, with broadly sculpted dorsal surface (if sculpture present) (0), or bar narrow, sculpting restricted to anterior surface (1).

27. (Clark, 1994: char. 33): Parietal: with broad sculpted region separating fossae (0), or with sagital crest between supratemporal fossae (1).

28. (modified from Clark, 1994: char. 36): Posterolateral process of squamosal: poorly developed and projected horizontally at the same level of the skull (0), elongated, thin and posteriorly directed, not ventrally deflected (1), or elongated, posterolaterally directed, and ventrally deflected (2).

29. (Clark, 1994: char. 37): Palatines: do not meet on palate below the narial passage (0), form palatal shelves that do not meet (1), or meet ventrally to the narial passage, forming part of secondary palate (2).

30. (modified from Clark, 1994: char. 39): Choanal opening continuous with pterygoid ventral surface except for anterior and anterolateral borders (0), or opens into palate through a deep midline depression (choanal groove) (1).

31. (Clark, 1994: char. 40): Palatal surface of pterygoid smooth (0), or sculpted (1). 32. (Clark, 1994: char. 41): Pterygoids posterior to choanae: separated (0), or fused (1). 33. (modified from Clark, 1994: char. 43): Primary pterygoidean palate: forms posterior half of the choanal opening (0), or forms posterior, lateral and part of the anterior margin of the choana (1), or completely enclose choana (2).

34. (modified from Clark, 1994: char. 44): Anterior edge of choanae situated between the suborbital fenestra (or anteriorly) (0), situated near the posterior edge of suborbital fenestra (1), or near posterior edge of pterygoid flange (2).

35. (Clark, 1994: char. 45): Quadrate: without fenestrae (0), with single fenestra (1), or with three or more fenestrae on dorsal and posteromedial surfaces (2).

36. (Clark, 1994: char. 46): Posterior edge of quadrate: broad medial to tympanum, gently concave (0), or posterior edge narrow dorsal to occipital contact, strongly concave (1).

37. (Clark, 1994: char. 47): dorsal, primary head of quadrate articulates with: squamosal, otocciptial, and prootic (0), or with prootic and laterosphenoid (1).

38. (Clark, 1994: char. 48): Ventrolateral contact of otoccipital with quadrate very narrow (0), or broad (1).

39. (Clark, 1994: char. 49): Quadrate, squamosal and otoccipital: do not meet to enclose cranioquadrate passage (0), enclose passage near lateral edge of skull (1), or meet broadly lateral to the passage (2).

40. (Clark, 1994: char. 50): Pterygoid ramus of quadrate: with flat ventral edge (0), or with deep groove along ventral edge (1).

41. (modified from Clark, 1994: char. 55 by Ortega et al., 2000: char. 68): Basisphenoid ventral surface: shorter than the basioccipital (0), or wide and similar to, or longer in length than the basioccipital (1).

42. (Clark, 1994: char. 56): Basisphenoid: exposed on ventral surface of braincase (0), or virtually excluded from ventral surface by pterygoid and basioccipital (1).

43. (Clark, 1994: char. 57): Basioccipital: without well-developed bilateral tuberosities (0), or with large pendulous tubera (1).

44. (Clark, 1994: char. 58): Otoccipital: without laterally concave descending flange ventral to subcapsular process (0), or with flange (1).

45. (Clark, 1994: char. 59): Cranial nerves IX-XI: pass through common large foramen vagi in otoccipital (0), or cranial nerve IX passes medial to nerves X and XI in separate passage (1).

46. (Clark, 1994: char. 60): Otoccipital: without large ventrolateral part ventral to paroccipital pricess (0), or with large ventrolateral part (1).

47. (Clark, 1994: char. 63): Mastoid antrum: does not extend into supraoccipital (0), or extends through transverse canal in supraoccipital to connect middle ear regions (1).

48. (Clark, 1994: char. 64): Posterior surface of supraoccipital: nearly flat (0), or with bilateral posterior prominences (1).

49. (modified from Clark, 1994: char. 65): Palpebrals: absent (0), or one small palpebral present in orbit (1), or one large palpebral (2), or two large palpebrals (3).

50. (Clark, 1994: char. 66): External nares: divided by a septum (0), or confluent (1). 51. (modified from Clark, 1994: char. 67): Antorbital fenestra: as large as orbit (0), about half the diameter of the orbit (1), much smaller than the orbit (2), or absent (3), or with salt gland fossa (4).

52. (modified from Clark, 1994: char. 68 by Ortega et al., 2000: char. 41): Supratemporal fenestrae extension: relatively large, covering most of surface of skull roof (0), or relatively short, fenestrae surrounded by a flat and extended skull roof (1).

53. (modified from Clark, 1994: char. 69): Choanal groove: undivided (0), partially septated (1), or completely septated (2).

54. (Clark, 1994: char. 70): Dentary: extends posteriorly beneath mandibular fenestra (0), or does not extend beneath fenestra (1).

55. (modified from Clark, 1994: char. 71): Retroarticular process: absent or extremely reduced (0), very short, broad, and robust (1), with an extensive rounded, wide, and flat (or slightly concave) surface projected posteroventrally and facing dorsomedially (2), posteriorly elongated, triangular shaped and facing dorsally (3), or posteroventrally projecting and paddle shaped (4).

56. (Clark, 1994: char. 72): Prearticular: present (0), or absent (1).

57. (modified from Clark, 1994: char. 73): Articular without medial process (0), with short process not contacting braincase (1), or with process articulating with otoccipital and basisphenoid (2).

58. (Clark, 1994: char. 74): Dorsal edge of surangular: flat (0), or absent (1). 59. (Clark, 1994: char. 75): Mandibular fenestra: present (0), or absent (1). 60. (Clark, 1994: char. 76): Insertion area for M. pterygoideous posterior: does not extend onto lateral surface of angular (0), or extends onto lateral surface of angular (1).

61. (modified from Clark, 1994: char. 77): Splenial involvement in symphysis: not involved (0), involved slightly in symphysis (1), or extensively involved (2).

62. (Clark, 1994: char. 78): Posterior premaxillary teeth: similar in size to anterior teeth (0), or much longer (1).

63. (modified from Clark, 1994: char. 79): Maxillary teeth waves: absent, no tooth size variation (0), one wave of teeth enlarged (1), or enlarged maxillary teeth curved in two waves (festooned) (2).

64. (Clark, 1994: char. 80): Anterior dentary teeth opposite premaxilla-maxilla contact: no more than twice the length of other dentary teeth (0), or more than twice the length (1).

65. (modified from Clark, 1994: char. 81): Dentary teeth posterior to tooth opposite premaxilla-maxilla contact: equal in size (0), or enlarged dentary teeth opposite to smaller teeth in maxillary toothrow (1).

66. (modified from Clark, 1994: char. 82 by Ortega et al., 2000: char. 120): Anterior and posterior scapular edges: symmetrical in lateral view (0), anterior edge more strongly concave than posterior edge (1), or dorsally narrow with straight edges (2).

67. (modified from Clark, 1994: char. 83 by Ortega et al., 2000: char. 121): Coracoid length: up to two-thirds of the scapular length (0), or subequal in length to scapula (1).

