This is an open access article under the CCBY-NC-SAThis license Universidad Nacional Autónoma de México. Peer Review under the responsability of Manuscript 30/05/2018 accepted: Corrected manuscript received: 28/05/2018 Manuscript received: 08/01/2018 BSGM2020v72n1a160819 72 (1), A160819.http://dx.doi.org/10.18268/ Boletín de la Sociedad Geológica Mexicana, Cretaceous marine fossilPuebla,site in Mexico: the San José de Gracia quarry, anewLate Villareal, V., 2020, Vertebrate fossilsfrom Cruz, J. A.Castañeda-Posadas, C.,Zavaleta- Alvarado-Ortega, J., Cantalice, K.M.,Díaz- To citethisarticle: [email protected] * Corresponding author:(J. Alvarado-Ortega) 72570, Mexico. Universitaria, Jardines de San Manuel, Puebla, Ciudad Bio-1, Edificio Puebla. de Autónoma Ciencias Biológicas, Benemérita Universidad 3 Alcaldía Coyoacán, CDMX,04510,Mexico. la Investigación S/N, Ciudad Universitaria, Nacional Autónoma de México, Circuito de 2 Alcaldía Coyoacán, CDMX,04510,Mexico. Investigación S/N, CiudadUniversitaria, Autónoma de México, Circuito de la 1 Carlos Jesús Cretácico SuperiorenPuebla,México Fósiles devertebradoslacanteraSanJoséGracia,unnuevositiofósilmarinodel fossil siteinPuebla,Mexico Vertebrate fossilsfromtheSanJosédeGraciaquarry,anewLateCretaceousmarine (https://creativecommons.org/licenses/by-nc-sa/4.0/) Posgrado enCiencias Biológicas, Universidad Instituto de Geología, Universidad Nacional Laboratorio dePaleontología, Facultad de Alvarado-Ortega Castañeda-Posadas 1,* , Kleyton M. 3 , Valentina cala Formation, vertebrates. quarry,Turonian, Puebla, Mex Keywords: San José deGracia Turonian age.los-Guerrero Basin)of to the Mexcala Formation (More canbe possibly referredGracia quarry theSanJosé de fossil-bearing strata of its fossilcontent and itslithology, the mosasaur isalso reported. Based on country. In addition, a yaguarasaurine record of known from Mexico; aswell as the first Enchodus described, includingrepresentatives of fossil fishremains from thislocality are plant remains. In this work, the first ostreids,indeterminate reptiles and few ammonites, belemnites, inoceramids, remains; fish italsocontains of mostly Josécomposed is deGraciaQuarry association recovered from the San previouslyreported. Althoughthe fossil theregional geological units those of fossilwith content donotcorrespond known; and the characteristic lithology sequence are this not lower limitsof thickness. Althoughthe upper and carbonated clays, withcentimetric poorly cream-brown strata of of are extracted from amarinesequence for commercial where purposes, slabs than the two hectares that is exploited occupiesanarea no larger quarry The thissite. mal scientificcontribution of decade ago. for paper the first is This paleontological site discovered about a Puebla, Mexico,southern a new is Molcaxac, within the Municipality of SanJoséThe deGraciaQuarry, located ABSTRACT Cantalice Zavaleta-Villareal Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín andclupeidbothpreviously Dercetis 1 , Jesús Alberto and Xenyllion 2 from this Díaz-Cruz - - - Este trabajo representa la primera contribución tológico descubierto hace cerca de una década. Estado de Puebla, México, es un sitio paleon dentro del Municipio de Molcaxac, alsur del La Cantera SanJosé de Gracia, ubicada RESUMEN vertebrados. Mexcala,Formación Puebla, José de Gracia, edad Turoniana, Palabras clave: Cantera San edad Turoniana. yson de (Cuenca Morelos-Guerrero) Mexcala son referibles posiblemente a la Formación fosilíferos de la Cantera San José de Gracia fósil ysu litología, se sugiere que los estratos saurinoMéxico. en basesu contenido Con en elhallazgo deun mosasaurio yaguara reporta de mexicano México; así comoelprimer registro clupeido, ya previamente conocidos en grupos destacan peces fósiles de esta localidad, entre losque se este trabajo sedescriben losprimeros restos de ymuyescasos restosreptiles de plantas. En nites, inocerámidos, ostreidos indeterminados, peces; ésta también contieneamonites, belem se compone ensu mayor parte de restos de recuperada de la Cantera San José deGracia Aunque la asociaciónpara fósil esta región. las unidades geológicas previamente reportadas conaquellos de contenido fósil no corresponden son conocidos; sus características litológicas y superior einferior, de esta unidad rocosa no varios centímetros. Apesar de que loslímites, cuyo alcanza grosor de color marrón-crema, en estratos carbonatadosparalelos, depositadas arcillosascompuesta de margas pobremente lajas de una secuencia de origen marino, tadasfines comerciales, con donde seextraen superficie nomayor a las dos hectáreas explo Esta cantera ocupa una científica formal. 2 , Dercetis Enchodus y /72(1)2020 Xenyllion , Laminospondylus . Aquí, también se yun 1 - - - -

