3764 (3): 240 278 Article · A NEW AETOSAUR FROM BRAZIL. Zootaxa 3764 (3) © 2014 Magnolia Press · 241. comprises Chilenosuchus forttae Casamiquela, 1980 (Estratos El Bordo) (Desojo

ZOOTAXA

ISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright copy 2014 Magnolia Press

Zootaxa 3764 (3) 240ndash278

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A new aetosaur from the Upper Triassic of the Santa Maria Formation southern

Brazil

LUacuteCIO ROBERTO-DA-SILVA1 3 JULIA B DESOJO2 SEacuteRGIO F CABREIRA3

ALEX S S AIRES1 RODRIGO T MUumlLLER4 CRISTIAN P PACHECO4 amp SEacuteRGIO DIAS-DA-SILVA45 1Programa de Poacutes-Graduaccedilatildeo em Ciecircncias Bioloacutegicas da Universidade Federal do Pampa E-mail lucio_bioyahoocombr 2Museo Argentino de Ciencias Naturales ldquoBernardino Rivadaacuteviardquo Av Aacutengel Gallardo 470 Buenos Aires Argentina E-mail julidesomacngovar 3Laboratoacuterio de Paleontologia da Universidade Luterana do Brasil Canoas-RS Brasil E-mail sergiofurtadocabreirayahoocombr 4Laboratoacuterio de Paleontologia Universidade Federal do Pampa Av Antocircnio Trilha 1847 Satildeo Gabriel-RS Brasil E-mails asschilleragmailcom rodrigotmullerhotmailcom crispachecorsyahoocombr paleospgmailcom5Corresponding author

Abstract

Aetosaurs are armored pseudosuchian archosaurs widespread in Upper Triassic units In South America four taxa were

previously recorded Aetosauroides scagliai Neoaetosauroides engaeus Aetobarbakinoides brasiliensis and Chilenosu-

chus forttae Herein we describe a new Late Triassic juvenile aetosaur from the Santa Maria Formation of southern Brazil

Polesinesuchus aurelioi gen et sp nov increasing the paleobiodiversity of this interesting group to five taxa in Western

Gondwana The holotype is composed of cranial (parietal and braincase) and postcranial elements (cervical dorsal sacral

caudal vertebrae both scapulae a humerus ilium pubis ischium tibia a partial right pes and anterior and mid-dorsal

paramedian osteoderms) It belongs to a juvenile individual as its neurocentral sutures are open in all vertebrae and also

due to its small size However future paleohistological investigation is necessary to fully corroborate this assumption

This new taxon is distinguished from all other aetosaurs by the presence of an unique combination of character states (not

controlled by ontogeny) such as cervical vertebrae with prezygapophyses widely extending laterally through most of the

anterior edge of the diapophyses absence of hyposphene articulations in both cervical and mid-dorsal vertebrae presence

of a ventral keel in cervical vertebrae anterior and mid-dorsal vertebrae without a lateral fossa in their centra expanded

proximal end of scapula anteroposteriorly expanded medial portion of scapular blade a short humerus with a robust shaft

and a dorsoventral and very low iliac blade with a long anterior process which slightly exceeds the pubic peduncle Re-

garding its phylogenetic relationships the present analysis placed Polesinesuchus as the sister taxon of Aetobarbakinoides

and both as sister taxa of the unnamed monophyletic clade Desmatosuchinae plus Typothoracisinae

Key words western Gondwana Carnian Archosauria Pseudosuchia Aetosauria Stagonolepididae

Introduction

Aetosaurs are heavily armored quadrupedal archosaurs restricted to the Late Triassic They have been recovered in continental deposits from South and North America Greenland Europe Morocco and India (Lucas amp Heckert 2000 Parker et al 2008 Desojo amp Ezcurra 2011 Desojo et al 2013) (Fig 1) They are nested within Pseudosuchia the crocodile lineage of Archosauria however their phylogenetic relationships with other high-level suchians are still subject of intense debate (for a complete set of references see Desojo et al 2012 and Desojo et al

2013) Formerly four South American taxa have been described (from Argentina Brazil and Chile) (Desojo amp Ezcurra 2011 Desojo et al 2013) The Argentinean record comprises Aetosauroides scagliai Casamiquela 1960 from the Ischigualasto-Villa Unioacuten Basin (Carnian to Norian Ischigualasto Formation) and Neoaetosauroides

engaeus Bonaparte 1971 (Norian to Rhaetian Los Colorados Formation) (Casamiquela 1960 1961 1967 Bonaparte 1969 1971 Desojo 2005 Desojo amp Baacuteez 2005 2007 Desojo et al 2012) The Chilean record

240 Accepted by R Benson 8 Nov 2013 published 11 Feb 2014

comprises Chilenosuchus forttae Casamiquela 1980 (Estratos El Bordo) (Desojo 2003) In Brazil Aetosauroides

scaglai is restricted to the red beds of the Santa Maria Formation in the central region of Rio Grande do Sul State (Langer et al 2007) The first Brazilian material ascribed to aetosaurs came from the vicinities of the municipality of Satildeo Pedro do Sul at the locality of Inhamandaacute It was described by Zacarias (1982) in her unpublished Masterrsquos degree dissertation and named ldquoAetosauroides subsulcatusrdquo (Lucas amp Heckert 2001) This specimen was never formally described (Desojo 2005 Desojo amp Ezcurra 2011) and its taxonomic validity has been dismissed in further contributions as follows Heckert amp Lucas (2000) and Lucas amp Heckert (2002) stated that all aetosaurs from Brazil are junior synonyms of Stagonolepis robertsoni in a recent reappraisal of the taxonomic status of Aetosauroides scagliai Desojo amp Ezcurra (2011) concluded that ldquoA subsulcatusrdquo presents the same autapomorphies observable in the Argentinean A scagliai Desojo et al (2012) based upon new postcranial material (dorsal vertebrae) also from the Inhamandaacute locality erected the new Brazilian genus Aetobarbakinoides

brasiliensis under collection number CPE2 168 (this material also had also been regarded as belonging to Stagonolepis robertsoni by Heckert amp Lucas 2001) Several osteoderms and bone remains were attributed to an indeterminate Aetosaurinae (Langer et al 2007 Desojo amp Ezcurra 2011) due to both the fragmentary nature and non-diagnostic condition of the remains In 2005 new material from a small aetosaur specimen was collected from the municipality of Satildeo Joatildeo do Polecircsine central region of Rio Grande do Sul State described in the present contribution It comprises skull fragments postcranial elements part of the carapace belonging to the new juvenile aetosaur Polesinesuchus aurelioi gen et sp nov

FIGURE 1 Paleogeographic map depicting worldwide occurrences of Upper Triassic aetosaur specimens (redrawn from

Blakey 2006) 1 Stratos El Bordo (Chile) 2 Ischigualasto-Villa Unioacuten Basin (Argentina) 3 Paranaacute Basin (Brazil) 4

Pranhita-Godavari Basin (India) 5 Timesgadiouine Formation (Morocco) 6 Chinle Group (USA) 7 Newark Supergroup

(USA) 8 Keuper Group (Germany) and Alpine marine Triassic beds (Italy) 9 Fleming Fjord Formation (Greenland) 10

Lossiemouth Sandstone (Scotland) (Modified from Desojo amp Ezcurra 2011)

Geological and biostratigraphical setting The material comes from an outcrop located 6 km southeast from the Municipality of Satildeo Joatildeo do Polecircsine central region of the Rio Grande do Sul State (Fig 2) This outcrop is included in the Santa Maria 2 Sequence (Zerfass 2003 Langer et al 2007 Dias da Silva et al 2011 2012) which mainly comprises mudstones and a high concentration of early diagenetic carbonate interpreted as deposited in a dry climate (Zerfass et al 2003 Langer et al 2007 Dias da Silva et al 2011) These sedimentary rocks were deposited in a system of shallow lakes and floodplains of anastomosed ephemeral rivers (Langer et al 2007 Dias da Silva et al 2011 2012) The occurrence of the rhynchosaur Hyperodapedon justifies the inclusion of the Buriol outcrop in the Hyperodapedon Assemblage Zone which is dated as Carnian (see Zerfass et al 2003 Soares et al 2011) (Fig 3) Lucas amp Heckert (2002) chronologically attributed this Assemblage Zone as late to latest Carnian based upon the presence of the rhynchosaur genus Hyperodapedon also found in the Upper Triassic Popo Agie Formation of Wyoming USA These authors defined a Hyperodapedon biochron (Otichalkian-Adamanian) based upon strata from North America Scotland India Zimbabwe Tanzania Madagascar Argentina and Brazil (Schultz 2005)

Institutional abbreviations CPE 2 Coleccedilatildeo municipal Satildeo Pedro do Sul Rio Grande do Sul Brazil MCP Museu de Ciecircncias e Tecnologia Porto Alegre Rio Grande do Sul Brazil NHMUK The Natural History Museum London United Kingdom PVL Paleontologia de Vertebrados Instituto ldquoMiguel Lillordquo San Miguel de

Zootaxa 3764 (3) copy 2014 Magnolia Press middot 241A NEW AETOSAUR FROM BRAZIL

Tucumaacuten Argentina PVSJ Divisioacuten de Paleontologia de Vertebrados del Museo de Ciencias Naturales y Universidad Nacional de San Juan ArgentinaPEFO Petrified Forest National Park Arizona USASMNS Staatliches Museum fuumlr Naturkunde Stuttgart GermanyTTUP Texas Tech University Museum of Paleontology Berkeley California USAULBRA PVT Universidade Luterana do Brasil Coleccedilatildeo de Paleovertebrados Canoas Rio Grande do Sul BrazilUCMP Museum of Paleontology University of California Berkeley California USA UMMP Museum of Paleontology University of Michigan Ann Arbor Michigan USA

FIGURE 2 Location map of the municipality of Satildeo Joatildeo do Polecircsine (in red) with the Buriol outcrop from where the holotype

of Polesinesuchus aurelioi was recovered (Modified of Reichel et al 2009)

Systematic Palaeontology

Archosauria Cope 1869 sensu Gauthier amp Padian (1985)

Pseudosuchia Zittel 1887ndash1890 sensu Gauthier amp Padian (1985)

Aetosauria Marsh 1884 sensu Parker (2007)

Stagonolepididae Lydekker 1887 sensu Heckert amp Lucas (2000)

Polesinesuchus aurelioi gen et sp nov

Holotype ULBRAPVT003 comprising cranial and postcranial elements a partial parietal and a well preserved basioccipital a partial axial skeleton (cervical dorsal sacral and caudal vertebrae) an incomplete pectoral girdle (partial scapulae coracoids and interclavicle) two partial humeri and both ulnae a right pelvic girdle (ilium

DA-SILVA ET AL242 middot Zootaxa 3764 (3) copy 2014 Magnolia Press

ischium and pubis) parts of both hindlimbs (femora tibiae fibulae both astragali and a left calcaneum) and pes (metatarsal III and V unidentifiable phalanges and two ungual phalanges) Ten isolated osteoderms from dorsal ventral and appendicular region of the armor (see Tables 1 2 3 and 4 below for anatomical measurements of skeletal elements of ULBRAPVT003)

TABLE 1 Measurements (mm) of the cervical vertebrae of Polesinesuchus aurelioi AbbreviationsAFH anterior

articular facet height AFW anterior articular facet width CL centrum length MH maximum preserved heightPFH

posterior articular facet height PFW posterior articular facet width incomplete () distorted

TABLE 2 Measurements (mm) of the postcervical vertebrae of Polesinesuchus aurelioi AbbreviationsAFH anterior

Type locality and horizon Buriol outcrop municipality of Satildeo Joatildeo do Polecircsine (coordinates 29ordm39rsquo342rdquoS 53ordm25rsquo474rdquoW) in the central region of the State of Rio Grande do Sul Southern Brazil This locality is included in the Sequence 2 of the Santa Maria Supersequence Hyperodapedon Assemblage Zone (Fig 3) (Langer et al 2007 Soares et al 2011)

Etymology The generic name is derived from the name of the municipality from where the specimen was recovered (Satildeo Joatildeo do Polecircsine Rio Grande do Sul State) The specific name is in honor of the physician Pedro Lucas Porcela Aureacutelio a paleontology enthusiast from Rio Grande do Sul

