Journal of South Ameriean arth Sciences 107 (2021) 103145, ELSEVIER Contents lists available at ScienceDirect Journal of South American Earth Sciences journal homepage: wwrw.elsevier.comilocateyjsames Triassic sauropodomorph dinosaurs from South America: The origin and ® diversification of dinosaur dominated herbivorous faunas Diego Pol”, Alejandro Otero”, Cecilia Apaldetti ®, Ricardo N. Martinez” {CONICET Museo Paowlige Epi enn Ave Forte 140, Tren, 8100, Cab, Argrin COMET ace de La Pla Puc da Bet a Plt, Burs Abe Arena sn y Mase de Css Nasier, UNS? CICGEOBO, CONICET, pe 400 (More) San Jy, 400, Son Jom Arnie ARTICLE INFO apsTRact Rowers Srutepadomorp Tine South mere Seuropodomeorpha isthe fist major dinosaurian group that radiated dusing the Tulasi. Dung this te the s1oup underwent major changes ln body plan, lneluding the nequlston of featutes elated to herbivory, lage body sie, and qurupedality. By the ea ofthe Late Tissc, pproxinitely 30 malin yt afte the ong of Ainosaus, samopodomorptis predominated the niches of lage esbivores in continental ecosystems throughout the world. The Tense siopadonionph diversity inches diverse linenges ith ste disparty in body sn, feeding blomcehanles, and locomerion types, raging fiom sal (~-10 hg) bipedal axa to the large (>5 tons) quadiupedal lesemsauids. The South Ameican teeocd has provided key information to undelstand certain stages of his votionty radiation. We review hee the diversity and composition of satopdomorp fas ftroughout the Late Trias of South Amesien and highlight their conerbuton fr understanding ce evolution of tis group, 1. Introduction, Sauropodomorpha is one of the three major groups of Dinosaur, along with Theropoda and Ornithischia (Beaton, 1983; Novas, 1996), ‘and became the most conspicuous herbivores of terrestrial ecosystems of the Mesozoic, Sauropodomorpha includes che gigantic quadmpedal sauropods and a diverse assemblage of early lineages with much staller body size, diverse dietary habits, and obligatory or facutatively bipedal locomotion, These early lineages are curently referred as basal or early sauropodomorphs (Bronzat, 2017) or more traditionally grouped in Prosauropoda (Seren0, 1999; Galton and Upchurch, 2008, a term now in disuse ns most phylogenetic studies inthe last decade agroe i the perapiyly of this group, The Late Triassic was a critical time for the history of Sat ropodomorpha for several reasons, Firstly, many early lineages ‘appeared during this time and became the most diverse dinoseutian ‘group in terrestrial ecosystems by the end of this period (Bonaparte, 1082; Benton, 1983; Langer et al, 2010). Secondly, during this time feeding habits markedly changed in the group towards the Tandmark acquisition of an herbivorous diet (Benton, 1983; Barrett etal, 2011, 2014). Thirdly, thete are important changes reeorded in thelr locomo. tion and body plan during the Late Triassic, including the earliest Coxsespoading autor Email adarese dpoltietog.ar (D, Pad, ps /do.org/ 10.1016, sames.2020.108145 acquisition of a quadrupedal stance among dinosaurs (Raut et al 2011), Finally, the Late Triassic is also important because the onset of| smuropedomorph gigantism (and the origin of Sauropoda sensu Yas, 2007) ean be traced back to this period (Apaldett etal, 2018), ‘The South American fossil record of Ssuropodomorpha es provided important information for understanding the diversity ofthe group and some of the most important evolutionary processes ofthe Late Trinssie Gonaparte, 1982; Preto eral, 20185 Apaldeti etal, 2018). There are 14 species described so far from differen Upper Teinsie beds of north western Argentina and southern Brazil (Fig. 1), and these constitute approximately half ofthe sauropodomorph diversity kuown worldwide for the Triassic, Moreover, some of these taxa represent lineages that are ‘unknown in other regions (except for southern Afren in somieenses) and thus provide wique information for understanding certain episodes in the evolutionary history of Sauropodomorpha, ‘The South American taxa include the earliest known sau ropodomorphs, which are recorded in late Carian deposits of the Ischignalasto Formation of Argentina snd the Santa Maria Formation of Brazil (Martinez etal, 2011, 201325 Langer etal, 2010). These early lineages are critical for understanding sauropodomorph origins and are not recorded in other continents. These species share many plesiomor ples with other Garian dinosturs, tothe exten shat certain taxa have Received 2 June 202; Received in revised form 28 Oetber 2020; Accepted 38 December 2020 Availabe online 11 January 2021 (0895.9811/© 2021 Eee Lic All ight serveddisputed affinities and are interpreted as early theropods in some studies (8 Boraptor; Sereno etal 19935 Baton etal, 2017) During the Norian and Rhaetian, sauropodomorphs became the most, ‘sbundant terrestrial vertebrates in many deposits of Pangea (Boar, 1982; Benton, 198%; Brasatte etal, 2010). The South American record of these times includes representatives of multiple lineages, most of ‘which have adaprations to omnivory or herbivory (Barrett, 2000, 20145 Button et al, 2017) and others show the earliest aequiston of quad. rupedality (¢., Bonaparte, 1972; MePhee ets, 2018). Some ofthese lineages share steking smarts with taxa recorded in southern Aftien (eg, Yates, 2006; 2007; Apaldett et a, 2012; 2018; MePhee et al, 20158), sueh as Tessensaurids tat show the earliest steps cowards ‘sigantism nod ace regarded as the earliest sauropods in some studies (Yates, 2007; Apaldeti et al., 2018; se below. In this contribution we review the sauropodomorph diversity recorded duting the Late Triassic in South America and analyze the temporal distribution of the different sauropodoniorph lineages. We dlsenss the appearance of diferent lineages through time based on 8 phylogenetic analysis chat sumatizes recent conteibtions (see Sup: plementary Information) and highlighting the contribution of the South American taxa for understanding Key stages in the evolution of ‘Sauropodomorpha Institutional Abbreviations: CAPPA/UESM, Centro de Apolo Pesquisa Paleontoldgien da Quarta Colonia da Universidade Federal de Sant Maria, Smo Joso do Pole'sne-RS, Brazil; MCN, Museu de Ciencias Naturais, Fundaeao Zoobotanica do Rio Grande do Sul, Porto Alegre, Brazil; MCP, Museu de Ciencia e Tecnologia, Pontifcin Universidade aril do Rio Grande do Sul, Porto Alegre, Bree; PL, Instituto Miguel Lillo, San Miguel de Tucunén, Tweumén, Argentina; PVSI, Divisién de Paleontologia, Museo de Ciencias Naturales de la Uni versidad Nacional de San Juan, San Juan, Argentinas UBRGS PV, Vertebrate Paleontology Collection at Universidade Federal do Rio Grande do Sul, Brazil; UFSM, Laboratorio de Estratgrafi e Paleo Diologia, Universidade Federal de Santa Maria, Brazil; ULBRA, Museu de (Ciencias Naturais, Universidade Luterana do Brasil, CanoasRS, Brazil 2. South American sauropodomorph faunas ‘The Triassic Fossil record of South American sauropodomorphs ranges from the Caran (0 the end-Trassie and is restricted to five Jal of Seth Ameren Bah Siecs 1072021 108445 formations that crop out in southem Brazil (Santa Maria and Caturite formations) and northwestern Argentina (Ischigualasto, Los Colorados, 1nd Quebrada del Barro formations). Recent advances in taxonomic and phylogenetic studies as well as progress in geochronologieal dating of these units allow dividing che known diversity into three different sa. ropodomorph faunas that are used here to structure this work: a late (Carian, an early Notian, and a mi Norian-Rhsetian (ig. 2. Although the phylogenetic arrangement of South American sau ropodamorphs still has some uncertainties, progress an agreement in recent years have led to the recognition of very distinet lineages that appear in different Triassic units (Cabrera et al, 20165 Otero et a. 2016; McPhee et al, 2017; Apaldetl et aly 20185 Miller e al, 2018). ‘This provides s basie pattern of taxonomic affinities for the different associations of sauropodomorphis recorded in Soutt Ameria. The ret oral component ofthe three sauropodoniorph fatinas sed in tis re ‘lew is based on previous studies that include radioisropie dating (see Martinez et a, 2011; Langer et al, 2018; Desojo et a, 2020) and paleomgneti studies (Kent ets, 2014) of sautopodonierph- bearing units, as well as the sauropodomorph content of other deposits (ex. Martinez et al, 2015) (See Fig. 2) ‘The division of sauropodomorph faunas used here only aims iden: tilfing the association of species recorded in similar aged deposit and thar share similar anatonieal features, reflected in most eases