Late Cretaceous vertebrates from the San José de Gracia quarry ABSTRACT Late Cretaceous vertebrates from the San José de Gracia quarry INTRODUCTION 1. Introduction of ServicioGeológico Mexicano, 2006). Figure 1 2 2 ok ihntelmt ftheir communal property rockswithin the limits of sedimentary Mexico, toexploit of began a deposit Puebla, central Molcaxac,State of Municipality of the San José de Gracia town, In 2006,people of dures usedduringtheir extraction have allowed us used to cover floors and walls.The artisanal proce its good appearance and resistance, these rocksare Given extracted. are veins reddish with flagstones “San Josécream-brown deGraciaquarry”, marly (Figuresite,this 1).In namedas herein formally / Mapof SanJosédeGraciaquarry showing thegeological formations presentinthisregion of Puebla, Mexico(based on map Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín http://dx.doi.org/10.18268/BSGM2020v72n1a160819 - /72(1)2020 quarry have scorned the scientific and aesthetic and scientific the scorned have quarry new this in extracted flagstones of traders and ers (Applegate locatedfromsouth about 10km Molcaxac town emblematic paleontological site Lagerstätte with thosefrom the Tlayúa quarry, aMexican theyare compared scarcepoorly and preserved if 2015). 2011; Romero-García, 2013;Zavaleta-Villareal, Alvarado-Ortega we partially report in thiscontribution (see also to discover a peculiar fossiliferous association that Fossilsfrom the San José are de Gracia quarry et al ., 2006).Forreason, this thework et al ., 2009,Pulido-González, Artícle A160819 # - Artícle A160819 Gracia quarryshowingmarl strata. the centimetric Figure 2 tiles), andscarce plantremains. rep and (fishes vertebrates crabs), and bivalves, invertebrates (ammonites, belemnites, gastropods, UNAM and BUAP,entific collections of including from have thisquarry in the sci been deposited and studying preserving them formally. of Currently, about 200 macrofossils goal final the with in order to recover fossilsfrom thisnewlocality, the San José de Gracia have worked together of Autónoma dePuebla (BUAP),well as workers as the UNAMand the Benemérita Universidad of first the thiswork. to Thus, in2008,researchers author of Gracia de José San the from fossils nal Autónoma de México (UNAM), gave some the Instituto de Geología, Universidad Nacio the Tlayúa Paleontological Project, supported by Molcaxac),andcollaborators of boring town of Tepexi Rodríguezinhabitants of town (neigh sources, different from flagstones of traders thy, theserocks. of Gracia specimensduringthe extraction and sale San José de a great quantity of destruction of their fossils;this hascausedthe lossor value of TheSanJoséde Graciaquarry;a)general viewof oneoftheexploited inthisquarry; b)Detailsof sections theSanJoséde In 2007,Messrs. FélixandSebastián Arangu http://dx.doi.org/10.18268/BSGM2020v72n1a160819 Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín - - - - showing gradation, normal and deposited in finely clays poorly carbonated, includes marl layers of the San José (Figure 2); which de Gracia quarry decimetric marly strata has been exploited in of to West and40mfrom NorthtoSouth(Figure 1). extends from about150m de Graciaquarry East ° 52’13.94”W. exploited The surfaceinSan José between the coordinates 18° 39’27.25”Nand97 1930m.a.s.l. and isat the altitude of the quarry the Cacique Stream. Currently,of the center of side tral Mexico. site is inthe southeastern This Molcaxac,Puebla, cen within the Municipality of homonymoustown, westkm from the center of while in the west this geomorphological featurewhile in the west is thisgeomorphological the strike isvariablequarry between N50-88°W, the an anticline. end of At the eastern presence of havethis quarry a strikeanddipthat the suggest with other lithological units; the strata exposed in possible faults andcontacts ognize the presence of José makingit impossible to rec de Gracia Quarry soil coversthe geological features around the San laminated parallel planes (Figure 2). Although the Up to now, a sequence of about 4 m thick about4m Up tonow, asequenceof San JoséThe islocated 2.5 de Gracia quarry /72(1)2020 3 3 - -