68. (Clark, 1994: char. 84): Anterior process of ilium: similar in length to posterior process (0), or one-quarter or less of the length of the posterior process (1).

69. (Clark, 1994: char. 85): Pubis: rod-like, without expanded distal end (0), or with expanded distal end (1).

70. (Clark, 1994: char. 86): Pubis: forms anterior half of ventral edge of acetabulum (0), or pubis contacting the ilium, but partially excluded from the acetabulum by the anterior process of the ischium (1), or pubis completely excluded from the acetabulum by the anterior process of the ischium (2).

71. (Clark, 1994: char. 89): Atlas intercentrum: broader than long (0), or as long as broad (1).

72. (modified from Clark, 1994: char. 90): Cervical neural spines: all anteroposteriorly large (0), only posterior ones rod-like (1), or all spines rod-like (2).

73. (modified from Clark, 1994: char. 91 by Buscalioni and Sanz, 1988: char. 37 and by Brochu, 1997: char. 7): Hypapophyses in cervicodorsal vertebrae: absent (0), present only in cervical vertebrae (1), present in cervical and the first two dorsal vertebrae (2), present up to the third dorsal vertebra (3), or up to the fourth dorsal vertebra (4).

74. (Clark, 1994: char. 92): Cervical vertebrae: amphicoelous or amphyplatian (0), or procoelous (1).

75. (Clark, 1994: char. 93): Trunk vertebrae: amphicoelous or amphyplatian (0), or procoelous (1).

76. (Clark, 1994: char. 94): All caudal vertebrae: amphicoelous or amphyplatian (0), or first caudal biconvex with other procoelous (1), or procoelous (2).

77. (Clark, 1994: char. 95): Dorsal osteoderms: rounded or ovate (0), or rectangular, broader than long (1), or square (2) or absent (3).

78. (modified from Clark, 1994: char. 96 and Brochu, 1997a: char. 40): Dorsal osteoderms: without articular anterior process (0), with a discrete convexity on

anterior margin (1), or with a well-developed process located anterolaterally in dorsal parasagittal osteoderms (2).

79. (modified from Clark, 1994: char. 97 by Ortega et al., 2000: chars. 107 and 108): Rows of dorsal osteoderms: two parallel rows (0), more than two (1), or more than four with accessory ranges of osteoderms (sensu Frey, 1988) (2).

80. (Clark, 1994: char. 98): Osteoderms: some or all imbricated (0), or sutured to one another (1).

81. (Clark, 1994: char. 99): Tail osteoderms: dorsal only (0), or completely surrounded by osteoderms (1).

82. (Clark, 1994: char. 100): Trunk osteoderms: absent from ventral part of the trunk (0), or present (1).

83. (Clark, 1994: char. 101): Osteoderms: with longitudinal keels on dorsal surfaces (0), or without longitudinal keels (1).

84. (Wu and Sues, 1996: char. 14): Jugal: participating in margin of antorbital fossa (0), or separated from it (1).

85. (modified from Wu and Sues, 1996: char. 23): Articular facet for quadrate condyle: equal in length to quadrate condyles (0), slightly longer (1), or close to three times the length of the quadrate condyles (2).

86. (modified from Wu and Sues, 1996: char. 24 and Wu et al., 1997: char. 124): Jaw joint: placed at level with basioccipital condyle (0), below basioccipital condyle about above level of lower toothrow (1), or below level of toothrow (2).

87. (modified from Wu and Sues, 1996: char. 27 and Ortega et al., 2000: char. 133): Premaxillary teeth: five (0), four (1), three (2), or two (3).

88. modified from Wu and Sues, 1996: char. 29): Unsculptured region along alveolar margin on lateral surface of maxilla: absent (0), or present (1).

89. (Wu and Sues, 1996: char. 30): Maxilla: with eight + (0), seven (1), six (2), five (3), or four teeth (4).

90. (Wu and Sues, 1996: char. 40): Radiale and ulnare: short and massive (0), or elongate (1).

91. (modified form Gomani, 1997: char. 32): Basioccipital and ventral part of otoccipital: facing posteriorly (0), or posteroventrally (1).

92. (0Buscalioni and Sanz, 1988: char. 35): Vertebral centra: cylindrical (0), or spool shaped (1).

93. (modified from Buscalioni and Sanz, 1988: char. 39): Transverse process of posterior dorsal vertebrae dorsoventrally low and laminar (0), or dorsoventrally high (1).

94. (Buscalioni and Sanz, 1988: char. 49): Supra-acetabular crest: present (0), or absent (1).

95. (Buscalioni and Sanz, 1988: char. 54): Proximal end of radiale expanded symmetrically, similar to the distal end (0), or more expanded proximolaterally than proximomedially (1).

96. (Ortega et al., 1996: char. 5): Lateral surface of the anterior region of surangular and posterior region of dentary: without a longitudinal depression (0), or with a longitudinal depression (1).

97. (Ortega et al., 1996: char. 9): Ventral exposure of splenials: absent (0), or present (1).

98. (Ortega et al., 1996: char. 11, 2000: char. 100): Tooth margins: with denticulate carinae (0), or without carinae or with smooth or crenulated carinae (1).

99. (Pol, 1999a: char. 134): Jugal: does not exceed the anterior margin of orbit (0), or exceeds margin (1).

100. (Pol, 1999a: char. 135): Notch in premaxilla on lateral edge of external nares: absent (0), or present on the dorsal half of the external nares lateral margin (1).

101. (modified from Pol, 1999a: char. 136): Dorsal border of external nares: formed mostly by the nasals (0), or by both the nasals and premaxilla (1), or by premaxilla only (2).

102. (Pol, 1999a: char. 138): Posterodorsal process of the premaxilla: absent (0), or present extending posteriorly wedging between maxilla and nasals (1).

103. (Pol, 1999a: char. 139 and Ortega et al., 2000: char. 9): Premaxilla-maxilla suture in palatal view, medial to alveolar region: anteromedially directed (0), sinusoidal, posteromedially directed on its lateral half and anteromedially directed along its medial region (1), or posteromedially directed (2).

104. (Pol, 1999a: char. 140): Nasal lateral border posterior to external nares: laterally concave (0), or straight (1).

105. (Pol, 1999a: char. 141): Nasal lateral edges: nearly parallel (0), oblique to each other converging anteriorly (1), or oblique to each other, diverging anteriorly (2).

106. (Pol, 1999a: char. 144): Dorsoventral height of jugal antorbital region with respect to infraorbital region: equal or lower (0), or antorbital region more expanded than infraorbital region of jugal (1).

107. (Pol, 1999a: char. 145): Maxilla-lacrimal contact: partially included in antorbital fossa (0), or completely included (1).

108. (Pol, 1999a: char. 146): Lateral Eustachian tube openings: located posteriorly to the medial opening (0), or aligned anteroposteriorly and dorsoventrally (1).

109. (Pol, 1999a: char. 147): Anterior process of ectopterygoid developed (0), or absent (1).

110. (Pol, 1999a: char. 149 and Ortega et al., 2000: char. 13): Small foramen located in the premaxillo-maxillary suture in lateral surface (not for big mandibular teeth): absent (0), or present (1).

