http://dx.doi.org/10.18268/BSGM2019v71n3a6 P. 727‒739 VOL. 71NO. 3 BOL. SOC.GEOL.MEX.2019 S.L.P., Mexico. Zona Universitaria Poniente, San Luis Potosi, Potosí,ma deSanLuis Dr. Manuel Nava 8, Facultad de Ingeniería, Universidad Autóno Lizbeth Nava-Rodríguez Mitras, 64348,Monterrey, N. L.,Mexico. Villa Nápoles 6506,ColoniaMiradordelas Vivas-GonzálezRafael 29000, Tuxtla Gutiérrez, Chiapas, Mexico. Natural. Calzada de los hombres ilustress/n, era”, Secretaría de MedioAmbiente e Historia Museo dePaleontología “Eliseo Palacios Aguil [email protected] Gerardo Carbot-Chanona Saltillo, Coahuila,Mexico. Desierto, Carlos Abedrop Dávila3745,25022, de Paleontología,Departamento Museo del Fernando Cabral-Valdéz Héctor E.Rivera-Sylva Fernando Héctor E. Pueblo Formation(Campanian),Coahuila,Mexico The firstcrocodyliformsremainsfromLaParritalocality,Cerrodel Manuscript accepted: April 21,2018. Manuscript April accepted: Corrected manuscript received: 10,2018. April Manuscript received: January 5,2018. Cabral-Valdéz Rivera-Sylva , Gerardo - - Carbot-Chanona Coahuila, Mexico. Globidonta, Late Cretaceous, Keywords: Crocodyliforms, during theLate Cretaceous. for the Cerro del Pueblo Formation on adelta plain with tropical climate environmentsfor stagnant to fluvial charophyte oogonia provide evidence odyliforms, turtles, dinosaurs, and croc association The of osteoderms. on one isolated tooth, vertebrae, and Cerro del Pueblo Formation, based remains from form La Parrita locality, this work we crocodyli report the first chians, and eusu goniopholidids, indeterminate last group isrepresented only by dinosaurs, andcrocodyliforms. This Coahuila, includes turtles, pterosaurs, (Late Cretaceous, Campanian), in the Cerro del Pueblo Formation land tetrapods of recordThe of ABSTRACT Brachychampsa montana Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín , Rafael Vivas-González . In - - - tortugas, pterosaurios, dinosaurios y tardío, Campaniano) enCoahuila, incluye del PuebloFormación Cerro (Cretácico enla de tetrápodos terrestres El registro RESUMEN tardío, Coahuila,México. formes, Globidonta, Cretácico Palabras clave: Crocodyli del Pueblo durante elCretácico tardío. de tipotropical para la Formación Cerro ambientes fluviales enuna llanura deltaica de cariofitas, proporcionan de evidencia tortugas,formes, dinosaurios yoogonias La asociación de crocodyli osteodermos. con base en undiente aislado, vértebras y del Pueblo,La Parrita, Formación Cerro de la localidad restos de crocodyliformes En este trabajo los primeros se reportan y indeterminados representado por goniofólididos, eusuquios está crocodyliformes. Este último grupo , Lizbeth Nava-Rodríguez Brachychampsa montana /2019 727 ,

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Crocodyliforms remains from La Parrita, Coahuila ABSTRACT INTRODUCTION / GEOLOGICAL Crocodyliforms remains from La Parrita, Coahuila SETTING AND PALEOENVIRONMENT 728 728 1. Introduction with similaritiesto turtle dinosaurs.record The includes a pleurodire turtles, squamates, crocodyliforms, pterosaurs, and Cerro del Pueblo Formation in Coahuila includes land vertebrates from the fossilrecordThe of ( Alligatoridae redundantis analyses withthat of posterior teeth. Itscontent in recent phylogenetic at least ancestrally, bulbous by the presence of an alligatoroidGlobidonta is clade characterized, assigned asEusuchia. the material be can only restCoahuila. The of the Cerroin delPueblodyliforms Formation, croco thisclade of thus isthe second record of and Globidonta, as classification the allows tooth the of and onebulbous morphology tooth. The amaxilla and lower jaw,of vertebrae, osteoderms, Saltillo (Figurefragments 1),andincludes west of from the La Parrita locality, located 54 kmto the materialThe described in the present work comes alligatoroid illon-Martinez (2010) referred to the globidontan that Agu isolated teeth, and isolated osteoderms table,1998), alongwithaskull two frontal bones, rial (Rodríguez-de la Rosa andCevallos-Ferriz, eusuchian andgoniopholididmate indeterminate in the Cerro del Pueblo Formationforms include crocodyliCarpenter, 2014a,2014b).Records of (Rivera-Sylva saurids, and andornithomimosaurs problematic coelurosaur the dromeosaurids topsian Velafrons coahuilensis dinosaurrecordThe the hadrosaurids comprises ornithocheiroid (Frey andStinnesbeck, 2014). Pterosaurs are represented by oneindeterminate Rincon Colorado (Aguillon-Martinez, 2010). sp., ataxon basedonvertebrae collected near squamate only 2014). The is man, and trionychidkinosternoid, remains (Brink trionychid coahuilaensis lys Compsemys / e.g. Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín , Brochu, 2011;Skutchas Coahuilaceratops magnacuerna and Adocus Brachychampsa montana , the kinosternid , the kinosternid Neurankylus , and indeterminate chelydrid,, andindeterminate and Tröodon Chedighaii Latirhinus uistlani and , the cheloniid Richardoestesia et al , the paracryptodires Saurornitholestes ., 2014;Salas-Gis . , ankylosaurids, Hoplochelys “Coniophis” , tyranno , the cera Mexiche , the , the ------/2019 paleoenvironment 2. Geologicalsettingand America (Gilmore, and Lindsay, 1911;Carpenter late Maastrichtian age throughout North western early Campanianthrough from several sites of 1972), and (Erickson, Alberta southern ian DinosaurPark Formation of Albertochampsa langstoni Formation, Alberta (Wu southern from the Early Maastrichtian Canyon Horseshoe Lauprasert, 2010), suchas earlier( analyses wereglobidontans excluded fromAlligatoridae in thermore, some Late Cretaceous North American Alligatoridae is node-based (Brochu, 1999). Fur Globidonta is explicitly astem-based group, and mondi conditions, which were influenced by sea-level eventsoscillations (Eberth andstorm by influenced were which conditions, ited under marginal marine andshallowmarine age. Cerro The del Pueblo Formationdepos was Eberth 71 and72.5millionyearsold (Obradovich, 1993; CerroThe del Pueblo Formationdated is between fromformation the Difunta Group (Figure 2A). Pueblo Formation, whichthe oldest is geologic the Cerro del La Parrita is part of locality of The Saltillo, Coahuila,Mexico. Coahuila), Museo del Desierto (MUDE), tion of ontológica de Coahuila (Paleontological Collec Institutional abbreviations: CPC, Colección Pale America. al. Montana (Gilmore, 1911;Bryant,1989;Norell B. Mexico (Williamson, 1996);andthe type species, the Kirtland Formation, San Juan Basin,New of sealeyi Twoof species 2009; Aguillon-Martinez, 2010; Irmis 1996; Sullivan and Lucas, 2003; Monroy Mújica, 1980; Bryant, 1989; Norell

, 1994) and possibly elsewhere North in western montana from the CampanianDe-Na-Zin Member et al. et al. fo h elCekFraino from the Hell Creek Formation of , 2004),which indicates Campanian , 2015; Hastings Brachychampsa e.g. , Brochu,2004; Martinand from the middle Campan Brachychampsa et al. Stangerochampsa mccabei are recognized: , 1994;Williamson, et al. et al. , 2016), but et al. et al. reported , 1996); , 2004). , 2013). B. B. et - - - - -

Figure 1 Formation supported is (Vivas-González, 2013). Pueblo del Cerro which in base the is layer first responding to the Parras Shale, which this means fig.5),352, this layer. According to Eberth weathered dinosaurbones are associated with as the ammonite tonalities.shales with greenish Marine taxa such ochre-gray 0.20mof the base, itiscomprisedof sandstone, siltstone and shale (Figure 2B).At of layers alternating Parrita locality iscomposed of sequence that stratigraphic outcropThe inLa berant vegetation (McBride, 1974;Hill,1988). lagoons, coastal swamps, andwetlands with exu and bays lakes, residual or channels, oxbow flow deltaic withshallow systems facies suggest The Location of theLaParritalocality(red star), Cerro delPueblo Formation,Coahuilastate. Sphenodiscus Sphenodiscus ispresent infacies 1cor and some isolated, et al. (2004:350- Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín - - ossso 1 fochre-gray shale characterized 10m of consists of gastropods liesabove next The layer the grit. of by 1mabove, with