Late Cretaceous vertebrates from the San José de Gracia quarry INTRODUCTION MATERIALS AND METHODS / Late Cretaceous vertebrates from the San José de Gracia quarry RESULTS 2. Materialsandmethods 4 4 ceous marinevertebrates andinvertebrates. SanJoséLateincludes Creta deGraciaquarry of thiswork,the hitherto knownfossil assemblage of respectively (Figure it 2). Asisnoted in the results layers isvariable between 23-35° S and 23-40° S close toN90°W. Atthesametime,these thedipin cited. 3 to8),arethe publications basedon respectively nomenclature and abbreviations usedhere (Figure taxa described here, aswell as the anatomical tions: andabbrevianomenclature Anatomical IGMasIGM-loc3876. cal sitesof paleontologi isrecordedquarry in the catalog of BUAP.Paleontología of San José The de Gracia theColección de PALZthat the acronym is of the Instituto de Geología, UNAM, as well as BUA Colección Nacionalde Paleontología, housed into under the acronymIGM that to the corresponds San José de Gracia are and catalogued deposited abbreviations:Institutional with afinebrush. diluted in methacrylate acetateplexigym applied late; they were hardened with weak solutionof necessary, the fossilswere with cyanoacry glued scavengersunder stereoscopic microscope. When ically prepared withneedles andodontological Preparationmethods: ical site. thispaleontolog describe the geological details of to report theSanJoséis deGraciafossilsand thiscontribution greater detail. Thus, the aim of fossil assemblage deserves to be recognized in Puebla. Therefore, we are convinced that this of distinctive lithology, never observedregionthis in represent a peculiar assemblage preserved into a José deGraciaquarry, we recognized that they / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Since our glancefirst at the fossils from the San h txnmcl dniiain ffossil of identification taxonomical The Fossilswere mechan Fossilsfrom the http://dx.doi.org/10.18268/BSGM2020v72n1a160819 ------/72(1)2020 3.1.1. SYSTEMATIC PALEONTOLOGY DE GRACIA 3.1. FOSSILVERTEBRATE ASSEMBLAGE OFSANJOSÉ 3. Results and counterpart, fairly complete specimen with and counterpart, material: Referred trunk is entirelytrunk is not possible scaled,it is to see more the Since ribs. 16 preural has fish Probably, this seem to be rectangular, slightly higher than long. somewhat distorted in IGM11190;however, they two urals. abdominal and caudal centra are The column, including18abdominals, 14caudals, and pierced bone. ceratohyalan elongated, is rectangular, andno are partially exposed showing that the anterior preserved in thisspecimen; hence, the hyoid bones of bothjawsare not the latter bone. bonesof The above just located is first the preserved, well moid region, boththemesethmid and vomer are separated bythe supraoccipital bone. Inthe eth temporal commissure and are laterally displaced, smooth. Inaddition, the parietals have the supra are bones possible toobserve thatroof theskull thetrunkwasabout13mm. maximum heightof head and ashallow caudal peduncle, inwhich the triangular a with chubbyfish, a be should this life In preserved. is fins its of none with (SL), length standard trunk stronglycurved, reachesof 40 mm preserved showsthe in part and counterpart Description: out thejaws andfins(Figure 3). Suborder ClupeoideisensuGrande, 1985 Order Clupeiformes sensuGrande,Order Clupeiformes 1985 There are thirty-four centra in the vertebral poorlyis Although the skull preserved, itis Genus andspeciesindeterminate Family ClupeidaeCuvier, 1817 This almost complete specimen This IGM11190AandB, part Artícle A160819 # - - - Artícle A160819 fused; u1-2,ural centra1and 2. anterior ceratohyal(no pierced); apt,analpterygiophore; dpt,analpterygiophore; pa,parietal;ur1+pu1, ural 1and pleural centrum 1 the SanJoséde Graciaquarry; a)general viewofthespecimen;b)closeup ofthecaudal fin. Abbreviations:abs, ac, abdominal scutes; Figure 3 projects back andforth from smallspines, aswell triangular; theyhave a thickened ventral edge that anal pterygiophores. abdominal scutesare The first the and isthmus the between extends scutes at least 14 abdominal in this fish. A ventral row of concentric circulli. scutesare No dorsal preserved scales, taller than long, with numerous ornamented fins are notpreserved. pelvic and caudal the SL (30 and 35 mm). The of 82.5% and 75 between located is fin anal the that There are 10visible analpterygiophores showing SL. the of middle the to near rises fin dorsal the visible pterygiophore single dorsal The shows that oneisovoiddorsal andthe ventral one isrod-like. Behind thisbone, there are two postclithra, the bone that is slightly widened at its ventral end. the pectoral girdle a Cshaped the cleithrum is bones. the vertebral column andassociated details of SpecimenIGM11190,representative ofanindeterminated genusand speciesbelonging toofthefamilyClupeidae from The trunk entirelyThe covered for ovoid cicloid however, in preserved; not is fin pectoral The http://dx.doi.org/10.18268/BSGM2020v72n1a160819 Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín relation with the mostcaudal preural centrum, ral centrum, the ural centra are relatively small in the first uroneuralthe withfusion of the first preu tionally, as inall clupeoideis, this specimenshows (see Grande, occur inall clupeiforms 1985);addi foramen, the beryciform as piercedof duethe loss tal bone and its anterior ceratohyal is not medially parietals are entirely separated by the supraoccipi its abdominal scute series asall clupeomorphs; clade, described by Grande, hasthe 1985). This this because it shows the diagnostic features of Clupeiformes) the Clupeoidei (Clupeomorpha, Remarks: diastema present between thehypurals 2and3. uroneural bone are fused andthere is a caudal first the and 1 preural the skeleton, caudal the In the ribs. leavingspace toobserve the extension of barely scutes These overlapcolumn. eachother, the abdominal area below the vertebral third of ward, directed upward, andcovering lest than one as alaminartriangular wing slightlycurved for IM110i eonzda ato IGM11190isrecognized as part of /72(1)2020 5 5 - - - -

Late Cretaceous vertebrates from the San José de Gracia quarry RESULTS Late Cretaceous vertebrates from the San José de Gracia quarry RESULTS 6 6 (15/32) is 0.46andmatches withthat known for IGM11190 ribs topleural vertebrae ratio of pleural numbersuperfamily Clupeoidea. The of the it is possible to consider IGM11190 as part of Suborder. to thementioned fish belongs this that confirms fin is not preserved not is fin mm 10 is head The (Figure 4). exposes its left lateral surface, inwhich the caudal that length mm 140 least at of fish longirostrine Description: its leftside(Figure 4). exposing fin caudal without fish complete almost material: Referred species wasnever illustrated andnow islost]. a species of Westphalia, Germany, Northweastern, as the type Baumberg, in of and Campaniandeposits Dercetis scutatus [Goody(1969) recognized England of deposits p. 258),from the Turonian andSenonian marine by1843, 1843 (inAgassiz, Munster and Agassiz, but its descriptionwaspublished ten years later genus waspublished by(1833, p. Agassiz XLIII) this of reference first The 258). (p. 1833 Agassiz, this genus is the type species of Type species: thisclupeidfish. of nomical determination materials are required to reach an accurate taxo 11190 deserves an accurate study; additional fossil Malabarba and Di Dario, 2017). Although, IGM Brazil (Figuereido, 2009;Lavoué the Morro de Chaves Formation, northeastern of (Earlyan-Aptian Cretaceous) in marine deposits with afossilrecord datingto the Barremi back thisclade(0.47to0.71). of members / nomen nudum Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín In addition, based on data from Grande (1985), The familyClupeidea is anancient living group The Family Dercetidae Woodward, 1901 Order Rosen, Aulopiformes 1973 Genus Dercetis bcuetesnl pcmno this specimen of because the single IGM11191isanelongated and Agassiz, 1834, from Agassiz, the Turonian Dercetis ; however, thiscanbe considered According Taverne (2005a) Dercetis IGM11191,anelongated Agassiz, 1833 Agassiz, sp. http://dx.doi.org/10.18268/BSGM2020v72n1a160819 Dercetis elongatus et al ., 2013; - - /72(1)2020