in that they ae phylogenetically close ro exch other. Where appropriate we cite references to vertebrate biostratigraphic assemblage zones (Albdale cal, 2001; Langer et al, 20078; Abdala and Ribeiro, 2010) or biozones (Martinez at al, 20115 2019s) recognized locally in certain sedimentary sequences (se Fig 2). The three sauropodomrph faunas discussed here fze temporally brond nnd far mae resrietive taxonomically chan those ‘ised in atenpts to define verebratebiochronologe units, sich as the tlobal ‘land vertebrate faunachtons’ proposed for the Triassic by some authors (Lucas, 1998; Langer, 2005). The validity of these, especially at fk Pangean scale, 1S dubions and lies beyond the scope of this contribution, 2.1, Lote Carnian sauropodomorph fauna ‘This founa includes the oldest dinosaurs known worldwide, which have been found in the lower levels of the Ischigualasto Formation (Marines et al, 2011) and the Sante Maria Fortuation (Langer et al. Fig. 1. Distibusion sauropodomoyph dinosaurs hom South America dung the Late Tse. All taxa shown lathe gue ae undisputed sausopodonoxphs, with he ‘exception of Grades tat has ben depicted asa esi heroped or satschian in ferent sues (see txtJal of Seth Ameren Bah Siecs 1072021 108445 cone mn ort af, Fig. 2, Ages ofthe five sauropodomorphbesrng formations inthe Tassie of South America. Fest column vepesents the age thieesausopedomuph faunas de ‘essed inthis connibation. For the Qhebrada del Baro Formation we show rhe bistatigtaphie age propased by Maines el. (2015) For Ishigsto Los ‘Coloaéos the let coluna shows the Martinez el. (201Sa) and Keat eta. (2014) proposal based on data fom the Iehigulaso Provinlal Pask region and che High column follows Desojo et al (2020) based on data ion Cao de as Laas locality. Vertebrate iosttatgtaphie zation ofeach propsal shown by each column (see abbcevistion) For the Sata Masia Catia formations we sho the proposal of Lage eal. (2018) Boundaies of formations and biozones te shaded die age wertainy, exept For those ofthe Iehigalasto Formation enelated using Bayesian age model (ee Desojo tl, 2020). Stas nant raolstople platen Gates and tangles represent possible detrital ages (following Lang eal, 2018). Date in light gay are Ar/Ar dats ad in dark gray are U/M dates) Abree ‘atlas: Az, assemblage zone; Bz, biozone; Exa, Euereodon: Hyp, Hyperedapeion; Joe, Jachalra, Rio, Rogrand: SEH, Seaptany Exuretodan Heerasars ‘ey, Teymbai 2010; Cabreira et al, 2016), ‘The Ichiguslasto Fortation is « luvial sedimentary saquence that ranges berween 700 and 1000 at thick in diferent ocerops located northwestern Argentina (Martinez etal, 20134; Desojo eta, 2020). “This nit ns been the source ofa least seven early dinosaurs, including ‘two ofthe best kaown and studied Carnian dinosauss:Herrerasauns and Eorapor. The sauropadoniorph diversity inthis unit includes thre taxa (Ge, Chromogisaurus, Eoapior, Penphagia, although some authors have interpreted Earaptor as a non-sauropodomorph (Sereno eal, 199%; Baron et sl, 2017). This diversity of early dinosaurs is the result of intensive paleontological prospection atthe type locality (Ischigualasto Provincial Park) over the lst decades (since the 1990's), whic has also yielded an extensive sample of over 2000 vertebrates collected throughout the sequence (Martinez etal, 2011, 2013). This allowed the distinction of three abundance-based biozones, the earlier af which ‘contains over 90% ofthe collected fosis, includes all the dinosaur re rains, and extends along the lower third ofthis nit. Tis isthe Sea Phony Beaeretodon Herrerasmirus biozone (Martinez ot al, 2011, 20138; Fig. 2) and an Ar-Ar radioisoropi date from an ash layer close ro the base ofthe Ischigualasto Formation yielded an age of 291.40 4 0.3 Ma, placing this assemblage inthe late Casnian (Marunez eal, 2041, Fig. 2). Desojo et al. (2020) studied a sequence of this formation at a different locality and published slightly younger U-Pb dates for the Ischigualaso Formation (Fig. 2), while recognizing two equivalent bio- zones aged between 229.20 + 0.10 Ma and 226.85 + 1.45 Ma. ‘The other unit that provided relevant materials of late Caran sau ropodomorphs is the Santa Maria Formation from southern Brazil Although fossil verrebrates (including the dinosaur Stauikosaurus) front this unit have been known for many decades (Huene, 19425 colbert, 1970), it was only during the last 20 years that dinosaur exploration resulted in the discovery of new significant dinosaur remains. Four early sauropodomorphs (Sonali, Pampadromaens, Burolstes, Bagualosau rus) have been deseribed go fa fom the Santa Maria Formation. These ‘remains come from a sequence less than 100 m thick deposited in an ephemeral fluvial lacustrine environment. These rocks are recognized as the Alenioa Member of the Santa Maria Formation, or alternatively roferred in a sequence stratigraphy seheme as the lower part of the Candelaria Sequence of the Santa Maria Supersequence (lon et al, 2014). These levels have provided a diverse assemblage of vertebrates characterized as the Hyperudapedon Assemblage Zone and have been traditionally correlated with the Ischigualasto Formation (Barberena, 197; Bonaparte, 1982; Abdala eta, 2001; Langer etal, 20078). & recent U-Pb radioisoropie age on detital zircons confirmed this idea (233.28 £0.78 Ma; Langer eta, 2018, ig. 2) and the late Carian age for the oldest dinosaur assemblages. The seven described sauropedomorphs from the late Carnian of South America (Earapror, Sarumali, Cromogisaurus, Panphagia, Pan adromaeus, Burilestes, and Bagualosaurs; Fis. 1 aa 3) comprise 50% ff the known ‘Triassic sairopodomorphs from South Americe and represent an easly diversity that has been revealed only in the last decade and (Table 1) These taxa are mostly known from single speci mens and large collection effors have shown thar non-dinosanrian groups (eg. rhynchosaurs, eynodonts, pseudosuechians) were signif cantly more abundant than seuropodomorphs during this time (see Martinez etal, 2011; 20138) In this sens, these studies provided solid data to consider the earliest sauropodomorphs as minor components of the terestrial fatnas of South Amerien during the late Carian, Sa ropodomorphs are absent front Carian deposits in ll other land miasses, n which sauropodomiorphs appear only during or after the Norian. The ‘only possible pre Norian sanropodomorph outside South America Is Nyasasaurus fromthe Middle Triassie Manda Formation of Tanzania “Tis taxon was orginally interpreted as a dinosaurform (Nesbit etal, 2013) but a recent analysis depicted it as an early sauropadomorph (Garon et al, 2017) Is status as a sauropodomorph, of even as dinosaur, is comtroversial and poorly supported given its highly frag mentary nature CLanger etal, 2017). Further remains are needed to assess the presence of sanropodomorphs in the late Carnian outside South AmericaJal of Seth Ameren Bah Siecs 1072021 108445 Fig. 3. Mateials of selected sau ropodomaxphe fiom the Carnian af South [xtetiea. ALG, Borarar hiner (PVST 512) A, skull In lateral ew, B, aneior vet In lateral lew, C, metatatsal Vin planar view, sale bats equal 5 em, 5 mi, and 1 em, Iespeetvely; D-F Panga pros (PVS) £874) Dy ight dearay in tera view, mid cervical verteba in ght lateral View, let ‘mandibular tooth ia buccal ve, eae bate fal 2 em, 1m, ae 1 a espeeively; 6G, Satumatia tpiiqui (MCP 3948.PV) patie in lateral view, scale bar eas 9 fm; H, Bapulsauras agudonss (UFRGS PV 1099-7) sal in ne ew (odie from Preto et ay 2018), stale bar equals 2 ent 1, Boles cts (CAPPA/UFSM 0035), 1 sh in lateral lew (odie os aes Cy 20180), J, ptemanillay tooth. scale ‘bar equals 5 om el 2, vespeivey KM, Panpadromeas "barbereet (GLBRAPVTOIG) K, maxilla, Lanteior ‘maxi teeth, and M, data evessed) in Inter ve (etfied tom Lange et a 2018), scale bars equal 1 em, 5m, and 8 mm (espectvey). ‘The seven species of early sauropodomorphs inthe late Carian of ‘South Americ also contrast withthe low diversity of other dinosaueian, lineages in the Ischigualasto and Santa Maria formations. Infact, the ‘setual presence of theropods or omithisehians has been debated given some authors interpret herrerasnids as non-aheropods (e.g, Lager, 2004) at Psanosauris as a non-dinosatrian dinosauriform (Aenalin ‘tal, 2017; Baron, 2019), The diversfiation in multiple ealy lineages ‘of Sauropodomorpha during the late Carnian is the fist striking difer fence between the evolutionary dynamies of this clade in comparison With Theropoda and Orithisehia ‘The taxonomic identity of the seven late Cernian sauropodomorphs have never been seriously questioned, even though