CL AFH AFW PFH PFW MH

Cervical 2 09 085 08 085 082 12

Cervical 3 09 09 088 08 08 243

Cervical 4 093 09 092 085 088 085

Cervical 5 10 092 09 085 088 195

Cervical 7 105 09 10 09 088 20

Cervical 8 10 092 11 085 088 10

Ant Dorsal 1 105 088 102 08 085 20

Ant Dorsal 2 108 076 08 08 09 07

Ant Dorsal 3 112 081 083 07 078 08

Ant Dorsal 4 12 09 085 075 07 09

Ant Dorsal 5 122 09 085 075 071 09

Mid Dorsal 13 128 093 088 088 09 23

Mid Dorsal 14 133 095 098 098 103 202

Mid Dorsal 15 138 095 098 098 10 16

Sacral 1 14 112 131 10 13 122

Sacral 2 142 108 125 10 121 105

Caudal 1 13 112 108 12 115 12

Caudal 2 125 115 118 115 12 115

Caudal 3 142 (108) (092) (10) (091) 123

Caudal 4 15 091 088 09 088 095

Caudal 5 14 062 058 06 056 09

TABLE 3 Measurements (mm) of postcranial elements of Polesinesuchus aurelioi (all comprising the maximum

preserved measure) incomplete () distorted

Scapula

Length 495

Proximal transverse width 24

Proximal anteroposterior depth 35

Distal transverse width 23

Distal anteroposterior depth 75

Coracoid

Length (28)

Interclavicle

Length 45

Humerus

Length (60)

Mid-shaft width (mediolateral) 075

Mid-shaft thickness (anteroposterior) 06

Length 45

Length of the iliac blade 505

Height of iliac blade above to acetabular crest 104

Transverse width of acetabulum 235

Height of acetabulum 195

Ischium

Length 37

Length (50)

Proximal transverse width (205)

continued on the next page

TABLE 4 Measurements in millimeters (mm) of paramedian osteoderms (A B and C) of the Polesinesuchus aurelioi

(all the measurements are the maximum preserved) incomplete

Diagnosis Polesinesuchus is distinguished from all other aetosaurs by the following unique combination of characters cervical vertebrae with prezygapophyses widely extending laterally through most of the anterior edge of the diapophyses absence of hyposphene articulation in both cervical and mid-dorsal vertebrae anterior articular facet width of cervical vertebrae measuring less than 12 times the posterior one presence of a ventral keel in

TABLE 3 (Continued)

Length 90

Distance from head to fourth trochanter 35

Head width 10

Head length 22

Thickness across fourth trochanter 12

Mid shaft length 12

Mid shaft width 9

Intercondylar thickness 75

Length 70

Proximal width (mediolateral) 13

Proximal thickness (anteroposterior) 185

Distal thickness 135

Fibula

Length 72

Proximal width 68

Distance proximal end of trochanter 193

Thickness of trochanter 6

Distal width 8

Distal thickness 12

Metatarsals III

Length 32

Proximal width 38

Distal width 55

Distal thickness 59

1mdashWidth behind the anterior bar 43 42 33

2mdashDistance to center of ossification from medial edge 17 16 17

3mdashLength along medial edge 25 3 ----

4mdashThickness at center of ossification 10 092 09

cervical vertebrae anterior and mid-dorsal vertebrae without a lateral fossa in their centra expanded proximal end of scapula anteroposteriorly expanded medial portion of scapular blade a short humerus with a robust shaft and a dorsoventral and very low iliac blade with a long anterior process which slightly exceeds the pubic peduncle Polesinesuchus differs from Aetosauroides scagliai by the presence of an anteroposterior projection of the scapular shaft absence of a lateral fossa in the presacral vertebrae tip of the anterior process of the ilium longer and very pointed in comparison femoral distal ends slightly twisted along the axial length P aurelioi differs from Aetobarbakinoides by the presence of a ventral keel in cervical vertebrae absence of a circular pit in the neural spine absence of a hyposphene articulation in dorsal vertebrae the humeral shaft is very robust in comparison the larger tuberosity of the humerus is poorly defined in comparison and the tibial shaft is very robust in comparison In Polesinesuchus the ornamentation of the paramedian osteoderms shows a radial pattern with circular pits grooves and ridges originated from a low eminence The radial pattern observed in Polesinesuchus is different from the reticular one of Typothorax Redondasuchus Chilenosuchus Ornamentation is randomly distributed in Desmatosuchus smalli D spurensis Longosuchus and Acaenasuchus The paramedian osteoderm of Polesinesuchus is subretangular with a widthlength ratio of about of 33

FIGURE 3 Biostratigraphic chart of terrestrial Triassic faunas from Gondwana showing the Riograndia AZ Dinodontosaurus

AZ (sensu Barberena et al 1985) Santacruzodon AZ (sensu Abdala amp Ribeiro 2010) Hyperodapedon AZ (sensu Abdala et al

2001) Ans Anisian AZ Assemblage Zone Crn Carnian Ind Induan Lad Ladinian Mad Madagascar Nor Norian Ol

Olenekian Rht Rhaetic S Africa South Africa and the position of Polesinesuchus level Modified from Abdala amp Ribeiro

(2010) Geological Time Scale based on Gradstein and Ogg (2004) Modified from Soares et al (2011)

DescriptionThe specimen is well- preserved as most of the thin delicate and small elements (eg interclavicle sacral and caudal vertebrae and pedal elements) present a high quality of fossilization Just a few bones are weathered and crushed (eg right coracoid left humerus a few paramedian and ventral osteoderms)

Ontogenetic stage The recognition of ontogenetic stages in aetosaurs is problematic due to the lack of information regarding juvenile individuals (Parker 2008 Taborda et al 2013) Moreover the ontogeny of crocodile-line archosaurs is poorly known (Schoch 2007) In archosaurs the usual practice addressing this issue is to consider individuals with closed neurocentral vertebral sutures as fully grown adults (Brochu 1996) The small size of an individual is not reliable information to establish its juvenile ontogenetic stage In recent years

paleohistological analyses of growth marks in osteoderms arose as a possible solution to the understanding of ontogeny in the Aetosauria (Cerda amp Desojo 2011 Taborda et al 2013 Torsten et al in press) Still the ontogenetic stage of Polesinesuchus was tentatively proposed based solely in the observation of neurocentral sutures in which they are widely opened Hence here we tentatively propose that Polesinesuchus is a juvenile specimen Future paleohistological studies are needed to obtain a more accurate idea regarding the ontogenetic stage of Polesinesuchus (meaning estimating how immature this individual was when he died)

Parietal Only the left parietal is preserved which is mostly broken A single identifiable small ridge separates this element in two parts with distinct angles (Fig 4g) Medial to this ridge it is almost flat showing a gentle radial ornamentation constituted of small and irregular fossae Laterally this bone protrudes ventrally and forms the dorsal margin of the supratemporal fenestrae The posterior limit of the parietal bears a thin and incipient overhanging flange (probably by its juvenile condition) which accommodates nucal osteoderms (as seem in Aetosauroides scagliai PVL 2059 Neoaetosauroides engaeus PVL 5698 Desmatosuchus smalli TTUP 9024) a feature that characterizes aetosaurs Due to the fragmentary nature of this element most of its features are not observable

FIGURE 4 Partial braincase of Polesinesuchus aurelioi holotype (ULBRAPVT003) A dorsal view B ventral view C right

lateral view D left lateral view E posterior view and F anterior view Abbreviations bo basioccipital bt basal tuber (a) of

the basioccipital mf metotic foramen np notochordal pit oc occipital condyle XII foramen for hypoglossal nerve G Dorsal

view of a fragmentary parietal preserved in Polesinesuchus holotype (ULBRAPVT003) Abbreviations m stf dorsal margin of

the supratemporal fenestrae r ridge o ornamentation Scale bar equals 10 mm

Braincase The braincase is represented by a partial but still well-preserved basioccipital which participates in both condylar neck and occipital condyle (Fig 4andashf) It is dorsally placed to the magnum foramen as in Tecovasuchus chatterjeei (holotype TTUP 545 Martz amp Small 2006) Typothorax coccinarum (TTUP 9214 Martz 2002) Desmatosuchus smalli (Small 2002) and Neoaetosauroides engaeus (PVL 5698) The articular facet of the occipital condyle is slightly projected dorsoventrally as in Calyptosuchus wellesi (PEFO 34516) In posterior view the occipital condyle is semicircular with a shallow notochordal pit in the center of its articular surface a condition also observed in Stagonolepis olenkae (Sulej 2009) Two non-contacting facets for articulation with the exoccipitals are present in the dorsolateral surface of the occipital condyle as in Longosuchus meadei (TTU 31185) and Desmatosuchus smalli (Small 2002) contrasting with the condition in Neoaetosauroides engaeus (PVL 5698) A pair of ventrally projecting knobs from the basioccipital together with the basisphenoid is present at the basis of the condylar neck and forms the basal tubera These tubera serve as areas to attach subvertebral muscles (Romer 1956 Desojo amp Baeacutez 2007) Despite the non-preservation of the exoccipitals the basioccipital presents two

channels in dorsal view The first one forms part to the metotic foramen which encloses nerves IX X XI and jugular vein as seen in Tecovasuchus chatterjeei (Martz amp Small 2006) Posteriorly the basioccipital also partially compose the ventral margin of the foramen for the hypoglossal nerve (XII)

Cervical vertebrae Six cervical elements including the centrum of the axis are preserved Based on the information about the position of their parapophysis the degree of prominence of their ventral keels and the size of the vertebral centra their placement along the axial skeleton was tentatively assigned The cervical segment is constituted by the axis and probably third fourth fifth seventh and eighth cervical vertebrae Thus two cervical elements are not preserved in the series as in aetosaurs nine cervical vertebrae are considered to be present (Walker 1961 Parker 2008) On the other hand a row of seven cervical paramedian osteoderms is present in the cervical region of Aetosaurus ferratus (SMNS 5770 S4 S7 S8 S16 S19) if each paramedian osteoderm is correlated with only one underling vertebra

FIGURE 5 Axis of the Polesinesuchus aurelioi holotype (ULBRAPVT003) A anterior view B posterior view C left lateral

view D right lateral view E dorsal view and F ventral view Abbreviations op odontoid process vk ventral keel Scale bar

equals 10mm

Axis This element is poorly preserved comprising only the centrum which is mostly constricted along its length The anterior facet of articulation is comparatively more distinguishable than the posterior one being ldquoUrdquo-shaped It is also possible to observe a prominent and dorsally displaced odontoid process (Fig 5) Its posterior facet is circular and concave In lateral view the ventral surface of the centrum is shorter than the dorsal one a small parapophysis almost ventrally displaced is also present In ventral view a conspicuous ventral keel is present as occurs in Aetosauroides scagliai (PVL 2059 MCP13andashb-PV) Neoaetosauroides engaeus (PVL 5698) Longosuchus meadei (TMM 31185) Stagonolepis robertsoni and Typothorax coccinarum (Long amp Murry 1995 fig 102c Desojo et al 2012) In contrast this feature is absent in Aetobarbakinoides brasiliensis (CPE2 168) and Desmatosuchus spurensis (Parker 2008) In lateral view the axis of Polesinesuchus does not present the well-

rimmed fossa observable in cervical vertebrae of A scagliai (PVL 2059 MCP13andashb-PV) (Desojo amp Ezcurra 2011) This absence is shared with Aetobarbakinoides brasiliensis (CPE2 168) and Longosuchus meadei (TMM 31185) Thus the presence of a ventral keel and the absence of well-rimmed fossa are characters extended to all preserved cervical elements of Polesinesuchus

FIGURE 6 Cervical vertebra (C3) of the Polesinesuchus aurelioi (ULBRAPVT003) in A anterior view B posterior view

C left lateral view D right lateral view E dorsal view and F ventral view Abbreviations dp diapophysis nc neural canal

ns neural spine pp parapophysis prz prezygapophysis psf post spinal fossa vk ventral keel Scale bar equals 10mm

FIGURE 7 Cervical centrum (C4 AndashF) of the Polesinesuchus aurelioi holotype (ULBRAPVT003) in A anterior view B

posterior view C right lateral view D left lateral view E dorsal view and F ventral view Cervical vertebra (C5 GndashL) in G

anterior view H posterior view I left lateral view J right lateral view K dorsal view and L ventral view Abbreviations dp

diapophysis nc neural canal pp parapophysis prz prezygapophysis vk ventral keel Scale bar equals 10mm