small tubercles. some thisbonehas above the orbit the surface of right frontalalmost entirely boneis lost;however, wide and high ascendingprocess.discernible The similarsize. premaxilla The has a teeth, conic of slightly curved, shows dentary twoThe rows of jawsare projected forward to the same level. highallalong.slightly lower anduniformly length) respectively.skull elongated trunk is The the of 60% and 40 (or mm 18 and 12 are lengths postorbital and preorbital (= snout) regions, which orbital- 30 mm;thisisdividedintothe length of skulllength is about high and38mmlength. The smooth andtriradiate scutes along the body; one sible to see that IGM 11191hasthree rowsof eight rays. thefishlength consistingof mm of between the 110 mm(= preanal length) and 118 short; its length isjutsabout 8 mmandit is located extremely is fin anal rays.The 40 about of sisting con length fish the of mm 106 and length) dorsal locatedlength is and between(= pre the 54mm mm 52 is fin dorsal The rays. 5 least at of sisting con fin dorsal the of part middle the to opposed girdle, is pectoral the behind mm 25 located is fin 11rays. pre-pelvic The distance is 55 mm; this of the trunk andconsists the abdominal border of pectoral girdle, between the vertebral column and are present allalongthetrunk. backward. Numerous and epipleurals epineurals areneural and hemalspines shortandcurved abdominal andattach withthinpreural ribs. The each abdominalcentra. At least 40 vertebrae are of processlaterally risesfrom the anterior half are slightly higher than long. transverse Asingle centra, preural first the probably specimen, this be less long. last two The vertebrae preserved in than highintheabdominalregion; theytend to Although, thevertebrae are about1.5timelonger while theirstrongly middlepart is constricted. tebral articulationaresurfaces wideandmassive thread reels, in which the interver the shape of This fossilThis preserves seventy vertebrae showing Although badpreserved, inthe head both Although theseare poorly preserved,pos itis ofthe the middle behind is fin pectoral The Artícle A160819 # - - - - - Artícle A160819 length +postorbitallength; trp,tranverse pcf,pectoralfin;prol,preorbitallength; process. sc,scutes; pvf,pelvic close up oftheabdominalregion; d)closeup of theposteriorpart of thetrunk; Abbreviations: af, analfin; df, dorsal fin;ol+psol,orbital Figure 4 trunk justabove thevertebral column. the tral body edge; and the last covers the flank of bodyalong the dorsal edge; another along the ven he fthesecharacteristics. three of Dercetis than mandible. IGM 11191 isincluded within this more (with fin 20 rays)and more than onetooth row inthe dorsal long a vertebrae, transverse processes onabdominal pair of a single ascendingprocessin the premaxilla, presence of Taverne, 2005b;these three genera share the of these groups includes of two monophyletic groups withinDercetidae. One Vernygora this family. these characterswithin supporting itsinclusion all shows 11191 IGM scutes. flank shaped arrow shallow head, long snout, and are the presence of reduced neural spines. Otherfeatures homoplasic the presence synapomorphy: non-ambiguous of point out that this familyissupported on asingle genera included in the clade Dercetidae; they 14 reviewed the phylogenetic relationship of Remarks: SpecimenIGM 11191, Cyranichthys - Ophiodercetis Recently Vernygora et al Taverne, 1987,and . (2017, suppl. 1) also revealed group because it shows the first Dercetis Dercetis sp.from theSanJosédeGraciaquarry;a)general viewof thespecimen;b)Detailsofhead;c) http://dx.doi.org/10.18268/BSGM2020v72n1a160819 assistergroup et al Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Ophiodercetis . (2017) -

Ruge, andPáramo Fonseca, 2017.In Candelarhynchus neurocranium), includingthe recently published the two timesthantheorbito-postorbital part of dercetidsconsiderablyis thesnout longer (at least the skull length. Contrary,exceed 60% of in other Dercetis the preorbital region is elongated;or snout in least four characteristics.all dercetids Althoughin that ec h ii f h rtcosi tefnigo of finding the Cretaceousif the of reachlimit the thisgenus could temporal distributionThe of Sahel Alma, Lebanon. Santonian sediments of and England of D. elongatus are included withinthe genus, the type species and Gallo, 2011);today, two nominalspecies species(see Goody,large number 1969;Silva of a representative of as identification its and features both show 11191 or isreduced to a tiny nodule. Here IGM again, developed whereas inother dercetids thisdisappear the premaxilla is well the ascending process of Taverne (2006, fig. 13, p. 41–42) pointed out pointed p. 41–42) 13, fig. (2006, Taverne Although oncethe genus Dercetis the snout is relativelyis thesnout shortanddoes not differs from other dercetid genera in at from Turonian-Senonian deposits Vernygora, Murray, Luque, Parra D. triqueter Dercetis /72(1)2020 isundeniable. Pictet, 1850, from Dercetis includeda Dercetis 7 7 ,