a0.40mthickconcentration (Vivas-González, layer 2013).Agrit continues top, which indicates marine costal environments gastropods at the with highconcentrations of hard, massive, sandstone layer coarse-grained gastropods. Above the siltstone lies a0.90m banks andabundant non-ostrean bivalves and brackish conditions, indicated by oyster istic of aresandstone and siltstone deposits character and Frey (2014), the Cerro del Pueblo Formation brackish environment. According to Stinnesbeck diverse size, which indicate a oysters of tion of exfoliated siltstones with a high concentra of Above the base follows a0.30mlayer consisting /2019 729 729 - -

GEOLOGICAL SETTING AND Crocodyliforms remains from La Parrita, Coahuila PALEOENVIRONMENT GEOLOGICAL SETTING AND Crocodyliforms remains from La Parrita, Coahuila PALEOENVIRONMENT 730 730 Figure 2 shale). Alargediversityofvertebrateand invertebrate fossilsoccursinthesecond layer of shale(from thebottom). Formation (A),andlocalstratigraphyofLa Parritalocality(B),showing the sequence ofalternantstrata(sandstone,and siltstone, of dark-gray shale and again by 0.35 m of fine- of m 0.35 by again and shale dark-gray of fine-grained sandstones followed above by 0.50 m massive, gray, 0.40 mof nextThe layer consists of González, 2013;Stinnesbeck andFrey, 2014). this layer indicate anestuarine environment (Vivas- marine and limnic faunal assemblages that coexist in the and plains, coastal overflooded of suggestive is assemblage between assemblage facies 10–12. The et al. well ascoprolites (Vivas-González, 2013).Eberth tyrannosaurids anddromeosaurids) are present, as (hadrosaurids, dinosaurs bone andteeth of forms, trionychid turtles, some goniopholididscrocodyli present, andamongvertebrate fossils, reptiles like (Vivas-González, 2013). Wood fragments are also the most commoninvertebrates inthis shale layer oysters ( nodosa by fossil diversity: the marine gastropods / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Stratigraphicchartshowing theageand divisionsoftheDifunta Group withthestratigraphic positionof theCerro delPueblo (04 5–5)pae hskn ffossil (2004: 350–352) placed this kind of and Flamingostrea Lissapiopsis sp.). mollusksrepresent These sp. are common,aswell as Cerithium Cerithium - /2019 (Eberth associated with the Cerro del Pueblo Formation whichdeposition, isconsistentwithother localities during to marineregressionsand transgressions events related rita locality underwent aseriesof theLaPar sediment suggests alternating This these top layers (Vivas-González, 2013). No fossils were observed or reported in both of sandstones.3 monthe top are medium-grained by the section are formed shales, andthe last m of other fossils are reported inthis layer. next The 18 Inoceramus vamuxemi bivalves, referred to with highconcentrations of massive sandstones medium-grained 3 mlayer of fossils. Above this shale layer,devoid of there isa shale, which is tured sandstone layer lie 32mof fractured sandstones. Above the frac 0.20 mof shale, then following 0.60mare composed of sandstones (Vivas-González,grained 2013).The et al. , 2004). i h o fthe layer. No inthe top of - - 4.1 EUSUCHIANS INDETERMINATED 4. Systematicpaleontology 3. Methods CorelDraw X8. images were processed usingAdobe Illustrator and taken with aCanonX5digital camera, andthe caliper with 0.01mmaccuracy. were Photographs dles. Measurements were obtained with adigital materialThe wascleaned usingbrushesandnee individual, despite the specimens’ match insize. speculate whether ornot it comes from asingle the material comes from area, asmall we donot turtle and remains.fish fragmentary Although and 15m in anarea of the LaParrita locality. Allmaterial wascollected vertebrate microfossil surface collection method at All specimens were recovered usingthe standard f ih ail;CC18,mda rgeto right maxilla; CPC 1481,medial fragment of of fragment right maxilla; CPC 1480, medial Material. 