111. (Pol, 1999a: char. 150): Jugal posterior process: exceeding posteriorly past the infratemporal fenestrae (0), or not (1).

112. (Pol, 1999a: char. 151): Compressed crown of maxillary teeth: oriented parallel to the longitudinal axis of skull (0), or obliquely disposed (1).

113. (Pol, 1999a: char. 152): Large and aligned neurovascular foramina on lateral maxillary surface: absent (0), or present (1).

114. (modified from Pol, 1999a: char. 153): External surface of maxilla and premaxilla: with single plane facing laterally (0), or with ventral region facing laterally and dorsal region facing dorsolaterally (1).

115. (Pol, 1999a: char. 154 and Ortega et al., 2000: char. 104): Maxillary teeth: not compressed laterally (0), or compressed laterally (1).

116. (Pol, 1999a: char. 155): Posteroventral corner of quadratojugal: reaching the quadrate condyles (0), or not reaching quadrate condyles (1).

117. (modified from Pol, 1999a: char. 156): Base of postorbital process of jugal: directed posterodorsally (0), or dorsally (1), or anterodorsally (2).

118. (Pol, 1999a: char. 157): Postorbital process of jugal: anteriorly placed (0), in the middle (1), or posteriorly positioned (2).

119. (Pol, 1999a: char. 158 and Ortega et al., 2000: char. 36): Postorbital-ectopterygoid contact: present (0), or absent (1).

120. (Pol, 1999a: char. 163): Basisphenoid: without lateral exposure (0), or with lateral exposure on the braincase (1).

121. (Pol, 1999a: char. 165): Quadrate process of pterygoids: well developed (0), or poorly developed (1).

122. (modified from Pol, 1999a: char. 166 and Ortega et al., 2000: char. 44): Quadrate major axis directed: posteroventrally (0), ventrally (1), or anteroventrally (2).

123. (Pol, 1999a: char. 167): Quadrate distal end: with only one plane facing posteriorly (0), or with two distinct faces in posterior view, a posterior one and a medial one bearing the foramen aereum (1).

124. (Pol, 1999a: char. 168): Anteroposterior development of neural spine in axis: well developed covering the neural arch length (0), or poorly developed, located over the posterior half of the neural arch (1).

125. (Pol, 1999a: char. 169): Prezygapophyses of axis: not exceeding anterior edge of neural arch (0), or exceeding the anterior margin of neural arch (1).

126. (Pol, 1999a: char. 170): Postzygapophyses of axis: well developed, curved laterally (0), or poorly developed (1).

127. (Pol, 1999a: char. 213): Unsculpted region of the dentary below the tooth row: absent (0), or present (1).

128. (Buckley and Brochu, 1999: char. 102): Surangular forms only the lateral wall of the glenoid fossa (1).

129. (Buckley and Brochu, 1999: char. 103): Anterior margin of femur linear (0), or bears flange for coccygeofemoralis musculature (1).

130. (modified from Ortega et al., 1996: char. 1 and Buckley and Brochu, 1999: char. 107): Dorsal edge of dentary slightly concave or straight and subparallel to the longitudinal axis of skull (0), straight with an abrupt dorsal expansion, being straight posteriorly (1), with a single dorsal expansion and concave posterior to this (2), or sinusoidal, with two concave waves (3).

131. (modified from Ortega et al., 1996, char. 2 and Buckley and Brochu, 1999: char. 108): Dentary compression and lateroventral surface anterior to mandibular fenestra: compressed and vertical (0), or not compressed and convex (1).

132. (Ortega et al., 1996: char. 7 and Buckley and Brochu, 1999: char. 110): Splenial: thin posterior to symphysis (0), or splenial robust dorsally posterior to symphysis (1).

133. (Ortega et al., 1996: char. 13 and Buckley et al., 2000: char. 117): Cheek teeth: not constricted at base of crown (0), or constricted (1).

134. (Ortega et al., 2000: char. 10): Ventral edge of premaxilla located: at the same height as ventral edge of maxilla (0), or located on a dental groove (1).

135. (modified from Ortega et al., 2000: char. 19): Maxillary dental implantation: teeth in isolated alveoli (0), or located on a dental groove (1).

136. (Ortega et al., 2000: char. 24): Caudal tip of nasals: converge at sagittal plane (0), or caudally separated by anterior sagittal projection of frontals (1).

137. (Ortega et al., 2000: char. 33): Relative length between squamosal and postorbital: squamosal is longer (0), or postorbital is longer (1).

138. (modified from Ortega et al., 2000: char. 34): Jugal portion of postorbital bar: flushes with lateral surface of jugal (0), anteriorly continuous but posteriorly inset (1), or medially displaced and a ridge separates postorbital bar from lateral surface of jugal (2).

139. (Ortega et al., 2000: char. 42): Outer surface of squamosal laterodorsally oriented: extensive (0), or reduced and sculpted (1), or reduced and unsculpted (2).

140. (Ortega et al., 2000: char. 47): Quadratojugal spine at caudal margin of infratemporal fenestra: absent (0), or present (1).

141. (modified from Ortega et al., 2000: char. 53): Quadrate condyles with poorly developed intercondylar groove (0), or medial condyle expands ventrally being separated from the laterally condyle by a deep intercondylar groove (1).

142. (Ortega et al., 2000: char. 70): Nasal participation in antorbital fenestra: yes (0), or no (1).

143. (Ortega et al., 2000: char. 75): Anterior opening of temporo-orbital in dorsal view exposed (0), or hidden in dorsal view overlapped by squamosal rim of supratemporal fossa (1).

144. (modified from Ortega et al., 2000: char. 98): Coronoid size: short and located below the dorsal edge of the mandibular ramus (0), or anteriorly extended with posterior region elevated at the dorsal margin of the mandibular ramus (1).

145. (Ortega et al., 2000: char. 130): Lateral contour of snout in dorsal view: straight (0), or sinusoidal (1).

146. (Ortega et al., 2000: char. 138): Pterygoidean flanges: laminar and expanded (0), or bar-like and elongated (1), or bar-like and poorly developed (2).

147. (Ortega et al., 2000: char. 30): Prefrontal pillars when integrated in palate: pillars transverselly expanded (0), transversally expanded in their dorsal part and columnar ventrally (1), or longitudinally expanded in their dorsal part and columnar ventrally (2).

148. (modified from Ortega et al., 2000: char. 156): Position of first enlarged maxillary teeth: second or third alveoli (0), or fourth or fifth alveoli (1), homodont (2).

149. (Pol and Apesteguia, 2005: char. 181): Posterior peg at symphysis: absent (0), or present (1).

150. (Pol and Apesteguia, 2005: char. 182): Posterior ridge on glenoid fossa of articular: present (0), or absent (1).

151. (Pol and Apesteguia, 2005: char. 186): Choanal septum shape: narrow vertical bony sheet (0), or T-shaped bar expanded ventrally (1), or massive and blocky (2).

152. (Pol and Norell, 2004a: char. 164): Cross section of distal end of quadrate: mediolaterally wide and anteroposteriorly thin (0), or subquadrangular (1).