Late Cretaceous vertebrates from the San José de Gracia quarry RESULTS Late Cretaceous vertebrates from the San José de Gracia quarry RESULTS 8 8 postorbital regions of the skull are 60% and 40% and 60% are skull the of regions postorbital the preorbital- the cranium, the lengths of of the preorbital region the skull, of proportions of reported by (2006)isconfirmed. Kadummi Chalk Marl Formation, Lebanon, near Harrana, Dercetis able wide rounded cavity withrough surface ament are preserved. palatine The has aremark grove northegroove for thepalate-maxillarylig this palatine; unfortunately, this neither the end of edge of the dorsal backward from the middle of for the palato-premaxillary projectedis ligament along andnear its ventral edge. Ashallow groove rounded andpracticallybeingwider smooth, thismassive bone is medial surface of 5). The slightly curved is straight and the dorsal (Figure long and 7.43mmhigh; itsventral edge is almost asquad-like structure.form palatine The 42.97 is tooth surface. stout bone and its terminal This complete palatine bonethatexposed is its labial on Description: 5). palatine bone bearing an incomplete tooth (Figure material: Referred (Goody, 1969). dle Chalk inLewes, England Sussex, southeastern the Mid 1822), Turonian and Senonian rocksof Type species: in America. genus this of finding first the represents specimen genus the of species referredis an indeterminate toas reachingAt thistime, aconclusion. IGM11191 fromspecimens San José de Gracia before D. triqueter the present62). In context,11191resembles IGM p. 1969, Goody, 184–185; p. 1901, (Woodward, in / Family Enchodontidae sensuFielitz,2004 Genus Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín D. triqueter ohnmnlseiso speciesof Both nominal i asrcta eiet fMuwaqqar inMaastrichtian sediments of Dercetis Enchodus ; however, it is desirable to have more while in ; however, it is noticeable that this IGM11192includesanalmost Enchodus

Agassiz, 1833(p. Agassiz, XLIV) Enchodus lewesiensis IGM11192,acomplete D. elongatus sp. Dercetis http://dx.doi.org/10.18268/BSGM2020v72n1a160819 hy r 50% are they differ in the in differ (Mantell, - - - /72(1)2020 palato-premaxillary ligament. Abbreviations:aca,anteriorfor carina;section. the ppg,groove Gracia quarry,showing acloseup ofthepalatinetoothincross Figure 5 (Maldonado-Koerdell, 1956; Blanco-Piñón 1998; San Luis Potosi, Hidalgo, Guerrero, andChiapas throughout Coahuila, Tamaulipas, Nuevo Leon, numerousin Albian-Maastrichtian deposits In Mexico, fossilof planet (Fielitz and González-Rodríguez, 2010). the thatof inhabitedpracticallyalltheseas active Upper Albian-Eocene marine predators speciesof The Aulopiformes). the family Enchodontidae (Order member of 70 nominal species, being the most diverse fossil about of consists fish, saber-tooth as named Remarks: (Figure 5). laterally,an anterior andhas cutting edge or carina the palatineovoid, tooth is compressed surface of broken thistooth. The does notproject in front of thepalatine clearlyis thistoothalso terminal, of the palatine. position The with the ventral edge of about 100°, anobtuse angle,forward of forming the palatine tooth is slightlytilted length. This of 50% about or mm 20 reach could length total its thepalatine; however,fromthe ventral surface of thistooth is 11mmmeasured preserved part of basaland should be straight The and smooth. tooth is not preserved; this ankylosedstructure just behindthepalatine tooth. the bone opening on the lateroventral surface of SpecimenIGM 11192, Although, the tip of the single stout palatine the single Although, the tip of The genus The Enchodus Enchodus Enchodus have been collected sp.fromtheSanJoséde Enchodus , commonly Artícle A160819 were were # Artícle A160819 recent systematic studieson only speciesrestricted toMexico. the genus (Albian-Cenomanian) and the known of remarkable because it represents the oldest species as México, which were reported by Giersch (2014) Vallecillo quarry,of deposits Nuevo León,northern of Marchese, 2013); as well as possible representatives Mexico,southern (Carbot-Chanona and Than- the Ocozocuautla Formation, Chiapas, of deposits and petrosus Mexico; isolated palatine teeth referred to the Muhi quarry, Hidalgo, central of deposits Rodríguez, 2010, from Albian-Cenomanian species Mexico at thespecieslevel. in identified been have species five however,only al Alvarado-Ortega González-Barba and Espinosa-Chávez, 2005; as including species, five shumardi in only carinated) ovoid, anterior carina (uni- andshowsa single thistooth is symmetrical, tooth, the crosssection of is multidentated. is a peculiar enchodontid fish, in which the palatine andCarbot-Chanona, 2016. Ortega Unicachichthys multidentatus the family Enchodontidae menus of all members only for palatinesharedcondition a homoplasic toothis Díaz-Cruz Considering the results2017; amongothers). of al Fielitz and González-Rodríguez, 2010;Cavin the last phylogenetic approaches(Fielitz, 2004; them were included into andjust14of others) andAgnolin,2010,among Chalifa, 1996, Bogan species have been considered (Goody, 1976, ., 2008;Díaz-Cruz ., 2012; Díaz-Cruz E. E. venator E. gladiolus Despite its taxonomicrichness, the most in the comprise complete specimens of These Among enchodontids palatinewith asingle cf. E. ferox E. Cope, 1874, Leidy, 1856; E. zimapanensis Rharbichthys ferox venator et al Arambourg, 1954from the Turonian Leidy, 1855,from Maastrichthian , . Amongthese, E. zimapanensis . (2016); the presence of a single asingle . (2016); the presence of et al E. dirus et al et al ., 2006a,2006b;Giersch E. gladiolus Fielitz and González- Arambourgh, 1954and ., 2016; Holloway Díaz-Cruz, Alvarado- ., 2016; among others); ., 2016;amongothers); , Leidy, 1857;aswell , andthe unnamed http://dx.doi.org/10.18268/BSGM2020v72n1a160819 E. zimapanensis Enchodus (Cope, 1875), Unicachichthys Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín , only 18 , only et al isa E. E. et et .,