Postaxial cervical vertebrae The third () cervical element is almost complete preserving both neural arch and spines (Fig 6) Two neural pedicles are preserved both higher than the centrum and laterally projected a condition shared with Aetobarbakinoides brasiliensis (CPE2 168) In lateral and medial views they present a slight concavity as in Longosuchus meadei (TMM 31185) The neural canal presents the same ratio and size as Aetobarbakinoides (its height is two times lower than the height of the anterior articular surface of the centrum) The transverse process is short and lateroventrally projected and its diapophysis is oval-shaped The infradiapophyseal lamina is absent in cervical elements of the holotype as in Neoaetosauroides (PVL 3525) and Longosuchus meadei (TMM 31185) Conversely in Aetobarbakinoides brasiliensis (CPE2 168) and Aetosauroides

scagliai (PVL 2059) the infradiapophyseal lamina is present (Desojo et al 2012) In Polesinesuchus the absence of both postzygapophyseal lamina and infradiapophyseal fossa are shared with Desmatosuchus spurensis (Parker 2008 Desojo et al 2012) Only the right prezygapophysis is preserved in the holotype of Polesinesuchus Its articulation facet is dorsally projected and medially restricted not exceeding the anterior edge of the diapophysis as in Aetosauroides scagliai (PVL 2059) Neoaetosauroides engaeus (PVL 5698) Typothorax coccinarum (TTU P-9214) Desmatosuchus spurensis (Parker 2008) and Stagonolepis robertsoni (NHMUK 4784) Conversely Aetobarbakinoides brasiliensis presents laterally projected prezygapophyses (CPE2 168) In Polesinesuchus a small oval-shaped fossa is visible between the prezygapophysis This fossa is known as the prespinal fossa which is not dorsally extended along the neural spine a condition similar to that found in Aetobarbakinoides (CPE2 168) Unfortunately none of the postzygapophyses are preserved in the cervical vertebrae of the holotype Its centrum is amphicoelous with a smooth lateromedial constriction Its length is longer than the height of the anterior facet with an lengthheight ratio of 12 This condition is similar to that presented in Aetosauroides scagliai (PVL 2059) (Fig 6) In Polesinesuchus the anterior facet of the centrum is wider than the posterior one (12 times) but in a comparative smaller degree regarding to Aetobarbakinoides (135 times) Desmatosuchus spurensis (Parker 2003 2008) and Typothorax (Martz 2002) also present a smaller degree regarding this ratio (Parker 2008 Desojo et al 2012) The lateral surface of the cervical centrum is slightly concave and lacks a well-rimmed fossa It is thus distinct from Aetosauroides scagliai (PVL 2059) and Longosuchus meadei (TMM 31185) The absence of a well- rimmed fossa is a condition shared with Aetobarbakinoides brasiliensis (CPE2 168) The parapophysis is situated in the ventralmost portion of the anterior margin of the centrum and placed in a low pedicle a similar condition to that found in Aetobarbakinoides (Desojo et al 2012) Additionally in ventral view Polesinesuchus presents a sharp ventral keel as in Aetosauroides scagliai (PVL 2059 MCP 13-andashb-PV) Stagonolepis robertsoni (Walker 1961 fig 7f) Typothorax (Long amp Murry 1995 102c) and Neoaetosauroides engaeus (PVL 5698) (Desojo et al 2012) Conversely in Aetobarbakinoides brasiliensis (CPE2 168) Longosuchus meadei (TMM 31185) and Desmatosuchus spurensis the ventral keel is absent (Parker 2008) The fourth () cervical element is represented only by a centrum (Fig 7andashf) In the fifth () cervical element the set of features is the same of the third one including the position of the parapophysis It comprises the centrum and the left neural pedicle (Fig 7gndashl) Only the prezygapophysis is preserved

The seventh () cervical vertebra preserves the centrum and also the left neural pedicle (Fig 8andashf) It also shows a ventrally projected transverse process longer than the one of the third vertebra Its parapophyses are dorsally displaced and close to the neurocentral suture The eighth vertebra only preserves the centrum that presents the same characteristics already mentioned for the previous elements Additionally it is possible to observe a diagenetic distortion in the right surface of the centrum which makes it more swollen than the parapophysis in comparison (Fig 8gndashl) In general the cervical centra of Polesinesuchus are robust with oval-shaped articular facets In contrast Desmatosuchus spurensis presents subrectangular facets on its centra (Parker 2008) Moreover the lengthheight ratio of the all cervical centra of the Polesinesuchus is about 12

Dorsal vertebrae The first dorsal vertebra presents the left neural arch pedicle and preserves its pre- and postzygapophyses transverse process and centrum (Fig 9andashf) In lateral view this element is anteroposteriorly shorter than the subsequent one (the more posteriorly placed dorsal vertebrae) Its centrum is amphicoelous laterally compressed and spool-shaped as in several aetosaurs such as Desmatosuchus spurensis and Calyptosuchus wellesi (Long amp Murry 1995 Parker 2008) Neoaetosauroides engaeus (PVL 3525) Aetosauroides

scagliai (Desojo amp Ezcurra 2011) The centrum of Polesinesuchus is longer than the height of its anterior articular facet with a ratio of approximately 17 resembling the condition found in both anterior and medial dorsal vertebrae of Aetosauroides scagliai (15 in PVL 2073) anterior dorsal of Stagonolepis robertsoni (145 in NHMUK

R4784 Walker 1961 fig 7i) and also of Sierritasuchus (146 in Parker et al 2008 fig 2g) (Desojo et al 2012) The parapophyses are present and dorsally displaced close to the neurocentral suture Additionally a well-rimmed fossa is absent in the lateral surface of the centrum the same condition of Aetobarbakinoides brasiliensis (Desojo et al 2012) and Desmatosuchus spurensis (Parker 2008) In contrast in Aetosauroides scagliai this structure is present (eg PVL 2073 PVL 2059 and PVL 2052) In ventral view the centrum is devoid of a ventral keel resembling the condition of Neoaetosauroides engaeus (PVL 3525) Aetosauroides scagliai Desmatosuchus

spurensis and Longosuchus meadei (Long amp Murry 1995 Parker 2008) The infradiapophyseal lamina that occurs in Aetosauroides is absent in the first dorsal vertebra of Polesinesuchus and Aetobarbakinoides (Desojo et al 2012) Its absence is shared with Desmatosuchus spurensis (Parker 2008) The transverse process is shorter than that from the middle dorsal ones but heavier showing a slightly anterior displacement in relation to the center of the neural arch a condition shared with Aetosauroides (MCP 13andashb-PV) (Desojo amp Ezcurra 2011) In posterior view the transverse process possesses a sharp postzygapophyseal lamina between the diapophysis and the postzygapophysis This lamina is also present in dorsal vertebrae of Aetobarbakinoides and Desmatosuchus

(Parker 2008 Desojo et al 2012) The diapophyses are suboval in shape and its long axis is anteroposteriorly placed The parapophysis is situated at the centrum just below the neural suture as seen in Desmatosuchus

spurensis (MNA V9300 Parker 2008 fig 8C) The parapophysis is poorly pronounced and presents an oval form In contrast in Desmatosuchus spurensis this structure covers almost a quarter of the lateral side of the centrum (Parker 2008) Four disarticulated centra probably proceeding from the anterior dorsal series are present These elements do not possess any distinctive difference from each other (Figs 9gndashl 10) presenting the same characteristics of the other centra The middle dorsal series is represented by three elements in which most structures are preserved (probably thirteenth fourteenth and fifteenth vertebrae) (Figs 11 12) Their position in the dorsal series is tentative based upon the location of their parapophyses as in this region of the column they are placed on the transverse process This feature is reported in all aetosaurs (Parker 2008)

FIGURE 8 Cervical vertebrae (C7 AndashF and C8 GndashL) the Polesinesuchus aurelioi holotype (ULBRAPVT003) in A anterior

view B posterior view C left lateral view D right lateral view E dorsal view and F ventral view In G anterior view H

posterior view I right lateral view J left lateral view K dorsal view and L ventral view Abbreviations dp diapophysis nc

neural canal pp parapophysis prz prezygapophysis vk ventral keel Scale bar equals 10mm

The better preserved neural spine belongs to the putative 13th dorsal vertebra (Fig 11) It is thin laterally and anteroposteriorly elongated as in Desmatosuchus spurensis (Case 1922 Parker 2008) Its apex is partially broken but it is possible to observe a well-marked lateral expansion of the neural spine This expansion could be the so-called ldquospine tablerdquo typical of aetosaurs as in Desmatosuchus spurensis (MNA V9300 Parker 2008 fig 8) Both transverse processes are partially broken (Fig 11) In dorsal view the transverse process is trapezoidal and laterally

projected In lateral view it is only possible to observe the diapophysis on both sides The infradiapophyseal lamina is completely absent in dorsal vertebrae of Polesinesuchus In contrast these laminae are present in the middle dorsal axial elements of Aetosauroides (PVL 2073) Stagonolepis robertsoni (NHMUK R4784) Desmatosuchus

spurensis (Parker 2008) and Typothorax coccinarum (Martz 2002) (Desojo amp Ezcurra 2011 Desojo et al 2012) This condition of Polesinesuchus is shared with Neoaetosauroides (PVL 3525) and the Stagonolepis olenkae

(ZPAL AbIII 50267) (Desojo amp Ezcurra 2011) Furthermore the infradiapophyseal fossa is absent in dorsal vertebrae of Polesinesuchus as in Aetosauroides (MCP 13andashbPV) (Desojo amp Ezcurra 2011) The neural spine is wide and posteriorly placed Unfortunately the apex of the neural spine is broken and its relationship with the centrum cannot be established

FIGURE 9 Dorsal vertebrae of the Polesinesuchus aurelioi holotype (ULBRAPVT003) D1 AndashF and dorsal centrum

probably between the first and thirteenth dorsal vertebrae GndashL) in A anterior view B posterior view C right lateral view D

left lateral view E dorsal view and F ventral view In G anterior view H posterior view I right lateral view J left lateral

view K dorsal view and L ventral view Abbreviations dp diapophysis nc neural canal pp parapophysis prz

prezygapophysis poz postzygapophysis pp parapophysis Scale bar equals 10mm

FIGURE 10 Dorsal centra of the Polesinesuchus aurelioi holotype (ULBRAPVT003) (probably mid dorsal vertebrae) in A

anterior view B posterior view C right lateral view D left lateral view E dorsal view and F ventral view In G anterior

view H posterior view I right lateral view J left lateral view K dorsal view and L ventral view In M anterior view N

posterior view O right lateral view P left lateral view Q dorsal view and R ventral view Scale bar equals 10mm

FIGURE 11 The thirteenth () dorsal vertebra of the Polesinesuchus aurelioi holotype (ULBRAPVT003) in A anterior view

B posterior view C right lateral view D left lateral view E dorsal view and F ventral view Abbreviations dp diapophysis

nc neural canal ns neural spine pp parapophysis poz postzygapophysis prz prezygapophysis psf post spinal fossa Scale

bar equals 10mm

FIGURE 12 The fourteenth () dorsal vertebra of the Polesinesuchus aurelioi holotype (ULBRAPVT003) in A anterior view

B posterior view C right lateral view D left lateral view E dorsal view and F ventral view The fifteenth () vertebra of the

Polesinesuchus aurelioi holotype In G anterior view H posterior view I right lateral view J left lateral view K dorsal view

and L ventral view Abbreviations dp diapophysis nc neural canal ns neural spine prz prezygapophysis pp parapophysis

Scale bar equals 10mm

The prezygapophyses are anteroposteriorly short and dorsally oriented as in Aetosauroides (PVL 2073 MCP 13andashb PV) Stagonolepis robertsoni (Walker 1961) and Typothorax coccinarum (Martz 2002 Desojo amp Ezcurra 2011) The longer axes of the prezygapophyses are lateromedial Only the left postzygapophysis is preserved in the thirteenth vertebra The hyposphene is absent in Polesinesuchus the same condition found in Aetosauroides In contrast this structure is present in Aetobarbakinoides and Desmatosuchus (Parker 2008 Desojo et al 2012) The postzygapophyseal lamina is present in the holotype Aetobarbakinoides and Aetosauroides (see Desojo amp Ezcurra 2011 Desojo et al 2012) Additionally a circular pit (located in both sides of the neural spine and present in Aetobarbakinoides) is absent in Polesinesuchus and Aetosauroides (Desojo et al 2012) Regarding the remaining dorsal vertebrae their centra are amphicoelous and anteroposteriorly elongated longer than the height of the anterior facet with an lengthheight ratio of 142 (in Aetosauroides 15 PVL2073) in the anterior dorsal of Stagonolepis robertsoni 145 NHMUK R4784 Walker 1961 and in Sierritasuchus 146 Parker et al 2008 fig 2g) (Desojo et al 2012)