9.2mm. with amaximum width of the crown isoval inoutline carinae. base of The the tooth. They lack atdeeper the distal side of that radiate from the apex to the base; they are striations irregular fine has It 3C). to 3A (Figures crown tooth inflated distomesialy and compressed Description. Formation, Campanian,Late Cretaceous. Coahuila.Cerro General Cepeda, del Pueblo of municipality the 101°20ʹ93.7ʺW,in 25°32ʹ68.4ʺN, Locality andhorizon. Material. Alligatoroidea Gray, 1844 Crocodyliformes Benton andClark, 1988 CPC 1483,isolated posterior tooth. Gen. andsp. indeterminated CC17,atrltrlprino CPC 1479, anterolateral portion of Globidonta Brochu, 1999 Crocodylia Owen, 1842 Eusuchia Huxley, 1875 CPC 1483 is small, labiolingually CPC 1483 is small,labiolingually 2 , inassociation with gastropods LaParrita islocated sensu Norell et al ., 1994 Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín - Locality andhorizon. with uncertainposition. vertebra;CPC 1489 dorsal CPC 1490, vertebra CPC 1487, lateral osteoderm; dorsal osteoderm; paravertebral CPC 1486,lateral dorsal osteoderm; CPC 1484, paravertebral CPC 1485, osteoderm; left dentary; left maxilla; CPC 1482, fragment of is ornamented with pits and grooves.is ornamented palatinal The surface maxilla (Figures external 4Aand4B). The the right responds to an anterolateral portion of Description. Cretaceous. Cerro del Pueblo Formation, Campanian,Late the Coahuila. General Cepeda, in municipality of 101°20ʹ93.7ʺW, 25°32ʹ68.4ʺN, located outline and are nearly confluent. The medial The confluent. nearly are and outline complete one. Allthe alveoli are semicircular in the tree preserved alveoli the middle isthe only surface.dorsal Three alveoli are preserved. Of oval vascular foramina are present on the latero withsmallpits andgrooves.ornamented is Two a semitriangularit hasshape. surface external The 10.3 mm inlength (Figure 4D). In transverse view the left dentary, CPC 1482 isa small portion of cating the sutural surface for the nasal (Figure 4C). on the medial margin,transverse has grooves indi surface. ventral The surface isslightlyrugose and, pitsandgroovesdeep 1480, ithas onitsexternal the left maxilla, and likeCPC 1479 andCPC the posterior part of CPC a fragment 1481 is of some vascular channels. perforatedcompact andis boneis imen. The by the spec transverse break at the anterior end of visible themaxillais at a structure of internal rugose, which the periosteum is lost.The suggests and rounded edges;the ventral surface isslightly pits, surface bears grooves, external maxilla. The the right tocorresponds amiddle portion of CPC 1480 isaquadrangular fragment that present awell-developed depression. preserved, but is smaller. Between both alveoli is eter circular other alveolus tooth. The is badly diam a6.4mm a root them stillcontains of of vascular foramina. Two alveoli are present. One concave, andslightly smooth surface is withtwo CPC 1479 is a fragment that cor The La Parrita The site is /2019 731 731 - - - - -

METHODS / SYSTEMATIC Crocodyliforms remains from La Parrita, Coahuila PALEONTOLOGY SYSTEMATIC PALEONTOLOGY / Crocodyliforms remains from La Parrita, Coahuila DISCUSSION 732 732 apical (B)and basal(C)view. Figure 3 ing the neural arch (Figures ante 5Aand5B).The CPC 1489isaprocoelous vertebral centrum lack the mandible. anterior portion of thatthe contact with the splenial suggest it isthe semitriangular shape and the evidencenial. The of striations indicating the contact surface for the sple the dentary is flat andsurface bearsof longitudinal marks andfour foramina (Figures 5Gand5H). with pits and grooves. The ventral surface has fiber angular outline. surface isornamented dorsal The CPC 1485isaparavertebral with arect osteoderm and hasfiber marks (Figures 5Eand5F). ventralnally runningkeel. The surface isplanar grooves. a small anteromedial It bears longitudi with pits and with asurface ornamented derm CPC 1484isaquadrangularparavertebral osteo suture surface isexposed andrugose. the neurocentral central part of 5C and5D).The served. centrum iswider than high(Figures The vertebra neural arch isuncertain. The isnot pre