153. (Pol and Apesteguia, 2005: char. 188): Lateral surface of dentaries below alveolar margin, at mid to posterior region of tooth rows: vertically oriented, continuous with rest of lateral surface of dentaries (0), or flat surface exposed laterodorsally, divided by a ridge from rest of the lateral surface of the dentaries (1).

154. (Pol and Norell, 2004a: char. 165): Palatine-pterygoid contact on palate: palatines overlie pterygoids (0), or palatines firmly sutured to pterygoids (1).

155. (Wu et al., 1997: char. 108.): Postorbital participation in infratemporal fenestra: almost or entirely excluded (0), or bordering infratemporal fenestra (1).

156. (Wu et al., 1997: char. 112): Posteroventral edge of mandibular ramus: straight or convex (0), or markedly deflected (1).

157. (modified from Wu et al., 1997: char. 119): Quadrate ramus of pterygoid in ventral view: narrow (0), or broad (1).

158. (Wu et al., 1997: char. 127): Depression on posterolateral surface of maxilla: absent (0), or present (1).

159. (modified from Wu et al., 1997: char. 128): Paired anterior palatal fenestra: absent (0), or present (1).

160. (Pol and Norell, 2004a: char. 178): Paired ridges located medially on ventral surface of basisphenoid: absent (0), or present (1).

161. (Pol and Norell, 2004a: char. 181): Orientation of quadrate body distal to otoccipital-quadrate contact in posterior view: ventrally (0), or ventrolaterally (1).

162. (Pol and Norell, 2004b: char. 185): Ridge along dorsal section of quadrate- quadratojugal contact: absent (0), or present (1).

163. (modified from Pol and Norell, 2004b: char. 186): Sharp ridge on the surface of the angular: absent (0), or present on the ventral-most margin (1), or present along the lateral surface (2).

164. (modified from Pol and Apesteguia, 2005: char. 220): flat ventral surface of internal nares septum: parallel sided (0), or tapering anteriorly (1), or expanding anteriorly (2).

165. (Pol and Apesteguia, 2005: char. 221): Perinarial fossa: restricted extension (0), extensive, with a distinctly concave surface facing anteriorly (1), or large concave surface facing anteriorly, projecting anteroventrally from the external nares opening toward the alveolar margin (2).

166. (modified from Sereno et al., 2001: char. 67): Premaxillary palate circular paramedian depressions: absent (0), present located anteriorly on the premaxilla (1), or present located at the premaxilla-maxilla suture (2).

167. (Pol and Apesteguia, 2005: char. 223): Posterolateral region of nasals: flat surface facing dorsally (0), or lateral region deflected ventrally, forming part of the lateral surface of the snout (1).

168. (Pol and Norell, 2004a: char. 183): Choanal opening opened posteriorly and continuous with pterygoid surface (0), or closed posteriorly by an elevated wall formed by the pterygoids (1).

169. (Zaher et al., 2006: char. 198): Ectopterygoid projecting medially on ventral surface of pterygoid flanges: barely extended (0), or widely extended covering approximately the lateral half of the ventral surface of the pterygoid flanges (1).

170. (Gasparini et al., 2006: char. 236): Evaginated maxillary alveolar edges: absent (0), or present as a continuous sheet (1), or present as discrete evaginations at each alveolus (2).

171. (Gasparini et al., 2006: char. 237): Foramen in perinarial depression of premaxilla: absent (0), or present (1).

172. (Sereno et al., 2001: char. 27): Frontal anterior ramus with respect to tip of prefrontal: ending posteriorly (0), or ending anteriorly (1), or subequal (2).

173. (modified from Sereno et al., 2001: char 68): Premaxillary anterior alveolar margin orientation: vertical (0), or inturned (1).

174. (Sereno et al., 2001: char. 69): Premaxillary tooth row orientation: arched posteriorly from midline (0), or angled posterolaterally at 120 degree angle (1).

175. (Sereno et al., 2001: char. 70): Last premaxillary tooth position relative to tooth row: anterior (0), or anterolateral (1).

176. (Gasparini et al., 2006: char. 242): Posterior teeth with rings of ondulated enamel: absent (0), or present (1).

177. (modified from Brochu, 1999: char. 108 by Gasparini et al., 2006: char. 243): Maxilla-palatine suture: palatine anteriorly rounded (0), or palatine anteriorly pointed (1), or palatine invaginated (2).

178. (Gasparini et al., 2006: char. 244): Lateral surface of postorbital bar: formed by postorbital and jugal (0), or only by postorbital (1).

179. (Gasparini et al., 2006: char. 245): Enlarged foramen at anterior end of surangular groove: absent (0), or present (1).

180. (Gasparini et al., 2006: char. 246): Shape of antorbital fossa: subcircular or subtriangular (0), or elongated, low, and oriented obliquely (1).

181. (Gasparini et al., 2006: char. 247): Prefrontal lateral development: reduced (0), or enlarged, extending laterally over the orbit (1).

182. (Gasparini et al., 2006: char. 248): Foramen for the internal carotid artery: reduced, similar in size to the openings for cranial nerves IX-XI (0), or extremely enlarged (1).

183. (Gasparini et al., 2006: char. 249): Squamosal posterolateral region, lateral to paraoccipital process: narrow (0), or bearing a subrounded flat surface (1).

184. (Gasparini et al., 2006: char. 250): Posteromedial branch of squamosal oriented: transversely (0), or posterolaterally (1).

185. (Gasparini et al., 2006: char. 251): Dorsal margin of squamosal occipital flange: straight (0), or dorsally concave (1).

186. (Gasparini et al., 2006: char. 253): Longitudinal depressions on palatal surface of maxillae and palatines: absent (0), or present (1).

187. (Gasparini et al., 2006: char. 255): Transverse process of sacral vertebrae directed: laterally (0), or markedly deflected ventrally (1).

188. (Gasparini et al., 2006: char. 256): Prefrontal and lacrimal around orbits: forming flat rims (0), or evaginated, forming elevated rims (1).

189. (Gasparini et al., 2006: char. 257): Nasal bones: paired (0), or partially or completely fused (1).

190. (Brochu, 1997: char. 3): Posterior half of axis neural spine: wide (0), or narrow (1).

191. (Brochu, 1997: char. 19): Axial hypapophysis: with deep fork (0), or without deep fork (1).

192. (Brochu, 1997: char. 27): Olecranon process of ulna narrow and subangular (0), or wide and rounded (1).

193. (Brochu, 1997: char. 29): M. teres major and M. dorsalis scapulae insert separately on humerus: scars can be distinguished dorsal to deltopectoral crest (0), or insert with a common tendon; single insertion scar (1).

194. (modified from Brochu, 1997: char. 53): Anterior dentary alveoli project anterodorsally (0), or weakly procumbent (1), or strongly procumbent (2).

195. (Brochu, 1997: char. 84): Dorsal and ventral rims of squamosal groove for external ear valve musculature parallel (0), or squamosal groove flares anteriorly (1).

196. (Brochu, 1997: char. 91): Ectopterygoid abuts maxillary toothrow (0), or maxilla broadly separates ectopterygoid from maxillary toothrow (1).