The fourteenth vertebra comprises a centrum and a partial neural arch including the base of the left transverse process (Fig 12andashf) Only the left parapophysis and left prezygapophysis are preserved The articular facet of the parapophysis is oval-shaped and placed on the transverse process The prezygapophysis is similar to that one from the first dorsal vertebra The centrum is amphicoelous without a well-rimmed fossa In dorsal view the centrum is compressed (spool-shaped) as in Aetosauroides (MCP 13andashb PV) (Desojo amp Ezcurra 2011) Its length is comparatively longer than the height of the anterior articular facet (lengthheight ratio = 133)

FIGURE 13 The first sacral vertebra of Polesinesuchus aurelioi holotype (ULBRAPVT003) in A anterior view B posterior

view C right lateral view D left lateral view E dorsal view and F ventral view The second sacral vertebra of Polesinesuchus

aurelioi holotype in G anterior view H posterior view I right lateral view J left lateral view K dorsal view and L ventral

view Scale bar equals 10mm

The fifteenth vertebra comprises a centrum with an open neurocentral suture the base of the neural spine a complete left transverse process the diapophysis and the parapophysis (Fig 12gndashl) The diapophysis is oval-shaped and the parapophysis is anteriorly placed In dorsal view the transverse process is trapezoidal with an anteroposteriorly longer proximal end as in Aetosauroides (PVL 2073 MCP 13andashb PV) The infradiapophyseal lamina is absent and the parapophysis is placed over the transverse process reaching half of its length As in many other centra described above the centrum is longer than the height of the anterior articular facet with a lengthheight ratio of the 136 This element is amphicoelous and transversely compressed Laterally the well-rimmed fossa is absent

FIGURE 14 Caudal vertebrae of Polesinesuchus aurelioi holotype (ULBRAPVT003) in A anterior view B posterior view

C right lateral view D left lateral view E dorsal view and F ventral view In G anterior view H posterior view I right lateral

view J left lateral view K dorsal view and L ventral view Scale bar equals 10mm

Sacral vertebrae The sacral vertebrae comprise two disarticulated centra both amphicoelous and considerably more robust than those described for the dorsal series as seen in Desmatosuchus spurensis (Parker 2008) Stagonolepis robertsoni and Lucasuchus hunti (Walker 1961 Long amp Murry 1995) (Fig 13) Both anterior and posterior sides are wider than tall as observable in sacral centra of Desmatosuchus supurensis Lucasuchus

hunti and Stagonolepis robertsoni (Walker 1961 Long amp Murry 1995 Parker 2008) These vertebrae possess slightly flattened anterior and posterior sides as in Typothorax coccinarum (TTUP 9214) and phytosaurs (Martz 2002) In lateral view it is possible to observe both areas for articulation of the neural arches almost reaching the midline of the centrum as in Typothorax and Lucasuchus hunti (Long amp Murry 1995 Martz 2002) However in both sacral centra these structures cover the dorsal half of the length of the centrum A well-rimmed fossa is absent in Polesinesuchus as in other sacral vertebrae of Aetosauroides scagliai (PVL 2073 PVL 2052) Neoaetosauroides

engaeus (PVL 3525) Stagonolepis robertsoni (Walker 1961) In ventral view both sacral centra are flattened and very broad as occurs in Desmatosuchus spurensis (Parker 2008) and other aetosaurs (eg Aetosauroides scagliai

PVL 2073 Stagonolepis robertsoni)Caudal vertebrae Five disarticulated centra are preserved in the holotype The first two are amphicoelous

and longer than the height of the anterior articular facet (lengthheight ratio 116) (Fig 14) These elements are quite robust with oval-shaped articular facets These facets form a thickened rim as in Desmatosuchus spurensis

(Parker 2008) In ventral view it is possible to observe a groove that is laterally limited by two ridges which anteriorly end in chevron facets as in Stagonolepis robertsoni (Walker 1961) In Polesinesuchus the ridges and grooves cover only the posterior half of the centrum In contrast in the anterior caudal element of Desmatosuchus

spurensis the ridges extend throughout its entire length (Parker 2008) Laterally the two first caudal centra are devoid of a lateral fossa Two other centra belong to the middle caudal region These elements present some degree of distortion and their lateral surfaces are broken (Fig 15) The centra are longer than the anterior ones with a length longer than the anterior articular facet (lengthheight ratio 131) In ventral view they present ridges with a groove in the anterior half of the centrum As in the other caudal elements lateral fossae are absent The distal caudal region is represented by a single centrum (Fig 16mndashr) This element is smaller when compared to the other caudal elements but with the same morphology It is longer than the anterior articular facet (length height ratio 233)

view J left lateral view K dorsal view and L ventral view In M anterior view N posterior view O right lateral view P left

lateral view Q dorsal view and R ventral Scale bar equals 10mm

Shoulder Girdle Both scapulae and coracoids of Polesinesuchus are preserved The left elements are more complete than the right ones (Figs 16 17) Both proximal and distal ends of the left scapula as well as the scapular blade are well-preserved In contrast in the right scapular blade both proximal ends and anterior margin are broken The coracoids present comparatively different states of preservation The right coracoid is broken The left element is more complete allowing its observation in detail As usual for aetosaurs the scapula is a slender element with a slight constriction along its center and expanded extremities In lateral view this bone is wide and slightly convex Its anterior margin is in comparison sharper than the posterior The posterior margin is thicker than the anterior one Dorsoventrally the scapular blade is thin presenting a large anteroposterior expansion to support the suprascapular cartilage which possibly becomes ossified in adult individuals (Walker 1961) The medial portion of the scapular blade is more expanded in comparison with of the condition found in Aetosauroides

(PVL 2073) The ventral expansion presents a marked acromial process It is thick and laterally placed as in Aetosauroides In contrast in Aetobarbakinoides (PVL 2073) Aetosaurus (SMNS 5770 S-2) and Neoaetosauroides (PVL 3525) this structure is less prominent (Desojo et al 2012) According to Romer (1956) the articular process would serve as a point of attachment with the clavicle Distally the subacromial tuberosity delimits a shallow anteroposterior groove as in Aetosauroides Conversely in Aetobarbakinoides and Neoaetosauroides the subacromial tuberosity delimits a moderately deep subacromial depression (Desojo et al 2012) Distally the scapula shows an articular facet for accommodation of the coracoids This articulation is lateromedially expanded towards dorsal region Anteriorly it is slender as in some basal archosaurs such as Prestosuchus chiniquensis (Huene 1938) and Rauisuchus tiradentes (Lautenschlager 2008) The glenoid fossa is dorsolaterally projected as in Aetobarbakinoides (Desojo et al 2012) Alongside the glenoid fossa there is a small process for the attachment of the triceps muscle as in Batrachotomus kupferzellensis (Gower amp Schoch 2009)

FIGURE 16 Left scapula and coracoid of Polesinesuchus aurelioi holotype (ULBRAPVT003) in A lateral view B medial

view C posterior view and D anterior view E Left coracoid in lateral view and in F left coracoid in medial view

Abbreviations ap acromial process cf coracoid foramen gf glenoid fossa sgl supra glenoid lip Scale bar equals 10mm

Coracoid The coracoids are laterally convex medially concave and longer than wide as in Aetosauroides

(PVL 2073) (Casamiquela 1961) Proximally they are thick as they attach to the scapulae to form most of the glenoid resembling Aetosauroides (PVL 2073) (Casamiquela 1961) Their glenoid shows a buttressed glenoid lip as occurs in Typothorax coccinarum (Heckert et al 2010) Posteriorly these elements are also thick and rounded and as usual there is a neck below the glenoid Dorsally a posterior process is present The coracoid foramen is quite visible in Polesinesuchus This foramen is circular and relatively large In contrast Neoaetosauroides

possesses a small foramen In Polesinesuchus it is situated 78mm from the glenoid (Figs 16 17) The subglenoid pillar present in some aetosaurs such as Typothorax coccinarum (Heckert et al 2010) Stagonolepis robertsoni

(Walker 1961 fig 12) Neoaetosauroides Lucasuchus Desmatosuchus Coahomasuchus and Aetosaurus

(Bonaparte 1971 Long amp Murry 1995) does not occur in Polesinesuchus a condition shared with Aetosauroides

scagliai (Casamiquela 1961)

FIGURE 17 Right scapula and coracoid Polesinesuchus aurelioi holotype (ULBRAPVT003) in A lateral view B medial

view C posterior view and D anterior view E right coracoid in lateral view and F right coracoid in medial view

Interclavicle This blade-like shaped element is disarticulated from the coracoids (Fig 18) It is an elongate element which is very thin anteriorly and becomes thick posteriorly forming a marked bulge as seen in Typothorax

coccinarum (Heckert et al 2010) Ventrally it is convex with its anterior end possessing two small articular facets separated by a tall crest as occurs in Aetosauroides scagliai (PVL 2073) and Stagonolepis robertsoni (Casamiquela 1961 Walker 1961) Walker (1961) described the articular facets as anterolateral pits to receive the medial process from the clavicles This element is anteroposteriorly expanded In Polesinesuchus its posterior end is well ossified and partially broken with a total length of 445 mm in comparison to Aetosauroides (PVL 2073 64mm) (Schoch 2007)

Humerus A fragmentary right humerus and just parts of the left one were recovered for Polesinesuchus The right element presents transversal fractures (close to its distal extremity) across the diaphysis The left element comprises both proximal and distal epiphyses Its diaphysis is missing (Figs 19 20) The right humerus is long and relatively robust with both proximal and distal ends transversally expanded and a robust diaphysis It measures about 12 times the length of the ulna (this ratio resembles that of Aetosaurus ferratus) (Schoch 2007) In contrast

in Aetosauroides (PVL 2073) Aetobarbakinoides Desmatosuchus haplocerus (UCMP A 26932168 Long amp Murry 1995) and Typothorax coccinarum (UCMP V 281634240 Long amp Murry 1995) the humeral diaphysis is a lot narrower than the two expanded ends of this bone The humeral head is prominent and presents a convex articular surface as in Aetobarbakinoides (Desojo et al 2012) The larger tuberosity is poorly defined as occurs in Aetosauroides (PVL 2052 2073 PVSJ 326) Neoaetosauroides (PVL 3525) Stagonolepis robertsoni (Walker 1961) and Typothorax (UCMP V 281634240 Long amp Murry 1995 Martz 2002) This structure is comparatively more conspicuous in posterior view as in Typothorax and other basal archosaurs (UCMP V 281634240 Long amp Murry 1995 Martz 2002) A prominent large tuberosity is present in Aetobarbakinoides brasiliensis and ldquoArgentinosuchus bonaparteirdquo (nomen dubium an indeterminate aetosaur according to Desojo amp Ezcurra [2011]) (Desojo et al 2012) The later authors described two tuberosities in the humeral head of Aetobarbakinoides The first one situated in the lateroventral corner and the second in the dorsal margin of the humeral head In Polesinesuchus these structures are either poorly developed or absent The deltopectoral crest is poorly developed ventrally and subtriangular in medial view This condition is shared with other pseudosuchian such as the ldquorauisuchianrdquo Batrachotomus and the aetosaur Aetosauroides scagliai (Desojo et al 2012) The distal end presents two ventrally placed condyles laterally the ectepicondyle and medially the entepicondyle These two condyles are rounded and separated by a medial groove In the dorsal surface of the distal end there is a shallow subtriangular depression as in Aetobarbakinoides brasiliensis (CPE2 168) In lateral view a distal deep groove is also present (ectepicondylar groove Schoch 2007) Along this groove there is a dorsal ridge probably to accommodate blood vessels and the radial nerve (Romer 1956) similarly to the condition present in several aetosaurs such as Aetobarbakinoides brasiliensis (CPE2 168) Aetosauroides scagliai (PVL 2073 PVSJ 326) Neoaetosauroides engaeus (PVL 3525) Stagonolepis robertsoni (Walker 1961) (Desojo et al 2012) and contrasting with a foramenfossa in Typothorax coccinarum (Heckert et al 2010) and Desmatosuchus haplocerus