the position of posterior width = 9.97mm).The (length = 12.11mm,anterior width = 13.31mm, CPC 1490isasmallprocoelous vertebral centrum mal, probably 1.5mtotal length. posterior width = 13.21mm)indicates asmallani (length = 19.84mm,anterior width = 12.30mm, CPC 1489 the centrum isnot visible. size The of rior surface isdamaged, andthe anterior socket of / Globidontan alligatoroid fromLaParrita site,Cerro delPueblo Formation.CPC1483, isolatedposteriortoothinlateral (A), Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín A ------/2019 B 5.1. TAXONOMY DETERMINATION 5. Discussion The crocodilianThe specimenCPC 1483 from La the clade Globidonta (Brochu,these form 1999). gatoroidea includes taxa with bulbous teeth, and Among North Americancrocodilians, onlyAlli butosteoderms, CPC1486hasseveral foramina. and grooves. face is planar in both internal The a rounded shape. They are sculpted with large pits with uresand 5L)are 5K lateral osteoderms dorsal CPC 1486 (Figure 5I and5J)CPC 1487 (Fig atr Nrh mrc, a b differentiated from those of be can America, North eastern bidontan the glo CPC 1483.Onlythe posterior teeth of to similar sections, cross oval with inflated and them, the teeth are striated nona, 2014).Inallof Albertochampsa globidontans basal North American Late Cretaceous teeth of the isolatedupon impossible, toothis becausethe to genus or species levelrate determination based gatorid based upon inflated tooth shape. Anaccu Parritabe referred locality can alli toglobidontan Bottosaurus , resemble each other (Carbot-Cha Brachychampsa Stangerochampsa , presently known onlyfrom , Stangerochampsa , Albertochampsa C and and ------1482, fragment ofdentary (D). Abbreviations:da,dentary alveolus; fo, left foramen;mxa,maxillary alveolus.. anterolateral portion ofrightmaxillaindorsal(A) and ventral (B)view;CPC1481,fragment of left maxillaindorsal (C)view, and CPC Figure 4 C D art,i h eodrcr faglobidontan Parrita, isthe second record of quence, the crocodyliform specimen from La diagnosticcharacters. Inconse to the lackof sp., oreven better, onlyasGlobidonta, due with caution, and be referred as although the identification shouldbe treated champsa montana Pueblo Formation wasreferred to as Previously, crocodilian material from Cerro del 1998). wrinkled appearance (Erickson, compressed andhave strong striations that give a Brachychampsa Eusuchiaindeterminated fromtheLaParritasite,Cerro delPueblo Formation.Skulland mandibular elements:CPC1479, , because the teeth are laterally by Aguillon-Martinez (2010), Brachychampsa Brachy Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín - -

and Clark,1988; Brochu, 1997;Molnar eusuchianCrocodylia including forms, (Benton the centrum, which characterizes the advanced of referred to Eusuchia based on the strong procoely CPC 1490 described in thiswork canonlybe On the other hand, the vertebrae CPC 1489 and Mexico. crocodilian diversity inthe Late Cretaceous of this taxon inNorthAmerica and increasing the mation, record extending the geographical of alligatoroid from the Cerro del Pueblo For 2015). The neurocentral2015). The suture surfaceexposed in B A /2019 et al. 733 733 - ,

Crocodyliforms remains from La Parrita, Coahuila DISCUSSION DISCUSSION / CONCLUSIONS / Crocodyliforms remains from La Parrita, Coahuila ACKNOWLEDGEMENTS 734 734 5.2. PALEOENVIRONMENTAL SIGNIFICANCE osteoderms toEusuchia level.osteoderms akeel in CPC 1484, allow referred the presence of aswell as the the osteoderms, the general shape of large (Brochu,individuals 1996).Inthe same way, rocentral sutures incervical vertebrae happens in neu specimen, because incrocodilians closure of anon-adult the vertebra CPC 1490 isindicative of fluctuating paleoenvironments of intermittent shal intermittent of paleoenvironments fluctuating cyclically by influenced was setting depositional The changing delta plain (Vogtpermanently in multiple stacked channelized sandbodies ona the Cerro del Pueblo Formation deposition suggest well asplant material structures andsedimentary in