rica, within a range between Late Cenomanian them are present only inNorth Ame since all of species noted above congruent geographically is IGM11192 withthe three nominal of among others). 6; 5, 3, figs. 1996; Chalifa, 4E; fig. 2004, Fielitz, 36; fig. (Goody, 1969, base tooth the of front in because this bonehasaprojection is notterminal as well as in Enchodus where the palatine is well known, of this work; it is noteworthy that in other species of be analyzed ingreater detail and beyond the scope palatine tooth.feature Althoughthis deserves to the position of species isthe anterior terminal feature shared byIGM 11192andthese same this fossilfrom SanJosé deGraciaquarry. Another the taxonomicidentity of attempt to determine 5); itisnotpossible toadvance further the in symmetrical, ovoid,uni-carinated andis (Figure itspalatineing asmooth thatcross in section is Enchodus shumardi shares the characteristicsabove discussed with 2017). E. dirus al (Fielitz and González-Rodríguez, 2010;Cavin the palatine toothshowslongitudinal striations and the unnamed pce fspecies of five fig.2004, Moreover, 4E). group of the among noticeably longer than the palate bone (Fielitz, forward andsoextremely developed that it is palatinebecause its single tooth is strongly tilted differs clearly from IGM11192andthe nominal species this however, carinated; this toothalsoovoidis incrosssection anduni- compressed andbi-carinated. In tooth is rounded without carinae or laterally this palatine tooth is known, the crosssection of contrasts,In inother enchodontidswhichin the et al Fielitz and González-Rodríguez, 2010; Cavin Enchodus ., 2012)whereas in Nonetheless, the taxonomical approximation IGM11192 Although the palatine tooth of ., 2012;Holloway this tooth is entirely thistooth is smooth(Parris speciesfrom Gavdos(Parris Enchodus Rharbichthys ferox , E. gladiolus noted above, in Enchodus E. shumardi et al /72(1)2020 ., 2017; among others). ., 2017;amongothers). , and species from Gavdos , the palatine tooth , Rharbichthys ferox E. gladiolus E. dirus E. zimapanensis Enchodus et al , includ Enchodus ., 2007; , and et al spp. 9 9 et ., -

Late Cretaceous vertebrates from the San José de Gracia quarry RESULTS Late Cretaceous vertebrates from the San José de Gracia quarry RESULTS 10 10 Order Sphenocephaliformes RosenOrder Sphenocephaliformes andPatterson, Although based on the present context, IGM 0001 andAngolin,2010). South America (Bogan of probably alsoreaches Maastrichthian deposits (Carbot-Chanona and Than-Marchese, 2013)and México part of the southern trichthian of deposits distribution of and EarlyMaastrichthian (Parris occipital crest,roofed andmainly bythe frontal triangular,is cracked, the skull withahighsupra fin(Figurethe dorsal 6). of part middle the to opposed base, fin anal the the distance between the pectoral and middle of the in located is fin this spine; (short) rudimentary ananterior plus rays seven about of consists fin located just is pelvic the trunk. The above and the ventral border of rays 10 about of consists fin the SL (preanal pectoral length = 14 mm).The plus eight rays, andlocated in the last quarter of at least finfour is thin short, spines is composed of anal the SL (predorsal length =8mm).The of half first the of end the at just located is it rays; seven 4 thin spinesplus six or fin consistsof short dorsal is about 7 mm, and the body height is 8mm.The standard length isabout 18 mm, the head length the fish this In upward. curved somewhat trunk the skeletoncompletely preserved but withthe Description: articulated specimen(Figure 6). material: Referred the FishScaleFormation, inAlberta,Canada. ray, 1996,from the Early of Cenomanian deposits Type species: American Enchodus species. thesethree North one of to some with orbelongs features that suggest it is relatedits morphological is considered as anuncertain species of / Family Patterson, Sphenocephalidae 1964 Genus Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín lhuhbnso the head are strongly Although bones of Xenyllion High diamond shaped fish, with fish, shaped diamond High

Xenyllion zonensis Enchodus gladiolus Xenyllion Wilson andMurray, 1996 1969 IGM11193,complete and sp. Wilson andMur http://dx.doi.org/10.18268/BSGM2020v72n1a160819 includes Maas includes et al ., 2007). The ., 2007).The Enchodus - - - /72(1)2020 ; seems to be open along a groove. posterior The the preopercularcanal, which sensory the wayof along flange lateral a of impression an is There about 90°. of preopercular anangle form limbs bone. this Both the vertical limb of the height of tal limb, whose length is contained four times in an impression;however, itsshowsa shorthorizon No teethare exposed. the premaxilla. projected up to the anterior end of shallow notch. maxilla is abar-like The structure as the articular process, which isseparated by a about triangular processwith ascending short as premaxilla the orbit. The is the posterior edge of bones. lower The jawprojected is upward from Formation,Alberta, described in Canada, was Scale the Fish Early of Cenomanian deposits zonensis Rosen andPatterson,1969. Recently, was described as speciesfrom Westphalia, Sendenhorst, German Baumberg, Westphalia, Germany. Later, another 1839 (p. 4),fromof the Campanian deposits (1964): the species Patterson by erected fishes marine acantomorph Remarks: the anteriorhalf. with crenate posterior edge and strong circuli on epineurals. trunkiscovered The with cicloidscales this, there are two short neural arch andabove of thin neural spine. preural The a triangular 1 has skeleton, the preural acomplete, 2 bears long, and the anterior abdominal centra. In the caudal of supraneural bone located above the neural spines number, andsizes are unclear. There is asingle areepipleurals preserved; however, their position, respective arespines and thin. Some epineurals vity. Neural and hemalarchesare smallandtheir the abdominal ca of just covering half the dorsal About 12 thin and short preural ribs are present ing about 13 abdominals, 11preurals, andtwo urals. posteroventral preopercular edge. twothe in longspines there are impressionsof Also, sinuous. is flange preopercular this of edge The preopercleThe is almostentirely preserved as etba ouncnit f26centra, includ Vertebralof consists column Wilson andMurray, 1996,from the Shncpaia safml fextinct a familyof is Sphenocephalidae Sphenocephalus Sphenocephalus fissicaudus Sphenocephalus brachypterygius Sphenocephalus Artícle A160819 Agassiz, Agassiz, Xenyllion # - -