197. (modified from Brochu, 1997: char. 103): Lateral margins of frontal: flush with skull surface (0), or elevated, forming ridged orbital margins (1).

198. (Brochu, 1997: char. 130): Capitate process of laterosphenoid oriented laterally (0), or anteroposteriorly (1) towards midline.

199. (Brochu, 1997: char. 153): Incisive foramen completely situated far from premaxillary toothrow, at the level of the second or third alveolus (0), or abuts premaxillary toothrow (1), or projects between first premaxillary teeth (2).

200. (Turner, 2004b: char. 129): Posterior process of cervical rib shaft lacks (0), or possesses (1) a posterodorsally projecting spine at the junction with the tubercular process.

201. (Pol et al., 2009: char. 278): Anterior half of interfenestral bar between suborbital fenestrae: lateral margins are parallel to subparallel (0), or flared anteriorly (1).

202. (Pol et al., 2009: char. 279): Posterior half of interfenestral bar between suborbital fenestrae: lateral margins are parallel to subparallel (0), or flared posteriorly (1).

203. (Pol et al., 2009: char. 280): Posteroventral margin of the angular straight or gently arched dorsally (0), or strongly arched dorsally (1).

204. (Buckley and Brochu, 1999: char. 106): Scapular blade width no more than twice the length of the scapulocoracoid articulation (0), or scapular blade very broad and greater than twice the length of scapulocoracoid articulation (1).

205. (Buckley et al., 2000: char. 115): Vomer exposed on palate (0), or not exposed or exposed as a consequence of an incomplete secondary palate (1).

206. (Turner and Buckley, 2008: char 287) Lateral expansion of posterodorsal edge of surangular anterior to glenoid fossa: absent (0), or present (1).

207. (Turner and Buckley, 2008: char. 288) In lateral view, anterior process of the squamosal extending to the orbital margin, overlapping the postorbital: absent (0), or present (1).

208. (Wilberg pers. com.) Prefrontal shape: quadrilateral with irregular outline (0), teardrop-shaped, with a convex outer margin (1), teardrop-shaped with a dorsal margin forming a 90 degree angle having a distinct triangular shape (2), teardrop- shaped with a smooth convex outer margin not exceeding the jugal bar in dorsal view (3), teardrop-shaped with the inflexion point directed posteriorly at approximately 70 degree angle from the antero-posterior axis of the skull (4), or teardrop-shaped with medial and lateral edges parallel with the inflextion point directed posteriorly at approximately a 50 degree angle from the antero-posterior axis of the skull (5).

209. (Wilberg, pers. com.) Radius: elongate (0), or greatly reduced (1). 210. (Wilberg, pers. com.) Humerus deltopectoral crest: present (0), or absent (1). 211. (Wilberg, pers. com.)Metacarpal I: elongate (0), or broadly expanded (1). 212. (Wilberg, pers. com.) Sacral rib curvature: little/none (0), or strong (1).

213. (Angle between medial and lateral processes of the frontal: approximately 90 degree angle (0), approximately 45 degree angle, or more acute (1), or approximately 60-70 degree angle (2)

214. Infratemporal fenestra in lateral view: considerably longer in length than the orbit (0), equal/subequal in length to the orbit (1), or shorter in length than the orbit (2).

215. Calcaneum tuber: well developed, with long neck (subequal in length to main body of calcaneum ± 5%), distal end wider than main body of calcaneum and projects inwards the body at > 80 degrees (0), poorly developed, short neck (< 50% length of the calcaneum main body), distal end < 50% the width of calcaneum main body width and projects straight from calcaneum (1), or absent/vestigial (2).

216. Transverse ridge crossing frontal rostromedial to orbits: absent (0), prominent anteriortly-curved shelf (transverse intraorbital crest sensu Andrade and Hornung, 2011) present (1), or intraorbital crest reduced to frontals only (2).

APPENDIX B DATA MATRIX USED IN PHYLOGENETIC ANALYSIS

Gracilisuchus 000??000000000000?0?00000000?0?000000?000000?0?0?000?100000?00000000?0?00 00?0?000001012?0????0?01?1000??101??000001002???0000???00???00000?000??00?? ??????0000?00?0?00?0?0??0000?0?00??1???001???0?10010000?010000?????0 Protosuchus 21000?0000110100021000001000001000201001100101103011?1102100010101000000 [1234]00?12001101011101101000[01]00000?01?01?10010[01]01010000???110000000 0000001?000???0?0100010??110?0?00?0000000?000000000000????0??0??0???0?0000 00????0 Hemiprotosuchus ?00???????10010?0??00?001010?01?0020?00?1001011?3?11??1?21????01???????0???? 1200?1?101??0??????000000??1000??00000??10???00???????0??0??00?0????00?????0 ???010?00?0???00?????00???00???00??00??????????????0???0000????? Orthosuchus 210001?0001001000[01]1000001000000000201100100??1?03011?0?0?0?00100010001 00000?120010010211411?1010?10000001010100000?000???00001?000???101000001? ?00????000?0010?01?0?00?0?000?0?0?0000?0000?00??????????????00??0000????? Zosuchus 200?00000??001010[01]110?0011022101012?1??01000?0?0311110????0?01111??????? ???????????1?12?3?1????001001101100010?0010112?0001????0?010001[01]000?[01]1 1?0??0???100?01011010?0000000000?0?00?0?0000?00????10?0?????0??000000????? Fruitachampsa 20000010001000010010000011022?110020112?0?0??01?3?31?????1?0111101011?000 11112?0??1???[01]00??101?0010?010010??11?0011?011??0??0010????001?00?0???00 0??????110??01???0??00?00?????0???????0?0?00????0010???100?1000000????0

Notosuchus 1001101001110001111110011002210100211120000010103111112?0101110001[01]11 2?200001000??0122011?10001[01]111[01]010010000111111111100111101010000011 0000110?000?110111100011000?10011[01]100000?0000?0000000????00101??0?01100 0000????0 Malawisuchus 100?20000?[01]10001[01][01]1100?1100221010011??20000?10?03?111[01]2?0101110 001?????210000010??01[12]2111?1?0??0110101?1100??110110101?0001????0100?10 100000110?00?0010111100000000?[01]?01000???00???00??00??00?????0?0?????0010 10000????? Simosuchus 100100000100010111110?01100211010011?110000010?030112121010110000?????02 100?2010?10002010?1?????111101212000110011100210120???10?000101[01]00[12]0 110?00[01]?11111111[01]000001210010[01][01]00100?000000000?0011??0010?????0? 0010000????0 Sphagesaurus 10002??00??100?????110??????210100?????0000??????13?2????????100??????1?????? ?????????312?0??????111110111111111111100101111?010?0??100??0?0????00????01 111?00000???1001?00?000000???????0?00????0?1??[02]?0???1??0000????0 Libycosuchus 200?2??00??10?010?1???01100?2101?0?111200?0??0?0?011??210001?1000?????????? ????????1[01]2010??????01?11???????????011?00????01????0?00??01?00?011??00??? 0?011?0000?1?0?[12]001?[01]0?000?????0??000?0?????00?0?????0????0000????? Araripesuchus tsangatsanga 200121?001110001011111011002210100111120000110102011212100011110?1111101 300010?0???110?000100010101001??100?00001111020001101?1?110010100210??0?1 01011010110000100100?0100000000?000000000?00?1??00?01?1??011000000????0