vertebrateThe andinvertebrate assemblages as crocodilian material. a remarkable amount of or frequentlywere flooded, and contain permanently likely assembled indepressions orchannels that concentrationsmm upto about 150mm.These were the specimens ranges between 0.5 size100 m².The of afew upto about surface clusters, covering anarea of lated inthe sandy layers, orintaxonomically mixed are fragments or isolated bones that may occuriso vertebrate remains at La Parrita vastThe majority of subaerial conditions. layers at LasÁguilas, terrestrial oreven alsosuggest sandstone cracks, repeatedly found onthe surfaces of climate. Mudcracks, here asdesiccation interpreted on adelta plain(Vogt environments fluviolacustrine for evidence provides dontan alligatoroid, aswell at asdinosaurs LaParrita, charophyte oogoniawith globi associationThe of cannot livetors inestuaries orcoastal areas. (Taplin intolerant to salt water because they lack salt glands the Alligatoridea are indicators. Recent of members it is possible to use fossil crocodilians as paleoclimatic tem indicators (Mazzotti Recent studies show that crocodilians are good ecosys global on extant crocodilians (Marwick,1998). distribution of influence principal the is Temperature / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín et al. , 1982).Consequently, the extant alliga et al. et al. , 2009).Inconsequence, , 2016) and a tropical , 2016)andatropical et al. , 2016). , 2016). ------/2019 6. Conclusions Acknowledgements in North America. The vertebrae and osteoderms vertebraein North America. The andosteoderms Globidonta recordextended the geographical of an isolated tooth. This of in the morphology rita locality, Cerro del Pueblo Formation, based alligatorid specimen from globidont La Par of evidence first the of presence the recognize We brackish environments (Vogt salt andfresh water, generating achanging zone of of the measured section indicates frequent mixing oysters throughout conditions. abundance The of low marine, brackishto fresh water, oreven subaerial Wikipaleo group for sharing bibliography, and supporting thisinvestigation.the GCCthanks Arturo González forDesierto and itschairman to thisproject. We are del to the Museo grateful Zaira Casas are acknowledged for their support Lucía Alfaro Ortíz,IsaíOyervidez Salazar, and al. tles, ( crocodyliforms anddinosaurs trionychid tur vascular plants, shell fragments of conditions isindicated by characean oogonids, other hand,continental to brackishto fresh water indicate shallow marine environments. Onthe and marine vertebrates (sharks andmosasauroids) banks, abundant bivalves, ammonites, gastropods, oyster brackish conditions. presence The of under deposition stones andsiltstones suggests sand at LaParrita. oscillating The sequence of environments depositional exists A mixture of scarce. crocodyliforms inMexico is Mesozoic record of but aredetermination, important because the characters to genus orspecies levelmorphological are be referred to Eusuchia, due to the lackof , 2008). et al. , 2016). e.g. , Meyer et et - - - Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín /2019

Figure 5 Eusuchia indeterminated from the La Parrita site, Cerro del Pueblo Formation. Postcraneal elements: CPC 1489 dorsal vertebra in lateral (A) and dorsal (B) view; CPC 1490, vertebra with uncertain position in ventral (C) and dorsal (D) view; CPC 1484, paravertebral osteoderm in dorsal (E) and ventral (F) view; CPC 1485 (I), 735 735 paravertebral osteoderms in dorsal (G) and ventral (H) view; CPC 1486, lateral dorsal osteoderm in dorsal (I) and ventral (J) view; CPC 1487, lateral dorsal osteoderm in dorsal (K) and ventral (L) view. Abbreviations: co, condyle; vc, vertebral centrum. Scale bar = 10 mm.

CONCLUSIONS / Crocodyliforms remains from La Parrita, Coahuila ACKNOWLEDGEMENTS Crocodyliforms remains from La Parrita, Coahuila ACKNOWLEDGEMENTS / REFERENCES 736 736 References editorial support. review andsuggestions, andFranciscoJ. 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