Artícle A160819 fin; pmx,premaxilla;pvf,pelvicfin. Figure 6 (Rosen andPatterson, 1969; Wiley and Johnson, monophyly and composition have been discussed et al. group wasnamedby2013). This Greenwoodrecord oldest the with dating back to the Cenomanian (Newbrey fishes, living of group Paracanthopterygii2016). The a problematic is (Nelson fins anal and dorsal the in spines a group well characterized for the presence of Acantomorpha, within inclusion its support fish and thegenus Sphenocephalidae thefamily a memberof 0005 is following paragraphs, it is demonstrated that IGM USA wasdescribed and added to Formation Mowry Vernal, in of deposits Utah, and Neuman,2013,from the Early Cenomanian stewarti member.as anewsphenocephalid Finally, Specimen IGM 11193, Specimen IGM The presence of dorsal andanal spines inthis dorsal presenceThe of (1966) asaSuperorder and sincethen its , Newbrey, Murray, Wilson, Brinkman Xenyllion Xenyllion . sp. from the San José de Gracia quarry. Abbreviations: af, anal fin; df, dorsal fin, pcf, pectoral pcf, fin, dorsal df, fin; anal af,Abbreviations:quarry. San JosédeGracia the from sp. http://dx.doi.org/10.18268/BSGM2020v72n1a160819 Xenyllion Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín . Inthe et al et al X. ., ., of a pelvic splint (= rudimentary anterior spine), splint (= apelvicrudimentary of include the presence Sphenocephalidae of ters and Newbrey thisorder. According toPattersonfamily of (1964) the single characters included in the diagnosis of Additionally, thisMexican specimen shows some ventral edge curved,unknown is IGM 11193. in these features, theopercle withpostero other of preopercle has large posteroventral spines. The Patterson (1969) are present in IGM11193,its described by RosenSphenocephaliformes and seen inthespecimenIGM11193. and Wilson352, 353)andMurray (1999) that are Paracanthopterygii (see Greenwood supraneural presentand asingle are features of preural, two in the caudal support, epipleurals acomplete second neural on spine presence of 2010; Borden n f w igotccaatr fthe order two diagnosticcharacters of One of et al. et al (2013),the diagnostic charac ., 2013; among others). The The ., 2013;amongothers). /72(1)2020 et al ., 1966,p. 11 11 - -

Late Cretaceous vertebrates from the San José de Gracia quarry RESULTS Late Cretaceous vertebrates from the San José de Gracia quarry RESULTS 12 12 Subfamily Yaguarasaurinae Palci, and Caldwell, because they share two characteristics: the trunk the genus here identified asanewrepresentative of is 11193 IGM results, their on Based exercisebetween species. thefour sphenocephalid Newbrey with four tofive spineseach. fin dorsal and anal as rays,well as fin sevenpelvic this premaxillary process isalmost entirely bar arises. whereangle Although the internarinal an acute rounded,is forming the posterior tapers is drop-like view;itsanterior edge shaped indorsal pemaxilla is anunpairedThe compound bone that availablespecimen is in Zavaleta-Villareal (2015). Description: maxilla andright(Figure 7). material: Referred America. the genus distributiongeographical of the of expansion the and Mexico in fish cephalid a spheno 11193 represents the first occurrence of IGM of finding the this, Despite precision. better itstaxonomic identity with to analysis determine 11193 prevents comparativeout adeeper carrying atriangularwave-form pattern. form apparently donotshow spines, andthe circuli scales whereas about 120°; aposteroventralform both limbs of obtuse angle thevertical preoperclar limb)and theheightof of atively long horizontal limb (its length is the half Sphenocephalus the preopercular vertical limb). Incontrast inboth short andcontained four times in the height of 90°andthe horizontal limb is of anangle form preopercle shows similar proportions (both limbs is covered with crenate cycloid scales, andthe / Superfamily Mosasauroidea Camp, 1923 Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín h orpeevto fthe skull inIGM poor preservationThe of Family MosasauridaeGervais, 1853 Orden Squamata Oppel,1811 S. fissicaudus S. et al species, the preopercles have a rel Papazzoni, 2013 Apeiiaydsrpino this A preliminary description of S. brachypterygius . (2013)provided acomparative IGM 11194, isolated pre has oval has cycloid spinoid are rather rounded, http://dx.doi.org/10.18268/BSGM2020v72n1a160819 Xenyllion Xenyllion in - - - /72(1)2020