Bernissartia 20212??00111000?00111?0010002???0001112?0100?0???1?1??410010102011?1?2?020 021110110100000????????1???1?????????0?10???0??0?????0?31?1001022101101011? 0?0?1???00?1?????010???0???????0??00??0010000010?10?0101000000????0 Borealosuchus 202120010111000100111?00100021012111112101001010?13100310001102011111211 3111?110?10?000001?10100?11??1111??00000010001???00001000310?00?02210???1 0[12]1?0?001100000100?000100??000000??00?0000000?010000?1000001000000????0 Eothoracosaurus mississippiensis [12]02120010??11?[01]100111?001000210?2?011?2??11????0?131??3100??20000???? ???[1234]111[12]1????1?00000??1???01?1??12110?000?00100?1???00???000010?0010 2210???00?200?001?0?00010??000?00010000100???0000?00????0100?0?000?1000000 0???0 Gavialis 212120011111101101111100100021012101112001101010113100310001200000111201 31112111100?000001?1010[01]?11?212110?0000001000101?00001000010000102210? 1000[12]200?001100000100?000100010000[12]00?00000001011100101000001010000 000???0 Asiatosuchus germanicus 202120000111000101111?001000210?21111?2???0????0?131003?000110?012?1?2?1? 11?????????000001???110111??1[12]110???0000100110??000??000?10?001022?0?1?1 0?100?0?110?00?10??0001?000000?000?0?0000?001011000001??0001000000????0 Crocodylus 2021200[01]01110001021111001000210?2111112001001010113100310001002012111 20131112021100?000001010100?11??12110?0000001001101?00001000310?00102210 ?101021?00001100000100?000100000000?00?00000000010110000110000010000000? ??0

Diplocynodon hantoniensis 202120010111000101111?001000210?211111200100?010?13100310001002011111201 31112021110?00000?01?1?0111??111[01]0??000001001101000000000310?00102200? 1?10?100?00110?00?100?000100?00??0000?000000?000011001011011001000000???? 0 Alligator 202?2?000111000102111100100021012111112001001010103120310001002012111211 31112021100?000001010100111??11100?0000001001[12]01?00001?00310100102200? 101021?00001100000100?000100000000000?00000000000111011110110010000000?? ?0 Hsisosuchus 21?????000000100001100011002210100[12]??12?0000?0?0[23]?11114?00[01]01?1??1 0?????000?1000???001?1011????0101?????00000?000?1011??001???10??0??0?00020? 10?00?0000?10000111[01]?1???010??0???0??000?000??00????00?0?????00?0000000?? ?0 Pelagosaurus typus 20112001102010110000000000102101000001100010010001201?300000200001101?00 0000120001110110010??100?11?21??1000?0000101011?100???0000??00011020010?0 2?20?0001100000100???110000000011000[01]1001?00???????0??????0????00000110 Stenosaurus bollensis 1011200110201001000100000010210100?00110001011?00120103?000?200001101100 000?12000111011?01000100111?210?1000?000010111??000001000010000110?00101 02?2000001101001100?0011000000101100001000100???????0??0??????00000?0000 Metriorhynchus superciliosus 1021200111201001000100000010210100?00110001011?001201?300010200000101100 000??????0?0012?0?0000?1111?210?1000?0?0010101?0?000?0?000?0000110?0000102 ?200000110?000100?0011000000002001111111100??????????????????210?10110

Metriorhynchus casamiquelai 0????0011?20?0010?010??00??0210101?0??1????0???0?1201????010?0000??????????? ???????001??0?0?????111?21??10?????0010??????0???????0???00110??00??0??2??0?0 ??0?0??1?????11?0?00?00???111?11??00??????????????????21???01?? Geosaurus araucanensis 002?20011120100100010000001021010000?1100010[01]1?000400?3?001020000????? ?0000????????0012?00000??1111?210?101??0?0010?[01]2???00????0?010000110?000 010?[01]20??00?10?00?100?001??0000000211111111??00??????????????????1????12? 0 Geosaurus suevicus 012????11??01?0100010?000010????????????????????0040??3??01020000??????????? ????????0?2?0?????01111?21??1????0?0010?02???0?001??00???00110???0??0??2???? 0??0?0????0?0?1??0?00000???11?11???00??????????????????111111220 Dakosaurus maximus 002????11??0??010?0???00?01?????????????????????01?0?????01??0000????????????? ????????2?0??????1?0??21??1????0?0011?????????????00??00011???????0???????0??? ?0????0?0??????00001??1?1????1100??????????????????4?1?11??? Dakosaurus andiniensis 00??20011?20000100010000001021010000??1???10???00?400???0010?0000????????? ?????????0012?0?0????1101?21??101??010011112???00001?0?0??000110?0000?0??0? ??00?1000????0???11????0001?11111111??00??????????????????4????11?0 Rhabdognathus 20??200??111000100110110101121011[01]011120011010?11?302?????????0????????? ?????????????1?10?0??????10?????0??00??0[01]10020?1?00????1????0?01[01]2200?0? 0002??00?11?00001?????0112?1???010??000000?11?????0001??000?1?00000?00?0 Sokotosuchus

2?212??10????001001???10101121?????1112??11??0?1?1?0?????????01?????????????? ?????????1?0????????1??????0??????0?10??????0????????????0?????????1??2???????? ????1????????2??000?????0??00??11??????????????????0000?00?? Dyrosaurus 202?2?010?11??010011??101011210110?11120011?1001113021???00?2?000????????0 0??????????1?10?0??????1??????0???0??0010020???0?????100?00001?0200???0??2??0 001?0?00?100?0?011201000010??00000???1????00101??01?????0000?00?0 Hyposaurus ?02?2??1???????1????0?101???2??????1?12001??10?1??3?????0???2?000??112???00?? ???????01??0?0??1?011?????????????0?1002????00????100100?0????00??????200?00? ???0??10???????2?????0?????0??????1????00?0?????0?1000000????0 Pholidosaurus 21212??1??1110?10011??00100121011001112?01??1000?131[12]???0???2?0????1?2?0 ??0??200????????0????????10??1?11????0?00100?????00?????00?0000102?10???00??? 000?11?000?1?????010??1????1???0?000??00??????????????????00000???? Sarcosuchus 202?200101?1000100110010100121011001?12?0100?00001312130?000210101?112??[ 01]00?1200?00?010[01]0?0[01]01?0?11??12110??0000010010???00???10?[012]11010[ 01]022[01]0?00000?00?00?10?00?100?010100011110100?0?0000?10????0010?00100? 1000000001?0 Terminonaris robusta 202?20010?11??01001???1010012101[01]0??????010??00?013[01][12]130000?210001 111200100?1200?10?0?0101?10100111??1?11??0?0?0010??????000011000???101[01]2 [12]10???00??00001??0?00?????0101020111101?0?0?00?0??01???00?01?00?00110000 0?00?0 Mahajangasuchus insignis