the morphological dlfferences between taxa are relatively minor and some taxa have been found at the same locality or in stratigraphic level (Martinez et al 2O13a; Miller etal, 2018b; Langer etal, 2019; Miller and Garcia, 2018). Anatomical distinctions between these taxa do exist and are re fected in the scorings of characters used in phylogenetic datasets. For Instance, in te pylogenetie dataset presented in dis contribution (see ‘Supplementary Information), « pairwise comparison ofthe scorings of these seven taxa reveal they differ on average sn 143% oftheir scored ratrix cells, with pairwise differences ranging between De and 24% (oF ‘a range between 4 and 87 matrix cells seored differently; see Table 2), “The only exception to these differences isin Samumiaia and Chromo sourus, which only differ in one scoring in the dataset used here (Table 2). This is due to the large amount of missing entries in Chro 1mogisaurs (869) and the face that some differences noted by previous authors (Ezcira, 2010; Martine? etal, 2013) are not represented as phylogenetic characters in the data matrix. Despite the anatomical diferences that support their taxonomic distinction, late Carnian sauropodomomphs share basic features oftheir anatomy, stall body size (>50 kg), and bipedal locomotion, chats terizing the body plan of early sauropodomorpls (Fi. 1). The well preserved remains of many of these taxa show three cranial and three Dosteranial apomorphies shared with other members of Saw ropodomorphe that supports their inclusion atthe base of this elade (ig. 6). These features include the anterior margin of the maxilla with & strong inflection a the base of the ascending ramus (char. 25.15 Fig. 3A, 1), quadretojugal ramus of squamosal more than four times as long as wide (chat, 61.1; Fig. 9D, dentary bearing over 18 teeth (ehar, 108.15 Fig. 3ND, middle and posterior cervical vertebrae (4-8) with poorly developed diapophyseal laminae (char. 143.1; Fig. SE) and lacking posczygodiapophyseal lamina (char, 142.1; Fg. 3E), and acetabulum partially open with straight ventral margin (ches. 251.1; fg. 96) (see Supplementary information)‘Table 1 ‘Stuopodosieph dinosass fon the Late Tense of South America. Atv “denotes eferal hachas been questioned. Gutbasumsisineued inthe table aldiough maay studies depicted this taxon aban easly theropod or sauischlan (see tex. Taxan/Age Specimens Deiptons Tarp ence PvSSI2, FYB! Serco (1990) = Seen et 2013) Parsee CAPPAUESM Carin a (2018) as Mier ea. (20308) Clvomagsavsnomst PIS) BS aca 2010) Mastin etal 20130) Parag re si 74 Mtns an leer (2000) Matinee etl 2012) Stomnake npg — MEPSEASPY Langer eal 999), Mcp cy Lange (209), 207, 2007 WMCP 354590 ont ern O17, 2018, Ponpairemaeas UIBRAUTOIS cobra eal 2012) Santer CAPPAAUFSM Langer eral 2015) Rapslncinaptonss UEROSVAGDRT Proto ah (28) “Guaararae MeN PV235S ——onpate a (1999) contrat CN P2356. onparte a (2007) MCN PVIOIT2 _——Langerees. 2011) Mawes tepid CAPPAVUSM Mle a. (20188) ote Mater (2020) ote capers ota Longanunetaemrae URAC L108 Lae (2008) MP es: (2019) Mid Novan-Rbaetan Colwadimrws bros PYLSI6T empare 1978) Pvt S904 pet et (202), 2014 Aesemsnns anoles PUL S22 empare (009) Palen Powel (2007) ‘oan mens PYLS8O8 Bonipre (1987), 1972 Pv 809 Ingots pe st 1086 Ape ot a (2018) on the other hand, their anatomy includes a large suite of dinose: ‘ian plesiomorphies thet distinguish them from the plateosaurian sa ropodomiorpls that are highly conspicuous in later faunas (es platwossurids, massospondslids Fg. 6). These include the presence (i most or all Carnisn sauropodomorphs) of features like proportionally long skulls (60% femoral lengths char. 1.0; Fig. 1) with small nares ‘Table 2 Jal of Seth Ameren Bah Siecs 1072021 108445 (char. 16.0; Fig. 3A), distally recurved tooth crowns (char 116.0; ig. 3), medially inset caudal end of dentary toothrovr (char. 97.0; Fig. 9M), presence of palatal teoth, relatively short anterior cervical entra (<2.5 times their dorsoventral helgh har. 131.0; Fig. 9), limited attack ‘ment fo the ium of non-primordial sacral vertebrae (char. 177.0), and narrow proximal end of metatarsal V (char. 399.0; ig. SC) (see Sup- plementary Information) Despite most recent studies agree in the position of Carnian sau ropodamorphs as close to the base ofthe clnde, their precise patter of| relationships nor well understood. Somte sts depict atleast several Carian sauropodonorphs in an early clade of Sanropodomorpha (eg. Ezeura, 20105 Mille ec, 20180; Mller, 2020; Langer e al, 2019). Other stiles, nstesd, depicted Carmian sturopodomorphs #8 & pare phyletie array along che basal nodes of the clade (eg, Maine eal, 2013b; Cabrera e al, 2016; Bronzati and Rankut, 2017; Chapelle and Ghoiniere, 2018; Bronzati etal, 20195 Chapelle et a, 2019; Peyre de Fabrigues et al, 2020; Peyre de Fabrégues and Allain, 2020; Zhang ea, 2020), resembling the result obtained in our analysis (ig. 6). A point of general agreement in most stdies, however, isthe placement of Eoraptor and/or Burioleses as the eerliest branchings of Sa ropodomorpha and the sister group relatonship of Saturnalia and Chromogisaures. Sauropodomorphs have been long regarded as «herbivorous group, bout recent discoveries and studies underscored the putative presence of| camivorous/faunivorous dietary habits in some Carian sat ropodomorphs as a retention of the predatory habits of early saws chlans (Cabreira et al, 2016; Miller et al. 2018b; Bronzati tal, 2017, 2019; Miller and Garels, 2019). Tooth morphology was the most common focus ofthese proposals such as te presence in Burioestes of @ combination of features indicative of carnivorous/faunivorous dietary habits: non overlapping crowns that are posteriorly recurved (Fig. 3D, with small mesial and distal serrations oriented perpendicular to the targins of the crown (Pig. 85; Cabreira et al, 2016; Mller et al 2o18b). Other Carnian sauropadomorphs have been regarded as omnivorous, a possible intermediate stage between early saurischians and herbivorous sturopodomorphs (et, Panphagia; Martinez and ‘Aleobet, 2009), based om the presence of an expanded crown base (ig. SDF) and large denticles that are oblique to the crown margin (ig. SEL), The precise dietary habits of many of these late Carnian axa, however, ate not entirely clear as they eae certain combination of features of unclear dietary significance or morphologies that vary along the toothrow (e.g, Eorptar, Sarwaala; Sereno etal, 20135 Brouzatl cal, 2019, Fig. 3A) Although the dictary assessment of these ently ssuropodomiorpis still inconclusive, multivariate analyses on che entire suite of phyloge neti characters of the feeding apparatus place most Carian species in & rmorphospace region different from that of later sauropodoniorphs (Button etal, 2017; Bronzat et al, 2019; Mller and Garcia, 2019). This suggests thet feding behavior was different from that of Inter Triassic ‘axa, with the notable exeeption of Bauatosaurus (see below). En feet some ofthese taxa (Buriolestes, Eoraptor, Panpadromaeus, Saturnalia) are placed in the morphospace closer to caraivorous theropods than to other Ainosaur Tineages QMiller and Garcia, 2019). Furthermore, the Proportion of chatacters sore diferent inthe phylogenetie dataset forte late Carman sauopodomorphs. Absolute uber of chatacters saved a ferent ase shown between parentheses. Scorags dfrences recalculated only for characters that are cored in the two tata beg compared, s that cataeets With missing ‘eam in at ent one of the two tata ae nor coxted for the otal upon whieh the proportion of eorng ie calelated Powe Cevad Ekneme babes Supaigein —echii —sRayedomms Ave Parag roe © bis@) ona) 0150) oa rst) cov os homers nat oma ozet) 012 oz) oi2i) —aa6t6) os Breyer ene ein —oaean 0 020030) onan ousisa a0) os Papen tebnes’ — 015(10) 0124) O00) b1205) Siogs) 033) ors ‘Simona epge e103) mag) a7) nas) ° outa) oan) ont Burolses ulet o1sa) 012) 05 (32) noe) onan ° 18 25) os Ranaloawseavions —_—_009(5) 04606) 020128) _—_—_—0a8LIS) ena2) sista) oneneuroanatomy of one of these taxa (Sazumalia) shows an enlarged floceuaefosste lobe, a feature thats consistent with the hypothesis of & predatory behavior 2017) Bagualosauris is positioned as more derived than all other late Car: ‘an sauropodomorphs (Fis. 6) due tothe presence ofa larger body size (feauur at least 259% longer than in other Carnian sauropodomorphs; ‘har, 59.1), supracetabular crest extending along pubic peduncle asa thin ridge (char. 371.1, orientation of the transverse axis of the distal ‘end of metatarsal one horizontal angled proximomedially (har. 333.1), More importantly, the