(TTU 9170 Small 2002)Ulna Two well-preserved ulnae are present in Polesinesuchus both presenting a slight fracture in their shafts

(Fig 21) They are lateromedially compressed with a slightly expanded and anteroposteriorly twisted proximal end which is wider than the distal one in comparison as in Neoaetosauroides engaeus (Desojo amp Baeacutez 2005) and Longosuchus meadei (Sawin 1947) Its olecranon is poorly-developed In contrast this structure is well-developed in Coahomasuchus kahleorum (Heckert amp Lucas 1999) Aetosauroides scagliai (PVL 2073) (Casamiquela 1961 fig 11ndash13) Neoaetosauroides engaeus (Desojo amp Baeacutez 2005) Longosuchus meadei (Sawin 1947) and Aetosaurus

(Schoch 2007) It is probable that the olecranon of Polesinesuchus olecranon would be filled with cartilage which is suggested by its poorly observable prominence Also it presents a gentle angle contrasting with the L-shaped one observed in Typothorax coccinarum (Heckert et al 2010) In ventral view the articular surface is oval-shaped as in Typothorax coccinarum (Heckert et al 2010)

FIGURE 18 Interclavicle of Polesinesuchus aurelioi holotype (ULBRAPVT003) in A dorsal view B ventral view and CndashD

lateral views Scale bar equals 10mm

FIGURE 19 Right humerus of Polesinesuchus aurelioi holotype (ULBRAPVT003) in A dorsal view B ventral view C

lateral view and D medial view E proximal and F distal views Abbreviations dpc deltopectoral crest enc entepictondyle

ect ectepicondyle gt greater trochanter Scale bar equals 10mm

Ilium Only the left ilium is preserved in Polesinesuchus Its iliac blade is long dorsoventrally short with two processes (one anterior and other posterior) Its outline resembles that of Desmatosuchus haplocerus (UMMP 73322) (Long amp Murry 1995) and Desmatosuchus spurensis (Parker 2008 fig 6C) (Fig 22) In lateral view the blade is proportionally lower than the iliac blade of Aetosaurus ferratus (Schoch 2007 S-20) Typothorax

coccinarum (UCMP V2816 122683 70Fa54) and Stagonolepis robertsoni (Walker 1961 Martz 2002 Schoch 2007) In Typothorax coccinarum the iliac blade is tall and shows in both sides two transversal buttresses above the acetabulum that is expressed as a strong thickening (Heckert et al 2010) This feature does not occur in the iliac blade of Polesinesuchus The anterior process of the iliac blade is short slightly exceeding the pubic peduncle a condition shared with Neoaetosauroides engaeus (PVL 3525) Longosuchus meadei (TMM 31185) and Desmatosuchus spurensis The tip of the anterior process is very slender with an oblique anteroventral direction as

in Aetosaurus ferratus and Desmatosuchus haplocerus (Long amp Murry 1995 Schoch 2007) and longer than in Aetosauroides scagliai (PVL 2073) Additionally the tip of the anterior process is also very pointed when compared with the rounded one present in Aetosauroides scagliai (PVL 2073 PVL 2052) The anterior process of the iliac blade serves as the insertion area for iliotibialis and iliofemurales muscles both directly related to the movement of hindlimbs (Desojo amp Baeacutez 2005) The posterior process is long and massive In dorsal view the iliac blade presents a ripple in its distal end Additionally the posterior process is mediolaterally inclined differently from the anterior process This condition is shared with Typothorax coccinarum (Heckert et al 2010) The acetabulum is a non-perforated shallow concavity It is almost entirely formed by the ilium and delimited by a supracetabular crest that forms an overhanging ridge to accommodate the head of the femur as occurs in Desmatosuchus spurensis (Parker 2008) Ventrally the ilium expands and ends in two articular surfaces

FIGURE 20 Left humerus of Polesinesuchus aurelioi holotype (ULBRAPVT003) in A dorsal view B ventral view C

ect ectepicondyle ect g ectepicondylar groove gt greater trochanter Scale bar equals 10mm

FIGURE 21 Ulnae of Polesinesuchus aurelioi holotype (ULBRAPVT003) AndashB right ulna in lateral and medial views CndashD

left ulna in lateral and medial views Abbreviation op olecranon process Scale bar equals 10mm

FIGURE 22 Left ilium of Polesinesuchus aurelioi holotype (ULBRAPVT003) in A dorsal view B ventral view C lateral

view and D medial view Abbreviations ap anterior process ac acetabulum ptp posterior process sac supra acetabular

crest Scale bar equals 10mm

FIGURE 23 Left pubis of Polesinesuchus aurelioi holotype (ULBRAPVT003) in A medial view B lateral view C dorsal

view In a fragment of right pubis in D medial E lateral and F dorsal views Left isquium in G lateral H medial and dorsal

views Abbreviation t tongue Scale bar equals 10mm

Pubis Represented by fragments of both pubial elements (Fig 23) The left pubis preserves both proximal and distal ends so the medial region is missing The right pubis only preserves its proximal end Walker (1961) described two foramina in the pubis of Stagonolepis robertsoni but these structures are not visible in Polesinesuchus as most of the aetosaurs The proximal end presents the characteristic twist a condition present in several pseudosuchian archosaurs (Walker 1961) The pubis is a short element in which the proximal end is stronger than the distal one in comparison which presents a massive rugosity A short pubis is also present in Aetosaurus ferratus (Schoch 2007) In contrast a longer pubis is present in Stagonolepis robertsoni and Desmatosuchus spurensis (Walker 1961 Schoch 2007 Parker 2008) The left proximal end is comparatively better preserved and allows the recognition of some structures For instance the articular surface for the ilium is almost flat and ldquoUrdquo shaped Anteroposteriorly the articular surface is long as occurs in Neoaetosauroides engaeus and other basal archosaurs (Lecuona amp Desojo 2011) Medially there is a bony tongue with a groove over its sagital length This structure is a thin proximal portion of bone for the obturador foramen which in Polesinesuchus is

difficult to observe due to its poor preservation This tongue-like structure is reported only for Postosuchus

kirkpatricki Chatterjee 1985 (Long amp Murry 1995 fig 135) and Gracilisuchus stipanicicorum Romer 1972 (Lecuona amp Desojo 2011 fig 4) Parts of the proximal end of the pubis are broken but there is some evidence of the presence of the obturator foramen A portion of the distal end of pubis is preserved in the left element It is constituted by a very thin lamina which becomes thicker distally to form a prominent pubic foot Distally the pubic foot of Polesinesuchus shows a marked rugosity as occurs in Typothorax coccinarum and Stagonolepis robertsoni

(Walker 1961 Martz 2002) Additionally Parker (2008) described (for the distal end) a club-like end in Desmatosuchus spuresis but stated that this structure is probably homologous to the pubic foot of other suchiansThis feature is usual in aetosaurs phytosaurs and sphenosuchians (Gower amp Schoch 2009)

Ischium Only the left ischium is preserved in Polesinesuchus in which most structures are preserved (Fig 23) It is L-shaped with the proximal end wider than the distal one in comparison The proximal end is also thicker than distal possessing two articular surfaces one medial and other lateral The medial articular surface is almost flat smaller in comparison and articulates with the posteroventral articular surface of the ilium The lateral articular surface contributes to the ventral margin of acetabulum It is larger and almost oval-shaped The anterior margin of the ischium is very thin and it is partially broken in Polesinesuchus It forms the puboischiatic plate Distally the ischium becomes posteriorly expanded and shows a curvature medial to the symphysis Apparently the symphysis is distributed along the length of the ischial shaft as in Aetosauroides scagliai (PVL 2073 PVL 2052) Typothorax coccinarum Stagonolepis robertsoni and many aetosaurs and pseudosuchian archosaurs as well (Walker 1961 Heckert et al 2010 Lecuona amp Desojo 2011 Nesbitt 2011)

Femur Both femora are present and well-preserved in the holotype They are robust with a typical sigmoid shape as in Aetosauroides scagliai (PVL 2073) and most archosaurs (Nesbitt 2011) (Figs 24 25) However in the femora of Polesinesuchus the extremities are slightly twisted along the axial length in comparison with those of Aetosauroides scagliai (PVL 2073) The distal end is anteroposteriorly twisted counterclockwise (to the left) in comparison to the proximal end Additionally the femoral shaft is oval in cross section as seen in Typothorax

coccinarum Stagonolepis robertsoni and Aetosaurus ferratus (SMNS 57771) (Walker 1961 Schoch 2007 Heckert et al 2010) Probably this feature is related to the type of insertion of the femur due to the ventral position of the acetabulum (Desojo amp Baeacutez 2005) The femoral head is medially displaced for connection to the acetabulum The greater trochanter is slightly developed Medially and below the greater trochanter there is a strong ridge that ends almost over the fourth trochanter This ridge is accompanied by a similar sized longitudinal concave area In the proximal femoral lateral surface there is another ridge but low and poorly-developed in comparison In dorsal view the proximal end of the femur presents in its articular surface an anteroposterior groove The fourth trochanter of Polesinesuchus is very high and forms a prominent crest as reported in Aetosauroides scagliai and Calyptosuchus wellesi In contrast the fourth trochanter of Aetosaurus ferratus presents a longitudinal bulge that forms a low platform Moreover Stenomyti huangae also show a low fourth trochanter (Schoch 2007 Martz amp Small 2013) These trochanters function as an insertion area for the M caudifemoralis

longus (Lecuona amp Desojo 2011) The distal end of the femur presents two ventral condyles The medial one articulates with the tibia and the lateral attaches with the fibula These condyles are separated by a longitudinal intercondylar groove for attachment of the M quadriceps femorallis

Tibia A complete left tibia and a proximal end of the right element are preserved Polesinesuchus both in a good degree of preservation (Fig 26) The tibia is a strong bone (Its length is 12 times the femur) with expanded extremities the proximal wider than the distal one in comparison The tibia of Aetobarbakinoides brasiliensis is gentle in comparison with that of Polesinesuchus (lengthwidth ratio = 10) The expansion of the proximal end of the tibia in aetosaurs is wider than those in non-aetosaur pseudosuchians (Parrish 1986 Martz 2002) However in Aetosaurus ferratus both extremities of the tibia present almost the same width contrasting with those of Stagonolepsis robertsoni Aetosauroides scagliai Neoaetosauroides engaeus Typothorax coccinarum and Desmatosuchus (Schoch 2007) The shaft presents a posteriorly projected curve as in Aetobarbakinoides

brasiliensis (CPE2 168) and Aetosauroides scagliai (PVL 2073) Tibia and fibula have almost the same length (12 times the femoral length) The shaft is lateromedially flattened and presents a slightly bowing towards its length Laterally the tibial shaft presents a longitudinal edge as in Aetobarbakinoides brasiliensis (CPE2 168) Dorsally the proximal extremities form two articular surfaces for articulation with the femoral condyles The distal end has an expanded and slightly elliptic outline which presents two ventral condyles both forming two divided facets for the astragalus as occurs in Typothorax coccinarum and also in other non-aetosaur pseudosuchians (Martz 2002)

FIGURE 24 Right femur of Polesinesuchus aurelioi holotype (ULBRAPVT003) in AndashB posterior and anterior views CndashD

medial and lateral view EndashF proximal and distal views Abbreviations fc condyle for fibula gt greater trochanter tc condyle

for tibia 4t fourth trochanter Scale bar equals 10mm

FIGURE 25 Left femur of Polesinesuchus aurelioi holotype (ULBRAPVT003) in AndashB posterior and anterior views CndashD