20[12]?21??0121100101111001100021011101112001001010??2121110100001111?112 ?140000100??0101[01]0011010101010?1211010010001110110001???01110100000210 000?00100110011010001002?001101000000000000000?01?????011??110?1?100000?? ??0 Stolokrosuchus 200020000111000101111000110121?1[01]?011120010010?0?021?0????0011[12]?????? ??????????????1?0000?0?????111101011110000001100111001???1??01110000210010? 0?110??00?110000100?0011?000000010?0000000?00?????01011?0?1?1010000????0 Argochampsa 20212001011???110[01]101?001001210?22??112?0110?0?0?1300????????00????????? ?????????????0000?0??????110?11110??00??010?0[12]0??00?????????0001022?0?1?00 2????0??1??0001???0?00??000?0010??0?0000??1???????000?0???1?00000????0 Piepehsuchus teleorhinus 011120011???1?0100000001?010210100?0?110011?????01201????????0????????????? ??????????02?0?0??????1??21??1?00?0??01?0??0??00??????????101??1?01??02?2??00 ?1???00110???000?0?2001?0??0001??1?00???????0???????1?00????00?? Stenosaurus brevoir 001020011???10010001?0000010210100??0110001????001201030000020000????????? ?????????0012?0?0????0?11?211?100??010010100??000???00?01?000110?0010?02?20 00001100001100?00110000001011000?11?0?00???????0?0???????00????00?0 Stenosaurus gracilorostris 000020011?20100100000000?010210101000?10001?????01101???????20000????????? ?????????0??2?0?0?????111?210?100??0000101?????00?????????00011?10010?02?2?? 00?11??00010??001100?0000??1?00?1001?00???????0?0???????00????0??? Geosaurus vignaudi

01???0?11?201?01000?0000101????????0????001?????0?20??3?001?2?0???????0?00?? ????????01??0?00???1??1??1??1010?????1??120??0?000???010??01102?00??0?12???0 0?10?0????0???1??0?0?????0111?111??001??????0?????????01??1?12?? Dakosaurus carpenter 001??0011?0???0100????01?010???????????????????00?40??3?0????00????????0000?? ??????????00??00????1?021??1????0?0011?????????????0???000?1???????0?????????1 ??0????????1??0?0?0??????1?1????00???????0?????????04????21?? Oceanosuchus 1011200101????110?10??10?0012?????0???2???0???0?0131???0??1?2?0???????001000 1200??0?000?0??1?????1??21110????0?0?100?????0?00??0?0?101010?1?0??00??20??? 1??0?00???2???0??20011101?0?000000?001???0??0?00??0?1?00????0??? Telosaurus cadomensis 00??200??1200?01000?00000012210101?01110101?11?00?200?????????????????????? ??????????1????0?????????????10010?1?0??10000100?????????0?011120010?0?0????0 ?010000000????1000?0???011?0011100?00???????00??????1?00????01?? Theriosuchus 201120100110000110111100110?2101001?11?0000???1?20211?410010101011011200 011?1200100101100??1011011?001?110?000?00110??0?0?00??0?0210100?0200010?1 011??0?011?0000??2?0?010??0000?0???000??0?00??0?0?011??00?0100000?????0 Alligatorium ?0???0000?1000010?111??0?10???0?00??11??000?????20?1????00101?101?0112000?? ?1?00100????????10?1????????????????0??????????????????1??????????????????????? ???????????????????0?????????????00??????????????0???000?????? Pachycheilosuchus 20?????????00??10?111???100???????01?1????????????2??????01?0?0?010111?0211?1 000??0????001?100?0?1?????????????0?1??0?????????0?0010??0??2?????????1????0?? 0?0???????????0???????????????????????0?????0??00?0??00??????

Vectisuchus 21?02??10???0?11001101?1000?210??0??????010???????31?03??00?2?????????0?000? 120???0????00?0???1?1?1?????0?????0?0100?1???0????0?????0?0102??0?1?0??1????0 ?10?00?1?????0??0???????0?????0???1????????1???????1?0????????? Calsoyasuchus 202010010?110?0102101100100011??00?1112?0?0??0?0?111?????????01???????????? ????????0?0100?0??????11?21111?0??0?0010001???00??????????001022?01??1??1??0 0?????1101??0000??000000??0?100?0?0?00?????0?01??????1?000000???0 Eutretauranosuchus 202?11210111000100111100100011010001112001001010?131204?00001010111????0 ??0?12????1??0000?01???0?110210111???01?01000110?00???00?3??000102210?1?1?1 1?0000?10010?100?00010000?00000??0?0000?00??????????????01?00?????0?1 Goniopholis simus 202110010111000100111?001000210100?11120010?10?021312?410000?01011?1???0? 0001200?11?0000010101101110210110?00010010001???00001100311000102210?1?1 011?0000110010?100?00010000?000000?0?0000?00??????????????010000000?0?1 Goniopholis baryglyphaeus 201?10010111000?0?11110010002101???1112??10??0?0?131??4??01?101????????0?0 001200?11?000?0?0??????1?021011????0??0100?????00??????31?00010?2?0?1?1??1?? ?0?110?10?10???00??000?000??????000???0?????????????????0????????? Goniopholis willetti 212?10010111001100111?001000210100?11120?10????021312????????01???????????? ?????????00000?0???????10210110???00?01?001?1?00??????????00112200?1?10?1??0 0?11?010?10??0001?000?00?00??0?0000?00???????0???????1?00?????0?2 Anetophthalmosuchus

20??10010?100001001111001000210100?11120??0????02?312??1001?1??????????0?0 0?1200?11?00?00?010??11?1?21?110????0?01?020???00???0??311??01022?0?1?10??? 0000110?10?1?0???01?0?0????00??0?0000?00???????0?????0?100000???0?0 "Goniopholis" felix 202??1210?1100000?111?001000?????????????????????131????????????????????????? ??????????0??????????1?21?11?????0??1?001?????????????????10221??1?1?????????1 ??1????????0????0?????0??0?0?0??00???????0?????????00?????0?1 Amphicotylus stovalli 202011210?11000100111?001000110100?11120010??0?0?1312?4??????01??????????0 0012????1??0000?0??????110210110???00?010001???00?????????000102210?1?1011? ?00?11?010?10??00010000?00000??0?0000?00???????0???????1?00?????0?1 Amphicotylus lucasii 202011210?11000100111?001000110100??112??10??0?0?13120????0??01???????????? ?12?0??1??0010?01?????110210110???00?010001???00???1?????0001022?0?1?10?1?? 00?11??10?10??0001?0?0?00?00??0?0000?00???????0???????1?00?????0?1 Amphicotylus gilmorei 202011210?11000100111?001000110?00??1120?10??0?0?1312????????01???????????? ??????????0010?0??????110210110???00?010001???00?????????0001022?0?1?10?1??0 0?11??10?10??0001?0?0?00?00??0?0000?00???????0???????1?00?????0?1 Sunosuchus junggarensis 201011?10?11000?0?111100100021??00111?20?10??01021312?41?00110101??11????0 001200?11??00?0?0??????1?02???1????????10001???0??????????00010?2???1???????? ???1??10?1?????0??000?000??????000???0???????????????1?0????????2 Siamosuchus 201??0?00?11000?0?1110001?0021???001112??10??0?0?131?????????01????11??0?00 01200?11?000?0?01?????1?021011????0??0100?????00???????1?0001??2?0?1?1??1??? 0??10?10?10???00??000?0000?????000???0?????????????????0????????2

Denazinosuchus 20?120010??1??01021??????00?2????????????????????131?1????102?1???????????0?1 200?11????10??1????101?212111???0??110????????????????10?01??1?????1??1?????0 ?0?0????0???0??0?2???0????0????0?10???????0?????0?1??3????0??0 Arlington Form 20????????1?0?????110?0100?02?0???011121??0???????3???3??01???210??11??0?000 1200?11?0??10?01?1?0?0?????10???????1000?1???00???0100??0?0??22?1?1?1??1?0?0 00???0??1?0???0??0?????000????000??10????0?????0???0??0?????????