feeding apparatis of Baguatosaurs is motpho- spatially positioned closer to post Carnian sauropodomorphs than (0 ‘other Inte Carian species (Bronzat) etal, 20195 Miller and Gare, 2019), Ragualosauras may represent the earliest trend towards large body sizes and omnivorous/herbivorous diets, whieh seem: to prevail among Norian sauropodomorphs. Infact, the similarity with ‘post Carian species might also be a reflection of a possible younger ‘geological age of Bagualasaurus (and Pampadronaeus) in comparison With other late Carian sauropodomorphs. The fauna found at the Bagualoeaumis site shows a predominance ofthe cynodant Exaeretodon (Mller et al, 2015a5 Langer eta, 2019; Miller and Garcia, 2019), asin the mid-evels of dhe Ischigualasto Formation (Exaererodon biozone of Martine et sl, 20138). Conversely, the rhynchosaur Hyperodapeon predominates at sites where all other sturopodomorphs ate recorded both inthe lower third ofthe Ischigualasto Formation (Martinez et al. 2011, 20134) and in the Santa Maria Formation (Langer eta, 2018, 2015), Using this biostratigraphic information, Miller and Garcia (2019) suggested Baquolosaurus and Pampadromaeus may be geologt cally more recent than other late Carninn smutopedomorp 2.2. Barly Norian sauropodomerph fauna Sauropodomorph assemblages from the early Norian are not as well, understood as the two other sasropardomorph faunas from the Triassic of South America (Fig. 1). They are currently only found ina sedimentary sequence deposited in a braided fluvial system in southern Brazil These ‘rocks are recognized asthe Caturvita Formation (sucdrels etal, 1980) oF ‘altematively referred asthe upper part ofthe Candelaria Sequence in the sequence stratigraphy scheme of Hor ct al. (2014). A recent U-Pb radioisotopic age on a single zircon from the Caturrte Formation po: Vided an early Norian maximum age of 225.42 0.37 Ma (Langer etal. 2018, Fig. 2). This unit has provide a diverse assemblage of small-sized vertebrates (Bonaparte etal, 2010) characterized as the Rlograndia ‘Assemblage Zone (Abdale and Ribeiro, 2010; Soares et al, 2011, Fig. 2. ‘The taxonomic coutent of these beds prompted correlation with the uppermost schigualesto and/or lowermost Los Colorados formation in “Argentina (Langer etal, 20078, 2007b; Miller etal, 2015b, 2017), ‘hich aze locally known 6 the Jachalera biozone (Martine? et al, 2018, Fig 2). The uppermost Ischigualasto Formation was dated its type locality at 225.9 Ma +: 0.9 Ma using AY/Ar (Martner eta, 2011, Fig. 2), but recent Uh dates from the top ofthis wnit at another locality yielded a younger date of 221.8 Ma 4 0.1 Ma (Desojo et al, 2020, ig, 2. Although there stil Is some uncertainty regarding the age of these asemblages, current dates constrain them tote early Nori. ‘Three dinosamrs have been named so far fron the early Norian of Brazil (Fig 1): Unaysauras (Leal eal. 2004), Maerocollum (tiller et 2018), and the conflitive Guaibasoums (nterpneted as a sie ropodomorph,theropod, or early saurischian by diferent authors see below lu comparison with other taxonomic groups, dinosaur diversity ‘and abundance is stil low (<58; Martinez ea, 2015: Fig. 6; Romo de Vivar etal, 2020, Fig. 8). Despite their searlty, the early Norian sa ‘opodomorpis from South America currently provide the best infor. mation on sauropadomiorphs faunas coming wp tothe large radition of the mid Norian (see below). Early Norian dinosaurian assemblages are rare worldwide (Mannion etal, 20115 Miller et al, 2017, 2018a), and sauropodontorph tecords putatively from this age ae impreesely dated Jal of Seth Ameren Bah Siecs 1072021 108445 We first comment on the closely related Unaysaurus and Macro: collin, which are definitive sauropodomorphs and subsequently we discuss the implications of the enigmatic Guaibasaurus for this faunal assemblage. Unaysaurs and Macrocllun are similar to each other and share features chat have supported their sister group relationship in some analyses (eg, promaxillary fenestra or foss (Fig. 4A, D), humerus distal end poorly expanded (char: 210.0; Fig. 48), manual phalans 1 longer than metacarpal (char. 2341; Fig. 4F and G; Miller, 2020). The ‘mst parsimonious trees of the phylogenetic analysis presented here (Wig. 6) corroborate the close affinities oF Unaysaurus and sacrocollum reported previously (iller, 2020), Their phylogenetic position in recent analyses has varied in diferent placements close 0 the base of Placeosauria (Miller et, 201895 MePhee ets, 2019) andor Mass: oda (tiller, 2020). All dese positions, however, are close 10 the bsalmost nodes of the Norian radiation of sauropotomorphs (Fg. 6) “The age ofthis radiation is congruent with the only available radioso topic date of the Caturrita Formation (see above), uiderseoring the Importance of this unit for understanding this key stage in. sa ropodamorph evolution, A suite of derived features that have long characterized Inter ssiropodomogphs aetvally make ther fist appear ance in these early Noran species from South America, These elude sand large external nares (cher. 15.1; Fig. 44), non-recurved teeth (ch 115.1 Fig. ¢B) overlapping to exch other (Char. 108.1; lg. 4D), and reduced second distal carpal (char. 220.1) broad distal end of femur (char. 387.1). Even though their affinities with plateosaurians are well established (Le, similarities wih sautopodomorphs of younger Norian faunas), their skeleton shows symplesiomorphies shared wih Carnian tax, such as an elongated rostrum and antorbitl fenestra (chat. 28.0; Fig. 4A, D), and a partially closed acetabulum (char. 250.2) (see Miller eral, 2018). ‘The anatomy of Unaysaurus and Macrocollum inthe eatly Notian of South America is important because it shows final stops in th assembly of the ssuropodomorph body plan. These modifietions include & Auplication in body size respect to Carnian sauropodomonphs (Miller cr al, 20188) and multiple festures that are possibly related to Increasing herbivory (McPhee et al, 2019), Neck elongation, for instance, has long been interpreted as providing a large feeding enve- lope and browsing heights for herbivores (see Barret, 201). Laneeolate teeth with coarse denticles predominate in Norian sauropodoniorphs and have been associated with atleast partial herbivory (Barret and Upchurch, 2007). Finally, large body sizes have also been postulated as related 10 the appearance of herbivory in Sauropadomorpha (Ret fo al, 2011), These South American sairepodomorphs reveal the fstablishment of herbivory and some basle features of the sa ropodomorph body plan in the easly Norian, predating the ecological predominance of sauropodomorphs in later faunas (see below. Guaibasaurs candelariensis was described on the basis of frag mentary skeleton (Bonaparte el, 1999) but is now known from four specimens, Its postcranial anatomy is relatively wellknown and approximately 60% of the phylogeneti characters ean be scored in most data matrices (despite the complete absence of cranial remains). Gu basaurs is smal bipedal dinosaur that resemble Carian saurischians in its body plan and size (Fg. 1) The proposed phylogenetic placement Df this taxon, however, has varied berveen an ently saurischinn (Son parte etal, 2007; Cabreita et al, 2016), an easly theropad (Langer, 2004; Yates, 2007; Langer etal, 2011, 2019; Martinez et al, 2013b), oF fn early sauropodomorph (Ezesrra, 2010; Novas eta, 20113 Chapelle tnd Choiniere, 20185 Chapelle eal, 2019) Irespeetive ofthese alter native positions, Guaibasauru salen’ depicted close to basal nodes of these clades du co its combination of multiple plesiomorphic features. ‘The phylogenetic analysis presented here retrieves this taxon allied with ‘Theropoda but with minimal support (see Supplementary Information). Regardless the uncertainty conceming its phylogenetic positon, Gua basaurus exemplifies that Norian faunas in South America included not only dinosaurs chat radiated during this time (es. plateoseurians, reotheropods), but also representatives. of earlier dinosauromorphFig. 