Fibula Both elements are preserved in Polesinesuchus They are deformed and damaged in some extent These elements are gentle measuring almost the same length of the tibia which attains 125 times the femur (Figs 27 28) In contrast Stagonolepis robertsoni Desmatosuchus smalli and Longosuchus meadei (lengthwidth ratio = 64) show the fibula stronger than the Polesinesuchusacutes (lengthwidth ratio = 96) (Sawin 1947 Walker 1961 Long

amp Murry 1995) Both proximal and distal ends are slightly transversely expanded and connected by a slender shaft In the anterolateral surface of the shaft there is a prominent crest-like iliofibularis trochanter which is longitudinally longer than in Aetosauroides scagliai (PVL 2073) Additionally the crest-like iliofibularis

trochanter of Polesinesuchus differs from the double trochanter of Longosuchus meadei (Sawin 1947) Additionally in Stenomyti huangae the iliofibularis trochanter is also well developed (Small amp Martz 2013) In contrast in juvenile specimens of Aetosaurus ferratus this structure is less pronounced than in all remaining aetosaurs (Schoch 2007) A huge iliofibularis trochanter is described for Desmatosuchus haplocerus (UCMP A26932392) and Typothorax coccinarum (UCMP V2816 34248 70G6) but in this last taxon it is placed almost halfway in the shaft (Long amp Murry 1995) The distal end of the fibula presents two articular facets one for the calcaneum (which is larger than the one for the astragalus) and one for the astragalus which is smaller in comparison

FIGURE 26 Left tibia of Polesinesuchus aurelioi holotype (ULBRAPVT003) in AndashB posterior and anterior views CndashD

medial and lateral views Proximal right tibia in EndashF posterior and anterior views Scale bar equals 10mm

Tarsus and pes Both proximal tarsals astragali and the right calcaneum are preserved in Polesinesuchus A few elements from the pes are preserved in the holotype two complete metatarsals (probably III and V) and fragmentary elements of other metatarsals phalanges The crurotarsal ankle joint of Polesinesuchus is typical ldquocrocodile-normalrdquo as in pseudosuchian archosaurs except in ornitosuchids (eg Sereno amp Arcucci 1990) This articulation consists in the rotary ankle joint in which the proximal tarsals move against each other with a peg-and-socket articulation (Parrish 1986 Brochu 2001)

Astragalus It is a massive bone that presents two articular facets (fibial and tibial) in its proximal region (Fig 29) These articulations are separated by a thick crest which forms an acute angle among them In Gracilisuchus

stipanicicorum these facets are divided by a sharp crest (Lecuona amp Desojo 2011) The tibial facet is subdivided into two facets by a small elevation and larger in comparison occupying almost entirely the proximal region of the bone Its anterior articular surface is strongly concave whereas the posteromedial one is almost flat These two facets diverge to form the screw-joint articulation for the tibia Diverging facets are also related in other aetosaurs such as Neoaetosauroides engaeus (PVL 3525) and Calyptosuchus wellesi (Long amp Ballew 1985 fig 82 Lecuona amp Desojo 2011) Anteriorly there is a poorly prominent astragalar hollow which does not contact the distal hollow According to Martz (2002) a small foramen located in the hollow is present in Desmatosuchus and Alligator

mississippiensis but not evident in Typothorax coccinarum Also this foramen is not observable in Polesinesuchus The distal hollow is large and articulates with metatarsals I II and the third distal tarsal In lateral view three articular surfaces for the calcaneum are present two forming the astragalar peg as in Gracilisuchus

stipanicicorum (Lecuona amp Desojo 2011) Typothorax (NMMNH P-36075 Lucas et al 2002) Stagonolepis

robertsoni (Walker 1961) Aetosauroides ( Casamiquela 1961) and Coahomasuchus (Heckert amp Lucas 1999)

FIGURE 27 Left fibula Polesinesuchus aurelioi holotype (ULBRAPVT003) in A lateral view B medial view C anterior

view and D posterior view Abbreviation tif trochanter for iliofibulares muscle Scale bar equals 10mm

FIGURE 28 Right fibula of Polesinesuchus aurelioi holotype (ULBRAPVT003) in A lateral view B medial view C

anterior view and D posterior view Abbreviation tif trochanter for iliofibulares muscle Scale bar equals 10mm

Calcaneum It is also a robust element with a wide shaft and a slight dorsoventral compression (Fig 29) The calcaneal condyle is very prominent Dorsally it presents a concave surface that forms the area that receives the fibula in its lateral half through a sliding articulation The astragalus articulates with the condyle in its medial length as occurs in Gracilisuchus stipanicicorum (Lecuona amp Desojo 2011) Ventral to the calcaneal condyle there

is a compressed tuber which serves for the attachment of the gastronemeus muscle (Martz 2002) This tuber is more prominent in Neoaetosauroides engaeus (Desojo amp Baeacutez 2005) Between the calcaneal condyle and the calcaneal tuber Polesinesuchus displays a well-marked notch as occurs in Aetosauroides scagliai (PVL 2052) and Gracilisuchus stipanicicorum (Lecuona amp Desojo 2011) In the posterior surface of the tuber of Polesinesuchus

there is the so-called shallow medial groove which holds the tendon of the M gastrocnemius externus reported in Gracilisuchus and other aetosaurs Distally the surface of the calcaneal condyle is flattened and contacts the fourth distal tarsal Below this surface there is a deep fossa in Polesinesuchus as in Typothorax coccinarum (Martz 2002) However in the former this fossa is ldquoUrdquo-shaped differently from Typothorax In lateral view other smaller oval fossa (in comparison with the distal calcaneal fossa) is present Medially the condyle of the calcaneum displays a deep socket which is dorsally supported by a protruding lip as in Typothorax coccinarum (Martz 2002) This socket receives the peg of the astragalus the typical articulation of the crocodile normal type of tarsus

FIGURE 29 Right (AndashC) left (DndashF) astragalus and left calcaneum of Polesinesuchus aurelioi holotype (ULBRAPVT003) in

A proximoanterior B distoposterior C lateral views In D proximoanterior E distoposterior F lateral views In G proximal

view H distal view I medial view Abbreviations cc calcaneal condyle cs calcaneal socket cdf calcaneal distal fossa daf

distal astragalar facet dr distal roller pag posterior astragalar groove pxcf proximal calcaneal facet s alt anterolateral

articular surface s fi articular surface for the fibula s pmt posteromedial articular surface for the fibula Scale bar equals

Metatarsal III It is a relatively long bone with expanded ends (Length 225 times the fibula) The proximal end is dorsoventrally compressed and more expanded than the distal one in comparison (Fig 30) Unfortunately the proximal end shows fractures that difficult the recognition of some structures Nonetheless in the proximal end the degree of expansion is longer than in Aetobarbakinoides brasiliensis According to Desojo et al (2012) in this taxon the degree of expansion of the proximal end is lesser than in Neoaetosauroides engaeus (PVL 3525) Aetosaurus ferratus (Schoch 2007) Stagonolepis robertsoni (Walker 1961) and Typothorax coccinarum (Heckert et al 2010) The shaft towards the distal end becomes thinner Distally the shaft presents a slight twist as seen in Postosuchus alisonae (UNC 15575) and Aetobarbakinoides (Peyer et al 2008 Desojo et al 2012)

Metatarsal V Medially the proximal articular surface forms a small condyle and its proximal end is quite expanded with the articular surface strongly concave (Fig 31) The proximal articular surface possesses small

foramina and ligament attachment marks Medially the proximal end shows a slight expansion as occurs in Aetobarbakinoides (Desojo et al 2012) In contrast Neoaetosauroides engaeus shows a typical archosauromorph hook-like shape medially in its proximal end (Desojo amp Baacuteez 2005) Additionally in Aetosaurus (Schoch 2007) Stagonolepis (Walker 1961) and Typothorax (Heckert et al 2010) this structure is also prominent (Desojo et al 2012) In Aetosaurus this expansion of the proximal end is smaller than in Stagonolepis (Walker 1961 Schoch 2007) The distal end is less expanded than the proximal one contrasting with the distal end of the Aetobarbakinoides in which the distal end is more transversely expanded and robust than the proximal one (Desojo et al 2012) The lateral surface is also similar to that of many archosauromorphs where the shaft is compressed and tall This characteristic is exemplified in Typothorax coccinarum (UCMP V281634255 Long amp Murry 1995 NMMNH P-56299 Heckert et al 2010)

FIGURE 30 Metatarsal III of Polesinesuchus aurelioi holotype (ULBRAPVT003) in A dorsal view and B ventral view

FIGURE 31 Metatarsal V of Polesinesuchus aurelioi holotype (ULBRAPVT003) in A dorsal and B ventral views Scale bar

equals 10mm

Phalanges Two phalanges are present In both the proximal and distal ends are wide (Fig 32) The first phalanx (Fig 32a) is longer in comparison with the second presenting a dorsoventrally compressed distal end Laterally there are small pits in both sides The anterior articular surface is trapezoidal and more expanded in comparison The second element (Fig 32b) is shorter than the first one robust and resembles the first element of the fourth digit of the Aetobarbakinoides (Desojo et al 2012) Its distal end is dorsoventrally higher and the pits are larger in comparison Furthermore this element shows the typical developed ginglymous articulation in archosaurs According to these features it is quite possible to infer that this element proceeds from the four digit of the pes (see Desojo et al 2012)

FIGURE 32 Indeterminate phalanges of Polesinesuchus aurelioi holotype (ULBRAPVT003) in A dorsal B ventral views C

dorsal D ventral views Ungual phalanges in E medial F lateral views G lateral view Scale bar equals 10mm

FIGURE 33 Left cervical paramedian osteoderm (A) right paramedian osteoderm (B) and (C) right posterior osteoderm of

Polesinesuchus aurelioi holotype (ULBRAPVT003) in A dorsal view B anterior view and C posterior view Abbreviations

ab anterior bar de dorsal eminence vin ventral inflexion Scale bar equals 10mm

FIGURE 34 Ventral osteoderms of Polesinesuchus aurelioi holotype (ULBRAPVT003) in AndashD Abbreviation ab anterior

bar Scale bar equals 10mm

Ungual phalanges They are quite curved and lateromedially compressed with strongly sharp tips (Fig 32endashg) In both sides close to distal end there are grooves for blood vessels (Martz 2002) The morphology these elements are very similar to those from other aetosaurs as Neoaetosauroides engaeus (PVL 3525) Aetobarbakinoides brasiliensis (CPE2 168) Aetosauroides scagliai (PVL 2052) and Typothorax coccinarum

(Desojo amp Baeacutez 2005 Desojo et al 2012 Heckert et al 2010)Dorsal paramedian osteoderms Three elements are present (Fig 33andashc) The osteoderm A is subrectangular

in dorsal view and wider than long (a widthlength ratio of about 276) (Fig 33) This ratio is higher than in Aetobarbakinoides (196) Unfortunately this element is quite damaged Even so on the external surface there is an anterior bar with a small elevation devoid of ornamentation This bar becomes anteroposteriorly longer towards the lateral edge as in Aetosauroides scagliai (MCP 13-andashb PV) (Desojo amp Ezcurra 2011) The lateral edge is partially broken but permits the observation of an anterolateral projection as occurs in a large specimen of Aetosaurus

ferratus (SMNS 12670 Schoch 2007) In Polesinesuchus this projection is comparatively lower and its medial edge is not projected Posteriorly the paramedian osteoderms of Polesinesuchus present a strong ventral flexion as occurs in typothorascisine and desmatosuchine aetosaurs (Parker 2007 Desojo et al 2012) The ornamentation consists of irregular circular pits and ridges with a radial pattern projecting from the dorsal eminence as occurs in Aetosauroides scagliai (PVL 2073 MCP 13-andashb PV) Aetobarbakinoides brasiliensis (CPE2 168) Neoaetosauroides engaeus (PVL 3525) Aetosaurus ferratus (Schoch 2007) Lucasuchus hunti Paratypothorax

andressi (Long amp Murry 1995) Coahomasuchus (Heckert amp Lucas 1999) Stagonolepis robertsoni (NHMUK R4788) and Calyptosuchus wellesi (Long amp Murry 1995) (Desojo amp Ezcurra 2011 Desojo et al 2012) However the overall ornamentation of paramedical osteoderms of Polesinesuchus shows poor density In small specimens of Aetosauroides scagliai this fact is probably attributed to its juvenile ontogenetic stage (Desojo amp Ezcurrra 2011) the same situation of Polesinesuchus In contrast Typothorax (Long amp Ballew 1985) Redondasuchus (Heckert et

al 1996) and Chilenosuchus (Desojo 2003) present a reticular distribution of ornamentation (Desojo amp Ezcurra 2011 Desojo et al 2012) Furthermore the random pattern is observed in Desmatosuchus smalli Desmatosuchus

spurensis Longosuchus and Acaenasuchus (Long amp Ballew 1985 Parker 2005 2008 Parker amp Martz 2010 Desojo et al 2012) The dorsal eminence is posteromedially displaced incipient and does not contact the posterior edge as occurs in Aetosauroides scagliai (MCP 13-andashb PV) (Desojo amp Ezcurra 2011) An incipient dorsal eminence occurs also in Aetobarbakinoides brasiliensis (CPE2 168) Typothorax coccinarum and Redondasuchus

reseri (Long amp Ballew 1985 Heckert et al 1996) The ventral view cannot be assessed due the resin used for reinforce the structure of the osteoderm before the description of the Polesinesuchus Due to these characteristics osteoderm A probably belongs to the left posterior cervical or to the anterior dorsal region of the armor Desojo amp Ezcurra (2011) claim that the pitting density in cervical and dorsal osteoderms presents differences according to their position in the armor (e g a diminution of density towards the sacral region)