APPENDIX C CHARACTER TRANSITIONS

Node A. Unambiguous synapomorphies: 38(10), 98(10) Synapomorphies only found in some trees: 31(01), 44(01), 57(02), 58(01), 117(01) Node B. Unambiguous synapomorphies: 16(01), 28(02), 32(01), 63(01), 102(01), 114(01), 135(01) Synapomorphies only found in some trees: 29(02), 30(01), 39(02), 68(01), 69(01), 93(10), 118(01), 123(01) Node C. Unambiguous synapomorphies: 24(01), 97(01), 99(01) Synapomorphies only found in some trees: 78(20), 157(10), 168(01) Node D. Unambiguous synapomorphies: 10(01), 12(01), 18(01), 155(01), 159(10) Synapomorphies only found in some trees: 14(10), 154(01) Node E. Unambiguous synapomorphies: 60(01), 63(10), 86(12), 100(01), 113(01), 141(01), 165(01) No unambiguous synapomorphies Node F. Unambiguous synapomorphies: 1(21), 17(01), 58(01), 150(01), 152(01) No umambiguous synapomorphies Node G.

Unambiguous synapomorphies: 112(01), 122(01) Synapomorphies only found in some trees: 21(01), 50(01) Node H. Unambiguous synapomorphies: 49(32), 88(10), 138(02), 153(10) Synapomorphies only found in some trees: 21(01), 35(20) Node I. Unambiguous synapomorphies: 6(01), 72(01), 98(10), 116(01), 130(01), 150(01), 200(01), 204(01) No unambiguous synapomorphies Node J. Unambiguous synapomorphies: 24(10), 54(10), 91(10), 120(01), 131(01), 148(01) No unambiguous synapomorphies Node K. Atoposauridae Unambiguous synapomorphies: 12(10), 59(01), 74(01), 75(01) No unambiguous synapomorphies Node L. Unambiguous synapomorphies: 26(01) No unambiguous synapomorphies Node M. Unambiguous synapomorphies: 18(10), 22(01), 62(10), 119(10), 123(10), 127(10) Synapomorphies only found in some trees: 3(01) Node N. Unambiguous synapomorphies: 8(01), 50(01), 51(23), 101(02), 115(10) No unambiguous synapomorphies

Node O. Unambiguous synapomorphies: 86(10), 130(03) Synapomorphies only found in some trees: 140(01) Node P. Unambiguous synapomorphies: 63(12), 78(21), 79(01) Synapomorphies only found in some trees: 3(12) Node Q. Unambiguous synapomorphies: 33(02), 34(01), 55(43), 60(01), 72(01), 74(01), 75(01), 196(10) Synapomorphies only found in some trees: 35(01) Node R. Unambiguous synapomorphies: 80(01), 82(10), 192(01) Synapomorphies only found in some trees: 66(12) Node S. Unambiguous synapomorphies: 15(01), 43(01), 145(10), 177(01), 199(10) Synapomorphies only found in some trees: 63(20) Node T. Unambiguous synapomorphies: 103(12), 172(01) No unambiguous synapomorphies Node U. Unambiguous synapomorphies: 61(10) Synapomorphies only found in some trees: 78(10), 79(12) Node V. Unambiguous synapomorphies: 140(10), 190(10), 196(01), 201(01), 202(01) No unambiguous synapomorphies

Node W. Goniopholididae Unambiguous synapomorphies: 5(21), 158(01) Synapomorphies only found in some trees: 83(01), 132(01) Node X. Unambiguous synapomorphies: 29(21) No unambiguous synapomorphies Node Y. Unambiguous synapomorphies: 6(01), 7(02) Synapomorphies only found in some trees: 216(01) Node Z. Unambiguous synapomorphies: 88(01) No unambiguous synapomorphies Node AA. Unambiguous synapomorphies: 21(10), 55(43), 60(12) No unambiguous synapomorphies Node BB. Unambiguous synapomorphies: 24(01), 188(01) No unambiguous synapomorphies Node CC. Unambiguous synapomorphies: 59(01), 83(01), 88(01), 113(01) No unambiguous synapomorphies Node DD. Unambiguous synapomorphies: 63(10), 95(10), 148(12) No unambiguous synapomorphies Node EE. Unambiguous synapomorphies: 28(01), 33(01), 172(21) Synapomorphies only found in some trees: 3(12), 54(01), 88(01)

Node FF. Unambiguous synapomorphies: 132(01), 134(01) Synapomorphies only found in some trees: 62(01), 173(01), 174(01) Node GG. Dyrosauria Unambiguous synapomorphies: 48(01), 189(01) Synapomorphies only found in some trees: 27(01), 49(01), 52(10), 105(10), 128(01), 169(01), 187(01) Node HH. Thalattosuchia Unambiguous synapomorphies: 1(20), 9(01), 19(10), 36(10), 39(21), 51(32), 87(02), 146(02), 160(01), 183(01) Synapomorphies only found in some trees: 13(01), 27(01), 52(10) Node II. Unambiguous synapomorphies: 42(10), 116(01), 167(01) Synapomorphies only found in some trees: Node JJ. Unambiguous synapomorphies: 84(10), 111(01), 184(01) No unambiguous synapomorphies Node KK. Unambiguous synapomorphies: 121(01), 160(10), 214(01) No unambiguous synapomorphies Node LL. Unambiguous synapomorphies: 186(01) No unambiguous synapomorphies Node MM. Unambiguous synapomorphies: 20(10), 111(10), 184(10) No unambiguous synapomorphies Node NN.

Unambiguous synapomorphies: 59(01), 142(10), 180(01), 181(01), 185(01), 209(01) Synapomorphies only found in some trees: 3(12), 96(01), 177(12), 208(02), 212(01) Node OO. Unambiguous synapomorphies: 51(24), 53(10), 213(01) Synapomorphies only found in some trees: 107(01), 118(12), 179(01), 210(01) Node PP. Unambiguous synapomorphies: 115(01) No unambiguous synapomorphies Node QQ. Unambiguous synapomorphies: 98(01) No unambiguous synapomorphies Node RR. Unambiguous synapomorphies: 50(10), 214(10) No unambiguous synapomorphies Node SS. Unambiguous synapomorphies: 2(01) No unambiguous synapomorphies

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