4. Materials of selected sauropodomorphs ftom the easly Norian of South ‘Ametica. Macroollon faqu A, skull (CAPPA/UESM O001a) (versed) 8, “dentny tooth (CAFPAAIFSMOOO1a in teal sew, and G ete vertebia in ater view (reversed, scale brs equal 5 em (Ay C) ad 3m (8), Urey srs tle (UESM 11065) D, maxilla in aeral view (reversed), E ight humerus in anterior view, F, metacarpal in dorsal view, G, anal phalanx 1 in doa view, scale bar eq 3 en (D) ad 2m (EC). Jal of Seth Ameren Bah Siecs 1072021 108445 lineages that diverged during the Anisian-Carnian (e.g. lagerpetis, silesaurids; Martinez etal, 2016; Langer and Ferigolo, 2013), 2.9, Mid Norian-Rhaston seuropodonomph fauna ‘This fauna includes numerous sauropodomorph specimens that have been found in the upper levels ofthe Los Colorados Formation (Bons parte, 1972) and a suite of specimens more recently discovered in the Quebrada del Barro Formation (Martinez etal, 2015; Apaldett et al, 2018) The Los Colorados Formation isa favial sedimentary sequence of over 600 mi thar overlies the Isehigualasto Formation (Caselli et al. 2001). This unit contains the best known dinosanr dominated faune from the Triassic of South Amtericn, This status stems fom the easly work by Bonaparte (1972), who reported a diverse vertebrate assent blage ftom the top levels ofthis unl, Known as La Esquins local fans Gonapart, 1982). This fauna included three very distinet sa ropodamorph taxa: Rigasairus, Coloradisaums, and Lescennsaurus. The ‘age ofthe top ofthe Los Colorados Formation was regarded as Norian Cc 4 Bonaparte, 1982), or more generally latest Triassic (eg., Arcucei ©, 2004), based on vertebrate biostratigraphy and comparisons with “Triassic faunas of other continents (e.g, lower Eliot Formation in South Africa; Bordy etal, 2020; Viglitti etal, 2020). Mare recently, paleo: magnetic data was used to correlate Los Colorados sequence with the CChrons E7+ to E15u ofthe Newark polarity imeseale (Kent et al., 2014) ‘The age of Los Colorados Formation was estimated by Kent et sl. 201) {o range between 227 Ma to 213 Mea, assigning an age to the La Esquina fauna as no younger than 213 Ma (mid-late Norian, Fig. 2). More recently, Desojo etal. (2020) noted the age of Los Colorados Formation could be several million years younger based on new U-Pb dates of the "uppermost Ischigualasto Formation (221.8 Ma + 0.1 Mi Fig. 2) other wn that has provided sina saopodomorph materia s the Quebrada del Barro Formation fom the Marayes-E Carrizal basin, also located in northern Argentina, arte etal, (2015) described & diverse vertebrate assemblage from this formation, including cynodonts, turtles, sphenodontians, pseudosuchians, pterosaurs, and dinosaur morphs. Its nique faunal composition, predominated by opistho- dontian sphenovionts, tritheledontid” cynodonts, and early smuropodomionpis, led to the tentative assignment ofthe Quebrada del Barro Formation Co the late Noria-Rhaetian (slighty younger than Los Colorados Formation; see Martinez ea. 2015).Sauropodomorph from the Quebrada del Barro Formation have close affinities with those af the Los Colorados Forniation, which we recognize here a the mid Nor jan-Rhaetian seuropodomorph fauna, Sauropodomorphs from the (Quebrada del Barro Formation inchide several istnet taxa, but so fa, {ie only formally deseribedssuropodonorph is Hrgentia prima (Apaldeat eral 2018). ‘There are four named mid Norian-Rthaetian sauropodomorphs in these units (Riojasaurus, Coloradiseurus, Lessemsaurus, and Ingenta; Fig 1), which comprise 308 ofthe known sauropadoniorph diversity in lhe Triassic of Son Ameriea (Table 1). During this time the taxonomic diversity of sanropodomorphs approximately matches that of other tetrapod groups (Le, aetosaurian pseudosuchians, cherapsid synapsis; (Otero etal, 2019), However, in contrast with previous faunas, the mid Norian-Rhetian sauropodomorphs from South Ameriea were conspie ‘ious components of their terrestrial ecosystems and predominated 1s the most common large herbivores (Bonaparte, 1982; Benton, 1983). In the Quebrada del Barto Formation sauropodomiorph specimens const tute approximately 20% of the collected and identifiable fossils (Mar inex el, 2015: Fig. 6). The abundance of sauropadomvorph dinosaurs n Los Colorados Formation Is even higher, exceeding 75% (Martinez cal, 2015). This is remarkably higher than in earlier sauopedomorpa faunas, as exemplified by the detailed census inthe Ischigualasto For tution that showed only 1.5% of the identifiable vertebrates were sauropodomorphs (Martinez etal, 20138). ‘The predominance of sauropodomorphsis similar to that of the lower Eliot Formation of South Africa, in which Ueirabundance has Deen recently estimated at 80% (Bory etal, 2020). ‘The striking similarity of the sauropodomorps dominated faunas of Los Colorado and the lower Eliot formations have long been noted Ce. Bonaparte, 1982), and recent improvements i the bistratigeaphy and ‘chronostratigraphy ofthe latter nit constrained its age between the mi Notian and Rbaetian (219.9 Ma to 204.9 Ma; Scseio eal 20175 Bory et al, 2020; Vigiot etal, 2020), Other Norian-Rhaetian terrestrial ‘sequences from northern and southem Ind masses (eg, Inds, central Europe, Greenland) also have sauropodomorphs as the predominant large herbivores during these times, in particular those Ioeated at high paleolatiudes (e4., >30") In turn, sauropodomorphs sem: tobe absent from Norian deposits at Tower paleolatcudes in North America (8 "upper Chinle inthe Colorado Place, Newark Supergroup; Olsen et 20025 Irmis eta, 2011; Remezani et al, 2071). Tis difference may be ‘due tthe position ofthe former localities in temperate lnunid belts and the latter localities in tropical arid belts predicted by paleoclimatic models (Kent etal, 20148 Fig. 6) Other dinosanromorph lineages recorded along with dhe mid Nor fan-Rhaetian saropodomorphs from South Amerien inchide three neotheropods (Zupaysaurus, Powelvenator, Luclanovenator; rice and Corie, 2003; Ezcurra, 20175 Martiner and Apaldet, 2017) and the lagerpetd Dromomeron (sari, 2016). In terms of abundance, these are minor components of the vertebrate assemblages (<2.59%: ‘Mastin et al, 2015) and ornithischians are so far absent altogether (although their ghost lineage extends back to the Carnian; Pacian, Jal of Seth Ameren Bah Siecs 1072021 108445 2012). The low diversity and abundance of theropods during this time is probably duet the naturally ow abundance of predatory species and to the existence of pseudostchinn archosturs groups that occupied cara vorous/faunivorous niches. Ths pate is ikely «global phenomenon, asics paralleled in eoeval terrestrial deposits from North America and Europe (Brusatte eta, 2010; Langer eta, 2010). ‘The sauropodomorphs from Los Colorados and Quebrada del Barro formations are South American representatives of the large Norian ra Aiston of sauropodomorph dinosaurs (Fig. 6). large mumber of line tages of sauropodomorphs originated throughout this radiation, Inchuding major inclusive clades such as Plateosauri, Massopeda, and Sauropodiformes, and the much less inelusive (ally evel Linnean categories) groups (Pig. 6). The deseribed taxa from the Los Colorados 1nd Quebrada del Barro formations are representatives of three ofthese less inchisve groups: Riojasmuridse, Massospondytidae, and Lessen sauridae, Other unnominated specimens have beea reported and may represent an even larger phylogenetic diversity, but their fragmentary nature (Ezeurrs and Apaldet), 2012) and/or undescribed status (Mia ‘inex et al, 2015) precludes their formal treatment at dhe moment. We discuss here the thre lineages mentioned above an comment on basic ‘aspects of thei paleobiology and relationships Rigjasaurusincerus was the first seuropodomorph to be described from Los Colorados Formation. Its holotype is relatively complete Posteranium (Fig. 1; Bouapare, 1967, 1972) and multiple referred specimens include posteranial materials (onaparte, 1972) and a sub- adult specimen with a complete cranium (Fig, 5 Bonaparte and Fig. 5. Matas of selected sav ropodomorphs from the mid Nor fan-Rhaetian of South. Americ. Ingeta ‘rina (PVSI 1086) A, cervical vetebrze in Jateral view (reverse) and Blanes in anterior view. sale bar equals 10 ent Les Semsaurssaropades PUL. 4822) C,antecor cervical newal ail la lateral view, D, bu eras in anterior view, ad Ey metacarpal in docsal vie, scale bas egal 3,6, and 3 fem (iespeetvely); Colorasanrs bres F ‘mld cervical vertebrae (PVL 590), 6, shall (PVL 3967) in lateral view, and, bunerss (PVL 5008) in antesir view, scale bass equal 2 em; Rigjsuras ince, Lzeferted skull (PULR 56) in lateral view and J, humerus (PML 3808) in anteir view (reversed sale ‘brs equal Sc aed 10 cn, respectivelyPomares, 1995). Rigjasaurus was allied with the two species of the gen ucnemesaurus fons the lower Elliot Formation of South Aftca (ates, 2007; McPhee et al.,2015a). Their affinities have been confirmed in most phylogenetic analyses and this clade fas been referred as Rioja ‘suridae (Yates, 2000). Potential records ofthis family have been re pported also from the Quebrada del Barro Formation, including an articulated pes originally referred to Riojasurus (Boss and Bonaparte, 1078) and