The osteoderm B comes from the right transverse segment and presents almost the same characteristics of the osteoderm A but differently its widthlength ratio is 215 (Fig 33) Its lateral ridge shows a rounded distal half the ornamentation is more conspicuous and the dorsal eminence forms an anteroposteriorly displaced ridge Due to the density of the ornamentation in osteoderm B this element is probably more anteriorly placed than A

The ornamentation of osteoderm C is distinct In dorsal view the ridges and pits are deeper in comparison with those from osteoderms A and B In the medial region the small pits merge increasing their size and being surrounded by anastomosed ridges (Fig 33) In posterior view the osteoderm C is not flexed According to this pattern this element was probably placed close to sacral region (Taborda personal communication 2012) Due to the position of its dorsal eminence that is medially displaced this element proceeds from the right side of the armor

Ventral osteoderms Seven elements are present in Polesinesuchus (Fig 34) They are almost squared slightly curved transversely and wider than long as occurs in Aetosauroides scagliai (MCP 13andashb PV Desojo amp Ezcurra 2011) Typothorax coccinarum (TTUP9214 fig440a Martz 2002) Coahomasuchus (Heckert amp Lucas 1999) and Stagonolepis robertsoni (Walker 1961 Heckert amp Lucas 2000 2002) The anterior bar is well-developed without ornamentation The pattern of ornamentation consists of pits and grooves in a radial fashion starting from the center of the osteoderm as in Aetosauroides scagliai (Desojo amp Ezcurra 2011) Among the preserved elements four are still enclosed in the matrix imbricated to each other which shows their original disposition in the ventral armor They are slightly flexed in posterior view

Phylogenetic analysis

In order to verify the likely position of Polesinesuchus within Aetosauria we used the data-matrix by Desojo et al (2012) which is constituted of 37 characters and 20 ingroup taxa plus the pseudosuchian ldquorauisuchianrdquo Postosuchus kirkpatricki and Revueltosaurus callenderi as outgroups Desojo et al (2012) modified the data-matrix from Parker (2007) and Parker et al (2008) The phylogenetic analysis was performed using the software TNT (Goloboff et al 2003 2008) The analysis was executed using Traditional Search The analysis recovered two most parsimonious trees (MPTs) of 67 steps each and consistency index of 0686 (Figs 35 36) The strict consensus tree is also depicted in Figure 37 See Appendix I for character coding of Polesinesuchus aurelioi

The phylogenetic analysis shows that Polesinesuchus is the sister taxon of Aetobarbakinoides brasiliensis and both form a clade that is a sister group of the large monophyletic clade composed of the subfamilies Desmatosuchinae and Typothoracisinae However this result must be taken with caution as the holotype of Aetobarbakinoides brasiliensis is quite incomplete with several missing entries in the data-matrix so the sister-group relationship between it and Polesinesuchus might be an artifact caused by the incompleteness of the former one and considered provisional Besides several postcranial characters present in are not preserved in the holotype of Aetobarbakinoides brasiliensis such as the pelvic girdle femora fibulae skull elements and tarsus

An interesting result worth pointing out is that the addition of Polesinesuchus in the data-matrix by Desojo et

al (2012) produced just slight changes in the topology found by these authors (only in the basis of the Aetosauria) The first most parsimonious cladogram (MPT Fig 35) shows Aetosauroides scagliai as the most basal aetosaur as seen in Desojo et al (2012) but not placed between Calyptosuchus wellesi and Stagonolepis robertsoni The second MPT (Fig 36) also presents Aetosauroides scagliai as basal taxon and the sister taxon of Coahomasuchus Neoaetosauroides engaeus occurs as the sister taxon of Calyptosuchus wellesi and Stagonolepis robertsoni The strict consensus tree of the present analysis (Fig 37) resembles the first topology of the Desojo et al (2012)

FIGURE 35 The first most parsimonious tree of 67 steps and consistency index of 0686 recovered in the analysis Note the

position of Polesinesuchus as sister group of the Aetobarbakinoides and the sister group relationship between them and

Desmatosuchinae plus Typothoracisinae

In the contribution by Desojo et al (2012 fig 19) the first tree shows the presence of the genus Coahomasuchus within a polytomy among Neoaetosauroides Coahomasuchus Aetosaurus and a derived group (also polytomic) comprising Stagonolepis Calyptosuchus and more derived forms (Typothoracisinae plusDesmatosuchinae) After a posteriory pruning of Coahomasuchus from the analysis the authors obtained a more resolved result where only the trichotomy among Calyptosuchus Stagonolepis and Typothoracisinae plusDesmatosuchinae remained As stated above in spite of the well resolved placement of Polesinesuchus as a sister taxon of Aetobarbakinoides this result should be seen with caution as we need to consider the small representativeness of postcranial characters in this phylogenetic analysis Previous phylogenies of Aetosauria (eg Parker 2007 2008 Desojo et al 2012 among others) are mainly based upon cranial and osteoderm characters as most members of this group lack information regarding postcranial skeletons The generalized use of osteoderms in such analyses could be distorting the ldquorealrdquo phylogenetic relationships of Aetosauria In fact the validity of the use

of osteoderms and their pattern of ornamentation has been discussed in several contributions (Martz amp Small 2006 Parker 2007 2008 Cerda amp Desojo 2011) Therefore as long as postcranial data are not available for many aetosaurs we should be aware that cranialosteoderm based phylogenies might embrace a great deal of biases due their homoplastic nature (Parker 2008) Moreover some characters used at the present moment can be individual variations due to ontogenetic development and anatomical position from where the osteoderm comes in the body (Harris et al 2003 Martz amp Small 2006) At this point these kind of problems cannot be avoided due to the overall lack of postcranial information in many aetosaurs such as Chilenosuchus forttae (Casamiquela 1980) Tecovasuchus chetterjeei (Martz amp Small 2006) Acaenasuchus geoffreyi (Long amp Murry 1995) and Redondasuchus reseri (Hunt amp Lucas 1991)

FIGURE 36 The second tree recovered in the present analysis Note the position of Polesinesuchus as sister group of the

Aetobarbakinoides and the sister group relationship between them Desmatosuchinae plus Typothoracisinae

FIGURE 37 Strict consensus tree recovered in the present analysis Note the unresolved polytomies among basal

stagonolepidids and also sister group relationship between Polesinesuchus plus Aetobarbakinoides and Desmatosuchinae plus

Typothoracisinae

Discussion

Most of the aetosaur studies (eg Heckert amp Lucas 2000 Parker 2007 Desojo et al 2012 Desojo et al 2013) have pointed out the importance of American aetosaurs and their role in the origin distribution and diversity of this group especially from Upper Triassic deposits from the Santa Maria Formation (Lucas amp Heckert 2001 Desojo amp Ezcurra 2011 Desojo et al 2012) The description of Polesinesuchus significantly increases the availability of data

regarding South American aetosaurs (particularly concerning postcranial data) This new species is based upon a unique combination of characters many of them shared with other two genera from the Santa Maria Formation Aetosauroides and Aetobarbakinoides As stated in the diagnosis some character states are shared with Aetosauroides whereas others with Aetobarbakinoides For instance in Polesinesuchus the cervical centra present a very marked ventral keel as occurs in Aetosauroides In contrast in Aetobarbakinoides this structure is absent Additionally the dorsal vertebrae of Polesinesuchus do not present both infradiapophyseal laminae and the well-rimmed fossa laterally placed in the centra as is seen in Aetosauroides The absence of these structures is shared with Aetobarbakinoides Furthermore the morphology of the appendicular skeleton is quite similar to that of Aetosauroides as both present robust limbs Conversely Aetobarbakinoides had gracile limbs In addition some derived features were recognized in the pelvic girdle of the new taxon which were not coded in the updated data-matrix by Desojo et al (2012) which is mostly based upon the morphology of the osteoderms

Despite the unequivocal position of Polesinesuchus in the topology generated here some problems arose during this study Among them the small representativeness of postcranial elements in phylogenetic studies of Aetosauria As Polesinesuchus adds a great amount of postcranial features regarding this group a new evaluation of postcranial characters of aetosaurs housed in different collections around the world as well as the inclusion of as many of them as possible in future phylogenetic studies is necessary in order to improve anatomical phylogenetic and paleobiogeographic knowledge of the Aetosauria

Acknowledgements

We thank many people who allowed us to study specimens under their care J Ferigolo (FZB) C Schultz (UFRGS) MC Malabarba (PUCS) AAS Da-Rosa (UFSM) J Powell (PVL) P Holroyd (UCMP) R Schoch (SMNS) and S Chapman (NHMUK) We are also thankful to Fabriacutecio Barreto (ULBRA) for his technical support on the photographic material and Jeremiacuteas Taborda (MACN) for discussion about this paper We thank the useful comments and suggestions that improved the manuscript provided by the reviewers Ignacio Cerda and especially Andrew Heckert CNPq Research Grants 500919 frasl 2009-7 and 3018012012-6 to SDS supported this research Access to the free version of TNT 11 was possible due to the Willi Henning Society

References

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Gondwana Mar del Plata 1967 Actas 307ndash324

Bonaparte JF (1971) Los tetraacutepodos del sector superior de la Formacioacuten Los Colorados La Rioja Argentina (Triaacutesico

Superior) Opera Lilloana 22 87ndash102

Brochu CA (1996) Closure of neurocentral sutures during crocodilian ontogeny implications for maturity assessment in

fossil archosaurs Journal of Vertebrate Paleontology 19 49ndash62

httpdxdoiorg10108002724634199610011283

Brochu CA (2001) Progress and future directions in archosaur phylogenetics Journal of Paleontology 75 1185ndash1201

httpdxdoiorg1016660022-3360(2001)075lt1185pafdiagt20co2

Case EC (1922) New reptiles and stegocephalians from the Upper Triassic of Western Texas Carnegie Institution

Publication 321 1ndash84

Casamiquela RM (1960) Noticia preliminar sobre dos nuevos estagonolepoideos Argentinos Ameghiniana 2 3ndash9

Casamiquela RM (1961) Dos nuevos Estagonolepoideos Argentinos (de Ischigualasto San Juan) Revista de la Asociacoacuten

Geoloacutegica Argentina 16 143ndash203

Casamiquela RM (1967) Materiales adicionales y reinterpretacioacuten de Aetosauroides scagliai (de Ischigualasto San Juan)

Revista del Museo de La Plata 5 173ndash196

Casamiquela RM (1980) Nota sobre restos de un reptil aetosauroideo (Thecodontia Aetosauria) de Quimal Cordillera de

Domeyko Antafogasta In Anonymous (Eds) Prueba de la existencia del Neotriaacutesico continental en los Andes del Norte

de Chile Actas Congreso Argentino de Paleontologiacutea y Bioestratigrafiacutea No2 y Congresso Lationoamericano de

Paleontologiacutea 1 135ndash142

Cerda IA amp Desojo JB (2010) Dermal armour histology of aetosaurs (ArchosauriaPseudosuchia) from the Upper Triassic

of Argentina and Brazil Lethaia 44 (4) 417ndash428

httpdxdoiorg101111j1502-3931201000252x

Desojo JB (2003) Redescription of the aetosaur Chilenosuchus forttae Casamiquela (Diapsida Archosauria) presence of

continental Triassic in Northern Chile Revista Geloacutegica de Chile 30 53ndash63

httpdxdoiorg104067s0716-02082003000100004

Desojo JB amp Baacuteez AM (2005) El esqueleto postcraneano de Neoaetosauroides (Archosauria Aetosauria) del Triaacutesico