new individuals (PVSJ 959, 907, 980; Martinez etal, 2015: Fig. 6) that resemble Riojascurus in averall morphology although they ‘ear somie minor differences that possibly imply they eannot be referred to Rijasaunis ince. All these remains share overall heavy constr: {Yow that indeate rojasaurids were large and robust sauropodomionphs that inhabiced in dlfferent regions of Gondwana. The lnib measure ‘ments and proportions of the most complete skeleton ofthe group (type of Riojasauus, see Figs. 1 and 53) have been used to estimate a body sass of 2.28 tons and to infer quarupedal locomotory habits (McPhee ct al, 2018). The skull anatomy isso far only known in an immature specimen (PULR 56; Fig. SD thar Bonaparte and Pumares (1995) referred co Rigjasaurus incertas, although some authors have recently casted doubts on the taxonomic identity ofthis specimen (McPhioe and ‘choinlere 2017), Inespeetive of the taxonomic identifiation of this specimen, an important feature of the skull PULR 56 is that it bears a unique combination of eraniodental features that separated It from sll ‘other Norian sauropodomorphs ina biomechanical morphospace anal ysis (Button etal, 2017). Rlojasauridae has been retrieved as a basal massopodan (Pi. 6) in multiple analyses (eg, Yates, 2007; Otero and Pol, 2013; Apaldett ‘tal 2018) slightly more derived placement was recently proposed (tePhee etal, 2018; Miller, 2020) that derives ftom excluding the Jal of Seth Ameren Bah Siecs 1072021 108445 cranial information ofthe referred specimen PULR 56, which bears some plateosaurian symplesiomonphies that are absent in massospondylids ud more derived sauropodoniorphs. These include a rounded subuarial foramen (char. 18.0), the anterior margin of antorbical fossa formed by sharp and elevated maxillary rge (char. 30.0; Fig. 5D, elongated anterior process of lacrimal (char. $9.0; Fig. SD), and a dorsoventrally low jugal below orbit (har. 51.0; Pig. SD. Despite this uncertainty, under both scenarios rojasaurids represent a relatively early branch of| the mid Norian radiation of Sauropodomorpha (Fg. ), represented by robust quadrupeds chat thrived in the latest THassic of western Gondwana CColoradisaurus brevis is known from a near-complete skull (ons: parte, 1978 Apaldett eta, 2014, Fi. 5D) and a fairly complete post tania skeleton (Apaldett etal, 2012). Goloradisaurus was likely @ facultative biped and the femoral circumference (Fe) of the largest specimen (PVL 5904) implies a body mass of approximately 370 kg (Fe = 191 my using the quadratie equation for non-avian bipeds of Can plone etal, 2014; see Peyre de Fabregues nd Allan (2020) for ad ‘ional comments). The relatively small boy size and gracile body plan of Coloradsaurus (with a proportionately short rostrum, extremely clongated cervical vertebrae, and slender limbs Fig. 9C-E) clearly sets this taxon apart from che body plan of other sauropodomorphs from the nid Norian-Rhsetian of South America, as well as from those of the similar fanna recorded in the lower Elliot Formation (MePhee et a, 2017). Coloratsauras vas originally allied to plateosaurids (Bonaparte, 1978; Galton and Upchure, 2004) because the sul ofthe type material bars a fre characters that are so far only kuown in this group (e.g, eescente and broad lamina of maxillary antorbiea fossa (char. 31.1; Fig, 5D), infratemporal fenestra does noc extend beneath the orbit (car. Fig. 6 Piylogesetic relavonships of Sav poder calibrated aginst Googie] Tine Seale. The topology shown a rede strict consensus obtained by ignovng sx ‘unstable taxa (See Supplementary Informs tion for furer deals), Clade names in cared by an atow are stembased phylogenece definitions aud those marked ‘witha black eile ate node based defi tions. The two alternative definitions forthe lade Sauropoda are indicated with YO7 (fates, 2007) and $97 (Slgato etal, 1997. ‘Tree calibration and pling used Paleouee (Gaps, 2012) and Strap (Bell att Loyd, 2015) R packages. 5 a a z 556.0), base ofthe basipterygoid processes and the parasphenoid rostrum ‘set below the level of the basal tuberae (char. 80.1}; see Apalde't eta 201). The posteranial skeleton, well preserved in de referred specimen, PVL 5904, contains a lrger number of similarities with mass: spondylids like the notably long mid-cervical vertebrae (char. 130.2) 1g. SC) and this information resulted in its phylogenetic placement ‘within Massospondylidae (Apaldett e ala, 2012), The incorporation of the portcranial anatomy of Coloradisaurus has resulted in the placeaient ‘of this taxon as the oldest massosponylid in most phylogenetic ana Iyses. tts inclusion in this clade pushes the diversification of mass. spondylids into che Late Triassic of South America (Pig, ©, asthe rest of the clade has « widespread distribution inthe Barly Jurassic (Chapelle and Choiniere, 2018; Chapelle tal, 2019) {essemsanride i the third sauropodomiorpa lineage recorded in the mid Norian-Rlhsetin of South Amerion andthe latest ca be recognized and named, Lessemsaurs sauropodes from the Los Colorados Formation was the first member ofthis lineage to be described, based on a ceri ccodorsal series (Bonsparte, 1999). Later, additional posteranial mate tials of dhe type material ofthis taxon was described (Pol and Powell, 2007), revealing multiple anatomical differences with other saw ropodomorphs from this unit and also revealed derived characters shared with Auttonirus from South Africa (Yates and Kitching, 2003), ‘sch as the nvably bros and shore metacarpal (char. 227-3; Fig. 5H). Lessenscuris nnd Anteonims were the only two kaown menbers of thiselade til recently, but new discoveries and studies have show the diversity of this group was higher than previously thought. Fest, anew lessemsaurid fom South Americ, Ingeniaprina, was recently described from the Quebrada del Barro Formation (Apaldeti eta, 2018). Soon ‘after, the lessemsauridLedumahad! was described from the Early Jurassic of South Affiea (McPhee ot al, 2018). Finally, two additional large bodied taxa from the Late Triassic oF southern AMtica, Meroktenos oyre de Fabrégues and Allain, 2016) and Kolunoluno (Poyre de Fabrogues and Allain, 2020) are reinterpreted by our analysis as mem ‘ers ofthis group (ig. 6; ee Supplementary information) as they share ‘with lssemsaurids relatively short cervical vertebrae (char. 130.0), high dorsal arch pedicels (char, 168.1), fourth woehanter located at midshat (char. 292.1). Lessemsaurids have recently provided important data for under standing the origin of gigantism among sauropodomorphs, as some of the new materials have distinctively large body size (pale etl, 2018 MePliee etal, 2018). New speciniens of Lessemsaurus confirmed the type material was immature and indicated the body mass of this taxon could have reached wp fo 7 tons (Apaldeti et al, 2018), This ‘approsches the 10 tons of ently eussurapods nnd is more than twice the size of other Triassic sauropodomorph linesges. Furtheremote, the recently described Ledumahadi yielded an even larger body mass est mate of 12 ons (MePhiee es, 2018), indicating lssenismurds achieved truly gigantic sizos by the Early Jurassic. Apaldet et al. 2018) noted that leseemsanrids attained gigantism through an acceleration of their ‘growth rate but maintaining @ plesiomorphic eyelieal growth pattern, lffering from the continuous growth strategy of giant eusauropods. Furthermore, lessemisautids show relatively short neck (Fig. SA) but Well developed cervical pneumatic cavities in their neural arches, ind cating an improved avlanstyle respiratory system (Apaldetti et al., 2018). These features, along withthe previously known high vertebrae indicative of large epaxial musculature aad robust limbs, were Likely ‘rtieal for the early appearance of gigantis in Sautopodomonpha. The anatomy of lessemsaurids is significant due 10 he presence of rmeros derived similarities with eusanropods in the vertebrae and limbs: high neural spines (char: 198.0 Pig. 5A, F) and neural arches (char 183.1), low deltopectoral erest (Fg. 58, G), and large pubic plate (char. 261.1). Features are absent inal other Triassic sauropedomvorphs ‘and have supported the position of lessemsanris as closer fo eusaro ‘pods than to all other taxa traditionally regarded as non-sauropods in autipe analyses (Yates, 2007; Pol eta, 2011; Otero and Pol, 20135 ‘Apaldett etal, 2018; McPhee ta, 2015b, 2018). This led Yates (2007) Jal of Seth Ameren Bah Siecs 1072021 108445 to emend the phylogenetic definition of Sanropoda, under sshich les semsuids are interpreted as the enlist branching of the stem defined smuropod clade (ee Fig. 6). The original node based definition of Sa ropoda (Salgado el, 1997) is much more restrictive, points tow node adjacent co Eusauropoda, and tothe same node defined as Gravisauria by llain and Aquesbi (2008). ‘The onset ofthe predominance and abundance of sauropodonorphs in the Norian vertebrate assemblages have long been regarded as @ landmark inthe history of inasaurfannas in South America Bonaparte, 1982; Hlenton, 1989). Advances in recent years have Inereased oe appreciation of the morphological disparity of the sauropodontorphs recorded during ths ime. The three lineages currently recorded in tis fauna ate chatacterized by body slaes that range over an order of ‘magnitude (370-7000 kg), have ver distinct body plans dfering in the longation of the neck, limb proportions and robuisticiy, and bipedal versis quadrupedal locomotion. Our current knowledge on the skull my of sauropodomorphs ofthis fauna is Limited to two skulls of the Los Colorados Formation (see above), but thelr anatomy indieates the ‘aceupation of disparate regions of the Fnctional mexphospace (Bon er al, 2017; Bronzati et al, 2019). The mid Norian-Rhaetian sat ropodomorph from South Ameriea clearly show one ofthe earliest cases of niche partitioning among herbivorous dinosaurs. & similar pattern has been noted for the lower Elliot Formation (McPhee et a., 2017), Uwhich is unsurprising given the similacy of dhe faunas nnd the shared presence of certain sauropodomorph lineages inthe Norian-Rheetian of South Antetica and southern Aftica. These two closely related and geographically proximate faunas of southem Pangea have been pivotal for recognizing the ecologieal diversiiention and preominsnce of sauropodomorphs prior (othe end Triassic (Benton, 19835 Barve and Upchurch, 2007; Langer etal, 2010). Many of these lineages sucess fully pass the TrissieyJurassc extinction event and farther diversify in the Early Jurassic throughout Pangea (Olsen tal, 2002), as recorded in South Ameen (Martinez, 2009; Apaldetti etal 2011; Pol etal. 2011, South Aiea (Bory eral, 2020; MePhee et al. 2017), India (Kuty etal. 2007), North America (Olsen et al, 2002; Marsh and Rowe, 2018), (hina (Wang a, 2017), and Antaretica (Smith snd Pol, 2007), 2. Discussion ‘3.1. Souropodomorph plylogenetics: recent progresses and gereements One of the major problems for understanding the evolution and di versity of Tiasi sauropodoniorps were their Iongstanding enficting phylogenetic affinities. A phylogenetic perspective provides valuable Insights on studies of past diversity and faunal changes (Norell, 1998; Edgecombe, 1993), sueh as the steps involved in the radiation of clades nd the time of appearance of lineages or sponorpies, Although the monophyly of Sauropodomorpha was never ques: tioned, basic aspects of the relationships between their early members Were telear (@ monophyly of Prossuropoda), Fortunately, some trons of agreement have now been realied thar define basi patterns of the evolution of snuropodomorphs. This agreement isthe resule of an Intensification of studies on early sauropodomoxph piylogenetis, largely prompted by discoveries and description of specimens from out Ameriea ad South Alien (eg, Leal etal, 20045 Yates, 2006, 2007; Martinez and Aleober, 2005; Yates eta, 2010; Ezeurra, 2010; Knoll, 2010; Apalderti eral, 2011, 2012, 2018; Cabrera etal, 2011, 2016; Pol et al, 2011; Otero and Pol, 201 Martinez eta, 20130; MePhee et al, 20152,b, 2017, 2018, 2019; Otero eta, 2015; Peyre de Fabrégues and Allan, 2016, 2020; Bronzat and Rauhut, 20173 Chapelle and Choiniere, 018; McPhee and Choiniere, 2017; Miller etal, 2018a, by Pretto etal, 2018 Bromzati et al, 2018, 2019; Langer eta, 2019, Chapelle etal 2019; Miller, 2020). There ae five points of agreement reached in recent years that are most relevant to Triassic ta: 1) the parapliyletic nature of Prosauropoda with respect to seuropods; 2) the placement of small bodied Carian texaat the base of Sauropodomorpha(outside Plateosauria); 3) the monophyly of four family-level clades: Plateosauridee, Riojasauridae, Massospondylidae, and Lessemsauridaes 4) the basal position of Plateosauridae relative to other three families; 5) the postion of Lassemsauridae as closer co ensauropads chan the other three families; Fg. 6). ‘The South American fossil record of ssuropodomorpis plays a crt ical roe in these advances, Firstly, the recognition of sauropodoniorphs in the late Carnian of South America (eg. Sauruaia, Panphagia) that ‘were unquestionably basal to plateosaurids and niassospondytids (ig, 6) helped in breaking the dichotomy berween Prosmitopoda and ‘Sauropoda within Sauropodomorpha, Secondly, the discovery of les semsetrids aud thelr auatomical information (rom South Amerie and ‘South Africa) elartied that some robust ‘Trini sauropocomoxphs were ‘certninly more closely elated to evsaropods than the mre lightly bail plateossurids and massospondylids (Pol and Powell, 20075 Yates, 20075 “MePhee etal. 2015), This in umn also revealed chat the acquisition oF ‘sigantism and quadrupedalism could be traced back to the Late Triassic, ‘during the earliest stages (or before) the origin of Sauropoda (Apslett ‘etal, 20185 MePhew etal, 2018), 8.2. Sauropodomorph founas: the inpact of recent discoveries ‘Sauropodomorphs from the Trnssc of Sth America only played aa Important role in studies of dinosaur early diversification ad faunal changes after the discovery of the Los Colorados fauna (Bonaparte, 1072). However, discoveries of Triassic sauropodomorphs from South ‘America were searce in the 30 years that followed Bonaparte's discov feries inthis formation in the early 1970's. Coloradsaurus (Bonaparte, 1978) and Lesemscurus (owaparte, 1999) were formally described well after Bonaparte (1972) but their discovery dates back to the early 1970's. Tao notable exceptions in the 30 year-long gap of discoveries were Eormpror (Sereno ea, 1998) and Saturnalia (Langer eal, 1999), although the forse ves regarded for mnny years asa theropod and both taxa were only briefly reported at that time (with detailed anatomical studies on them published such more recently) This situation changed radically due to new discoveries made in the last 15 years that put souropodomiorpis from the Late Triasic of South America back in the limelighe ‘Our knovledge of the Ite Carian sauropodoniorph fain increased, drastically over the last 15 years. During this time five out ofthe seven noun taxa were discovered and the 1vo previously known taxa were desribed in detsil. A thorough monograph vine published on Eoraptor (Gereno e€ al, 2013) and a series of papers on Satwnatia documented ‘many aspects ofits anatomy and paleobiology (Langer, 2003; Langer ‘etal, 2007; Bronzati etal 2017, 2018, 2019). These discoveries and ‘studies allowed for the recognition ofthis early stage in th evolution of ‘Smuropedonionpha, chatseterized by n rather Inrge taxonomic diversity in comparison with their much more moderate morphological disparity (Langer eta, 2019). One ofthe most recent and significant discoveries was that of Baguaosaurs agers that inereased the body size range ‘and the ecomorphologieal morphospace of Carnian sauropodonorphs ‘approaching those occupied by the Norian taxa (Pretto al, 2018 Bronzat et al, 2019) ‘The early Norian seuropodomorphis were completely unknown prior to the intensive research on the vertebrate fans ofthe Caturtita For tation that started inthe turn ofthe century (Bonaparte ets, 199 2001, 2003). Definitive seuropodomorph dinosaurs were limited t0 ‘naysaurs (Leal et al, 2004) wnt the discovery of the beautifully preserved and complete specimens of Macrocolla (Miller etl, 20188, Miler, 2020). The latter taxon will landmark our knowledge of early souropodomiorpis ofthe early Norian due the paucity of the dinostrian fossil record from this time at a global scale, highlighting once again how recent discoveries are highlighting the importance of Triassic sa ‘opodomorphs from South America, Inthe case of the mid Norian-Rhsetian sauropodomorp fan fom South Amerie, no new lineages or clades have been discovered in the Jal of Seth Ameren Bah Siecs 1072021 108445 past 15 years. Progress has undoubtedly been made through discoveries Of new taxa that belong to previously Known lineages (e.g, lessen sourids, riojasauriés) and through studies that revealed evolutionary hovelties chat appeared during this time. New discoveries inehude the promising faa of the Quebrada del Barto Formation (Martine? etal. 2015; Apaldet etal, 2018), which reeords a sauropedonorph fauna that is related but distin ro that of the Los Colorados Formation. An important advance vas the recognition that the oldest giant sau ropodamorphs appeated during this ime, concomitant with the devel opment of axial pneumaticty and large epasial musculature (Apaldeat eC al, 2018). Furthermore, these new renuans strengthened the close link between the South American seuopodomorphs from: this fauns with those fom the lower Elliot Formation of South Africa (MePhiee cal, 2018; Bordly et al, 2020; Viglet e al, 2020). 