Superior Del centro-oeste de Argentina Ameghiniana 42 115ndash126

httpdxdoiorg105710amghv48i1(265)

Desojo JB amp Baacuteez AM (2007) Cranial morphology of the Late Triassic South American archosaur Neoaetosauroides

engaeus evidence for aetosaurian diversity Paleontology 50 267ndash276

httpdxdoiorg101111j1475-4983200600608x

Desojo JB amp Viscaiacuteno SF (2009) Jaw biomechanics in the South American aetosaur Neoaetosauroides engaeus

Palaumlontologische Zeitschrift 83 499ndash510

httpdxdoiorg101007s12542-009-0032-6

Desojo JB amp Ezcurra MD (2011) A reappraisal of the taxonomic status of Aetosauroides (Archosauria Aetosauria)

specimens from the Late Triassic of South America and their proposed synonymy with Stagonolepis Journal of Vertebrate

Paleontology 31 596ndash609

httpdxdoiorg101080027246342011572936

Desojo JB Ezcurra MD amp Kischlat EE (2012) A new aetosaur genus (Archosauria Pseudosuchia) from the Early Late

Triassic of Southern Brazil Zootaxa 3166 1ndash33

Desojo JB Heckert AB Martz JW Parker WG Schoch RR Small BJ amp Sulej T (2013) Aetosauria a clade of

armoured pseudosuchians from the Late Triassic continental beds Geological Society London Special Publications 379

203ndash239

httpdxdoiorg101144sp37917

Dias-da-Silva S Cabreira SF amp Roberto-da-Silva L (2011) Occurrence of giant stereospondyl remains in the Santa Maria

Formation (Middle frasl Upper Triassic of southern Brazil) Alcheringa An Australasian Journal of Palaeontology 35 11ndash19

httpdxdoiorg10108003115511003793538

Dias-da-Silva S Sengupta DP Cabreira SF amp Roberto-da-Silva L (2012) The presence of Compsocerops

(Brachyopoidea Chigutisauridae) (Late Triassic) in southern Brazil with comments on chigutisaurid palaeobiogeography

httpdxdoiorg101111j1475-4983201101120x

Goloboff PA Farris JS amp Nixon K (2003) TNT Tree analysis using new technology Version 11 Program at

documentation Available from httpwwwzmucdkpublicphylogenyTNT (accessed 23 December 2013)

Goloboff PA Farris JS amp Nixon K (2008) TNT a free program for phylogenetic analysis Cladistics 24 774ndash786

httpdxdoiorg101111j1096-0031200800217x

Gower DJ amp Schoch RR (2009) Poscranial anatomy of the Rauisuchian Archosaur Batrachotomus kupferzellensis Journal

of Vertebrate Paleontology 29 103ndash122

httpdxdoiorg10108002724634200910010365

Heckert AB amp Lucas SG (1999) A new aetosaur (Reptilia Archosauria) from the Upper Triassic of Texas and the

phylogeny of aetosaurs Journal of Vertebrate Paleontology 19 50ndash68

httpdxdoiorg10108002724634199910011122

Heckert AB Lucas SG Rinehart LF Celeskey MD Spielmann JA amp Hunt AP (2010) Articulated skeletons of

aetosaur Typothorax coccinarum Cope (Archosauria Stagonolepididae) from the Upper Triassic Bull Canyon Formation

(Revueltian Early-mid Norian) Eastern New Mexico USA Journal of Vertebrate Paleontology 30 619ndash642

httpdxdoiorg10108002724631003763524

Heckert AB amp Lucas SG (2000) Taxonomy phylogeny biostratigraphy biochronology paleobiogeography and evolution of

the Late Triassic Aetosauria (Archosauria Crurotarsi) Zentralblatt fuumlr Geologie und Palaumlontologie 11ndash12 1539ndash1587

Irmis RB (2007) Axial skeleton ontogeny in the Parasuchia (Archosauria Pseudosuchia) and its implications for ontogenetic

determination in archosaurs Journal of Vertebrate Paleontology 27 350ndash361

httpdxdoiorg1016710272-4634(2007)27[350asoitp]20co2

Langer MC Ribeiro AM Schultz CL amp Ferigolo J (2007) The continental tetrapod-bearing Triassic of South Brazil

New Mexico Museum of Natural History and Science Bulletin 41 201ndash218

Lautenschlager S (2008) Revision of Rauisuchus tiradentes (Archosauria Rauisuchia) from the Late Triassic (Carnian) Santa

Maria Formation of Brazil and its implications for rauisuchian phylogeny Ludwig-Maximilians-Universitaumlt Munich

Lecuona A amp Desojo JB (2011) Hind limb osteology of Gracilisuchus stipanicicorum (Archosauria Pseudosuchia) Earth

and Environmental Science Transactions of the Royal Society of Edinburgh 102 105ndash128

httpdxdoiorg101017s1755691011000181

Long RA amp Murry PA (1995) Late Triassic (Carnian and Norian) tetrapods from the southwestern United States New

Mexico Museum of Natural History and Science Bulletin 4 1ndash254

Long RA amp Ballew KL (1985) Aetosaur dermal armor from the Late Triassic of southwestern North America with special

reference to material from the Chinle Formation of Petrified Forest National Park Museun of Northern Arizona Bulletin

47 45ndash68

Lucas SG amp Heckert AB (2001) The aetosaur Stagonolepis from the Upper Triassic of Brazil and its biochronological

significance Neues Jahrebuch fuumlr Geologie und Palaumlontologie Monatshefte 12 719ndash732

Lucas SG amp Heckert AB (2002) The Hyperodapedon biochron Late Triassic of Pangea Albertiana 27 30ndash38

Martz JW (2002) The morphology and ontogeny of Typothorax coccinarum (Archosauria Stagonolepididae) from the Upper

Triassic of the American southwest MS Thesis Texas Tech University Lubbock TX 200 pp

Martz JW amp Small BJ (2006) Tecovasuchus chatterjeei new aetosaur (Archosauria Stagonolepididae) from the Tecovas

Formation (Carnian Upper Triassic) of Texas Journal of Vertebrate Paleontology 26 308ndash320

httpdxdoiorg1016710272-4634(2006)26[308tcanaa]20co2

Martz JW amp Small BJ (2013) Anatomy Phylogeny and Paleobiology of Early archosaurs and their Kin Geological Society

London Special Publications 1ndash379

Nesbitt SJ (2011) The early evolution of archosaurs relationships and the origin of major clades Bulletin of the American

Museum of Natural History 352 1ndash292

httpdxdoiorg1012063521

Parrish JM (1986) Locomotor adaptations in the hind limb and pelvis of the thecodontia Hunteria 1 1ndash33

Parker WG (2007) Reassessment of the aetosaur Desmatosuchus chamaensis with a reanalysis of the phylogeny of the

Aetosauria (Archosauria Pseudosuchia) Journal of Systematic Paleontology 5 1ndash28

httpdxdoiorg101017s1477201906001994

Parker WG (2008) Description of new material of the aetosaur Desmatosuchus spurensis (Archosauria Suchia) from the

Chinle Formation of Arizona and a revision of the genus Desmatosuchus Paleo Bios 28 1ndash40

Parker WG Stockert MR amp Irmis RB (2008) A new Desmatosuchinae aetosaur (Archosauria Suchia) from the Upper

Triassic Tecovas Formation (Dockum Group) of Texas Journal of Vertebrate Paleontology 28 692ndash701

httpdxdoiorg1016710272-4634(2008)28[692andaas]20co2

Parker WG amp Martz JW (2010) Using positional homology in aetosaur (Archosauria Pseudosuchia) osteoderms to evaluate

the taxonomic status of Lucasuchus hunti Journal of Vertebrate Paleontology 30 1100ndash1108

httpdxdoiorg101080027246342010483536

Perez PA amp Malabarba MCSL (2002) A Triassic freshwater fish fauna from the Parana Basin in southern Brazil Revista

Brasileira de Paleontologia 4 (27) 33ndash54

Peyer K Carter JG Sues HD Novak SE amp Olsen PE (2008) A new suchian archosaur from the Upper Triassic of North

Carolina Journal of Vertebrate of Paleontology 28 363ndash381

httpdxdoiorg1016710272-4634(2008)28[363ansaft]20co2

Reichel M Schultz CL amp Soares MB (2009) A new traversodontid cynodont (Therapsida Eucynodontia) from the Middle

Triassic Santa Maria Formation of Rio Grande do Sul Brazil Palaeontology 52 229ndash250

httpdxdoiorg101111j1475-4983200800824x

Romer AS (1956) Osteology of the Reptiles University of Chicago Press Chicago Illinois 772 pp

Sawin HJ (1947) The psedosuchian reptile Typotorax meadei Journal of Paleontology 21 201ndash238

Sereno PC amp Arcucci AB (1990) The monophyly of crurotarsal archosaurs and the origin of bird and crocodile ankle joints

Neues Jahrbuch fuumlr Geologie und Palaumlontologie 180 21ndash52

Soares MB Schultz CL amp Horn BLD (2011) New information on Riograndia guaibensis Bonaparte Ferigolo amp Ribeiro

2001 (Eucynodontia Tritheledontidae) from Late Triassic of Southern Brazil anatomical and biostratigraphical

inplications Anais da Academia Brasileira de Ciecircncias 83 329ndash354

httpdxdoiorg101590s0001-37652011000100021

Sulej T (2009) The skull of an early Late Triassic aetosaur and the evolution of the stagonolepidid archosaurian reptiles

Zoological Journal of the Linnean Society 158 860ndash881

httpdxdoiorg101111j1096-3642200900566x

Scheyer TM Desojo JB amp Cerda IA (in press) Bone histology of phytosaur aetosaur and other archosauriform

osteoderms (Eureptilia Archosauromorpha) The Anatomical Record

Schoch RR (2007) Osteology of the small archosaur Aetosaurus from the Upper Triassic of Germany Neues Jahrbuch fuumlr

Geologie und Palaumlontologie 246 1ndash35

httpdxdoiorg1011270077-774920070246-0001

Schultz CL (2005) Biostratigraphy of the non-marine Triassic is a global correlation based on tetrapod faunas possible In

Koutsoukos EAM (Ed) Applied Stratigraphy Springer 123ndash145

httpdxdoiorg1010071-4020-2763-x_6

Small BJ (2002) Cranial anatomy of Desmatosuchus haplocerus (Reptilia Archosauria Stagonolepididae) Zoological

Journal of the Linnean Society 136 97ndash111

httpdxdoiorg101046j1096-3642200200028x

Taborda JA Cerda IA amp Desojo JB (2013) Growth curve of Aetosauroides scagliai Casamiquela 1960 (Pseudosuchia

Aetosauria) inferred from osteoderm histology In Nesbitt SJ Desojo JB amp Irmis RB (Eds) Anatomy Phylogeny

and Paleobiology of Early archosaurs and their Kin Geological Society London Special Publications 379 413ndash423

Walker AD (1961) Triassic reptiles from the Elgin area Stagonolepis Dasygnathus and their allies

PhilosophicalTransactions of the Royal Society of London 244 103ndash204

httpdxdoiorg101098rstb19610007

Zacarias JD (1982) Uma nova espeacutecie de tecodonte aetossaurio Aetosauroides subsulcatus sp nov da Formaccedilatildeo Santa

Maria Triaacutessico do Rio Grande do Sul Brasil Dissertaccedilatildeo de Mestrado Universidade Federal do Rio Grande do Sul

Porto Alegre 66 pp

Zerfass H Lavina EL Schultz CL Garcia AJV Faccini UF amp Chemale Jr F (2003) Sequence stratigraphy of

continental Triassic strata of Southernmost Brazil a contribution to Southwestern Gondwana paleogeography and

paleoclimate Sedimentary Geology 161 85ndash105

httpdxdoiorg101016s0037-0738(02)00397-4

APPENDIX

Scores for Polesinesuchus aurelioi in the cladistic character matrix of Desojo et al (2012)

Polesinesuchus aurelioi

0 110 00000 1 010 00

Abstract

Introduction

Ant Dorsal 1

Ant Dorsal 2

Ant Dorsal 3

Ant Dorsal 4

Ant Dorsal 5

Mid Dorsal 13

Mid Dorsal 14

Mid Dorsal 15

Sacral 1

Sacral 2

Caudal 1

Caudal 2

Caudal 3

Caudal 4

Caudal 5

Discussion

References

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