3.3. Radioisotope dings: large scale correlations ane dng the faunal Recent advances in radioisotopie methods are having a deep impact, ‘on paleontological studies of continental sequences and fortunately the Lae Triassic of Soult Ameriea has not been an exception. ‘The eto nostraigraphie control of most sauropodomorph bearing sequences is now reasonably well constrained and this has allowed for more precise correlations and determination of significant evolutionary events, The Ischigualasto Formation was the fist of these units to be radioisoopically constrained (Rogers et al, 1999) and more recent studies have improved the precision of the age of the Tate Carnian dinosaur bearing beds at its type locality (251.4 + 0.3 Ma; Martinez cal, 2011) and ether auterops (229.20 4.0.1 Ma through 226.85 1.4 Maz Desojo et al, 2020, Fg. 2). Dates obtained forthe Satumnalia locality of the Santa Matin Formation provided maxinuim depositions age (233.25 + 0.73 Ma) that indicates the dinosaurs from. bodh units are probably approximately coeval (Langer etl, 2018; Desojo tal, 2020, Fig 2). The temporal correlation ofthese units was expected, but these dates validated traditional biostratigraphic corelatons between them based on their vertebrate fauna (Barberens, 1977; Bonaparte, 1982). Furthermore, the new radiometric dates of the Very distinct fatal assemblage inthe underlying Chaiiares Formation are only three to five rillion years older (ca. 296 Ma; Marsicano et al., 2016; Ezcurra et a. 2017), which constrains the timing of the enset of the peculiar Ischigualasto Santa Marla fauna that records the earliest sal ropodamorphs. The upper extent of the Inte Carnian saropodonsorpa fauna is less precisely constrained due tothe lek of dinoser records in the mid t0-upper levels of the Ischigualasto Formation, the top of which was frst dated at 225.9 + 0.9 Ma (Martine et al, 20138) and more recently at 221.8 Ma 0.1 Ma (Desojo eal, 2020, Fig. 2). The single svailable date from the Santa Maria Formation only provides ‘axinium depositional age ata single site, but future efforts may expand this to other localities and may help with determining the temporal extent of ths fauna. For instance, resting the biostratigrapheally based hypothesis that dhe site of Baguaasaurus ad Panpadromaeus is actually younger than those of Sanumalia and Burolsts (Miller ea. 201585 Langer etl, 2019; Miller and Garcia, 2019) ‘The publication of the new early seuropodomorphs and the radio Isotople dates of the Caturrita Formation occurred slinost simul neously (Langer et al, 20185 Miller et al, 20188). The maxinaam depositional age of 225.42 £0.37 Ma froma single zircon (Langer etal. 2018) only provides a lower Doundary for this unit, which may be equivalent to the top of the Ischigualasto Formation as suggested by vertebrate biostratigraphy (Langer etal, 20074, 20071 Miller et a 2015}, 2017, Fig 2). Thus, the fist fal change in sauropodontorplis (hich included the acquisition of herbivorous adaptations) occurred at some point in the eaely Norian, about eight million yers late than the record of the previous sauropodomorph fauna. Prior to these recent contributions the sauropodomorph components of this unit were known from limited material and the age was solely based on large scalebiostratigraphie correlations. Several important questions are still open ‘about this fauna, sch as the relative abundance ofthe afferent dino saurian components or the changes in sauropodomorph faunas "houghout the frst half of the Norn. Finally, regarding the latest sauropodomorph fauna of the Triassic, recent studies have also impacted our understanding on the time at ‘hich dinosaurs predominated in terrestrial ecosystems for the frst time. This event can now be placed a the mid Norian based on advances ‘mace both in South Amerie and South Africa. The paleomagnetie data ‘gathered from Los Colorados Formation sugested the sauopodomorph fauna from the top ofthis unit (ie, the local La Esquina fauna sensi Bonaparte, 1982) was no younger Han 213 Ma (Kent etl, 2014 Fig. 2) bur other authors noted this fauna could be even younger Des0j0 et, 2020, Fig. 2). This is roughly congruent with recent radioisorpie dates from the lower Elliot Formation (219.6-204.9 Mas Hordy etal, 2020), ‘which contains a very similar fans. The long timespan of the lower Eliot Formation throughout the mid Norian and Rletian coupled wit the continuity of he sauropodomorph record throughout ts seamen. tary sequence (McPhee ot al, 2017; Bordy etal, 2020) is pivotal for recognizing the relative uniformity of the mid Norian-Rhaetian se ‘opodomorph fauna of southern Gondwana, The sauropodoniorphs fom Los Colorades and the Quebrada del Barro formations thus represent South America members ofthe sane fanna that spread ehronghoue SW Pangea during approximately 15 milion years. The mid Nor lan-Rhaetian sauropodomorph fauna would not only be distinguished by the predominance of this group over other vertebrates (Bonaparte, 1982) but also by its duration in the Late Triasi, which may have been extended over a period of ine thac potentially Is up co three times longer than the duration ofthe previous faunal assemblages 4. Conclusions ‘Our knowledge on the seuropadomorph fauna from the Triasste of South America hasinereased significantly inthe las 15 years Based on 8 ‘review of available information we underscore here hovr important the South American sauropodomorphs have been for understanding the origin and early diversification ofthis group as wel as for exemplifying their predominance over vertebrates dnring the Late Triasi. ‘Three distinct sauropodomorph faunas are recognized inthe Triassic terrestrial sequences of South America: late Carninn, early Norian, mi NNoriatt-Riaetan, The fst cwo lave no parallels outside this continent ‘and records relatively short and successive periods of ime (en 3-8 MY), “The mid Norian-Rlssetisn smiopodomorphs, in contrast, have remark ‘ably close similarities with those recorded inthe lower Elliot Formation fof Sout Afri (evince et al, 2017), which is now well constrained _geochronologically (Sordy eta. 2020) and extends over approximately lon years up to the end Triassic: Sauropodomorphs are nique among dinosaurs not nly du to thee ‘characteristic body plan and gigantic body sizes, bur also because during ‘he firs 30 millon yeats of dinosaur history, thei diversity, abundance, ‘and predominance in terestrial faunas were far higher than those of ‘ther dinosauromorph lineages. The South American record has been the centerpiece in defining this pattern in de past decades and will likely keep providing new data on the ssaropodomogphs that thrived in “Triassle ecosystems inthe years to come. Declaration of competing interest ‘The authors declare that they have no known competing nancial interests or personal relationships that could have appeared to influence the work reported inthis paper. Acknowledgments ‘We would like to thank the editors ofthis specialise forthe invi tation to partielpate, We would like 10 thank A. Ribeiro (MCP), M. Jal of Seth Ameren Bah Siecs 1072021 108445 Langer (USP), C. Schultz (UERGS), L Leal (UESMD, B. Vaceari and 6. Cisterna (PULR), and Jaime Powell and Pablo Ortiz (PVL) for access co material under their care, We are grateful to R. Miller for providing photographs of certain specimens used in figures of this manuscript. “Thanks ate also extensive to M, Bronzati and an anonymous reviewer whose comments greatly improved this manuscript. We would like to tank the support from the Agencia de Promoeién Cientifico y Tee nolégiea grants PICT 2015-0504 and 2016-0236. References Abt, F ibe, A, Sula, CL, 200A eh eyoton far of Sata Ce os (Suns Mara fran mene la) outer ral Nee aback Gea (Therapie, Cynodents) in Gondwans, Palacogceg,Palacoclimatol Palseoeol en Rol 207. ype emma of amine Pract 1, aS. Alin, AqueN, 2008 Antony snd phylopentic relia of Todas in Deri Stopes) fom he te ny Jr a Merce, Geotiese ‘Andre, RRL, Boul, G., Montardo, DK, 1980, 0 Grupo roxio de su (Teste) io Gane do ul In Anais o OK Congreso Base de Geog. Cambor 2, ples. Mane, RN, Aeoer, OA Po 201. 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