anterior and posteriorly, withthe ventral edge (Figure 7). line; while the two posterior regions have noteeth previousregions two a teeth are presented forming thetwo bone anditprojects backwards. Ineachof this tudinal that rises behind the anterior edge of versal that runs from sideto side and other longi into four regions framed by two crests, onetrans nerve. the ophthalmic and exposes numerous pores of thisbone is curved dorsal-lateral surface of The the premaxilla. bar is at least as long as the rest of preserved asanimpression;the thin internarinal h curneo tooth rootslargely exposed in the occurrence of Accordingamong others). to Palci ofthe amember Family (see LeblancMosasauridae as identification its supports premaxillae) with four teeth in IGM11194 of acouple of the fusion premaxilla (resulting of Remarks: near tothetip(Figure 7). a weak anterior cutting edge or carina and form are rounded at the base, oval in the middle part, strongly curved backward. Incrosssection, teeth edge.maxillary All teeth are entirely smoothand they have broad bases protruding from the alveolar teeth have6, x,8,9,11.The crowns; ankylosed ordered following 1, 2,3,4,x, the dental formula alveolar placed on the maxillary border; these are thisbone(Figure 7). edge of this bone and are parallel to the ventral arranged arowform below the medial longitudinal edge of these nerve; the maxillary nine of rous pores of the maxilla shows nume Although the surface of the bone. axis protruding near the ventral edge of labial surface iscurvedThe and hasalongitudinal -ventral process where there are no more teeth. top, but its bottom develops athinandlongpostero thisbone is inclineddownward from the edge of extends over posterior twoteeth. The maxillary premaxilla-maxilla suture border is straight and onesomewhatstraight and the dorsal sinuous. The The right maxilla is anelongated bone taperedThe Ventrally, divided is and flat rather is bone this Eight conical teeth and three empty alveoli are Tepeec fanunpaired presence The of Artícle A160819 et al et al . (2013), ., 2012, # - -

Artícle A160819 premaxilla; pms,premaxilla-maxillasuture; ua,unoccupied oremptyalveolus. Figure 7 yaoopisspotn h oohl fsupporting themonophyly of synapomorphies the jaw bonesrepresents oneof medial view of fthissubfamily(Figure 7). of in IGM11194supporting itsrecognition as part Alfaedo Villages. characteristics These are present theS.di Anna middle Turoniannear of deposits and Fumanevillages) and the S.di Alfaedo Anna near of Santonian deposits fumanensis northen Italy, which recently were named as Verona,collected in Late Cretaceous localities of Romeosaurus the genusTexas, USA;aswell as to species of Ranch Limestone, at Cedar Hill, DallasCounty, Park Shale, approximately 15 cm above Kamp the Arcadia from themiddleTuronianof deposits 2000); Colombia (Páramo, 1994; Páramo-Fonseca, Villeta Formation alongthe Magdalena Valley, collected in Turonian sedimentsbelonging to monospecific genera, two include There genera. three the subfamily Yaguarasaurinae, which gathers SpecimenIGM 11194,indeterminated Yaguarasaurinaethe SanJosédeGraciaquarry.Abbreviations:mx,maxilla;pmx, from Russellosaurus coheni Russellosaurus Yaguarasaurus columbianus (from themiddleTuronian toearly Palci, and Papazzoni, Caldwell, 2013, Polcynand Bell, 2005, R. sorbinii http://dx.doi.org/10.18268/BSGM2020v72n1a160819 Páramo, 1994, Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín (from the R. maxillary teeth on maxillary teeth in lary suture isover maxillary-premaxillary three maxil teeth. The maxillary as the number and shapeof suture, thepremaxillary-maxillary well as the size of Yaguarasaurinae differ from of bers each other by and posterolateral carinae whereas teeth, entirelyand bearing weak smooth, anterior however, in curved backward inboth species of teeth are subcircular in cross section and strongly Rusellosaurus dinal striations, andprobably have no carenae in are teeth longitu contrary, medial coveredfine curved,strongly with and 3); fig. 2000, Fonseca, and withnocarenae in moderately curved backward, probably smooth and Bell, 2005, p. teeth are maxillary 323). The et al Fonseca, in in p.127),13–15 2000, 14–15 is teeth illary (Palci Among other characteristics, the nominal mem ., 2013,p. 600),and16in et al ,21,p 0) h ubro premax ., 2013,p. number 600).The of (Polcyn fig.Bell, 2005, and Finally, 2). R. fumanensis Romerosaurus Yaguarasaurus /72(1)2020 while it is over only two there are 15 maxillary Yaguarasaurus Yuguarasaurus Rusellosaurus and Romeosaurus R. sorbinii Russellosaurus Romeosaurus (Páramo- (Páramo- (Polcyn (Palci there 13 13 - - - - ;

Late Cretaceous vertebrates from the San José de Gracia quarry RESULTS Late Cretaceous vertebrates from the San José de Gracia quarry RESULTS FROM SANJOSÉDEGRACIA 3.2. OTHER ELEMENTSINTHEFOSSILASSEMBLAGE 14 14 this Mexican fossil the maxillary-premaxillary anypreviously known yaguarasaurine genus. In features that avoid itsbelonging to determining (Palci longitudinal ridges andapparentlycarinae withno with areteeth that 13maxillary are ornamented distorted, compressed inadorsoventral plane, but carbonate crystals. Practically allthefossilsfrom this siteare calcium with filled are cavities Bony contents also preserve similar phosphatic mass. gastrointestinal the fishes, small some In planes. as phosphatized on the bedding deposited masses San José these are preserved de Gracia quarry; plants, andvertebrates. scarce impressions of ostreids,inoceramids andother indeterminate that includecomplete ammonites, belemnites, mens. Untilnow, theyrepresent a fossilassemblage lated and fractured elements to complete speci iso from conservation, of degrees different show dada Autónoma de Puebla (BUAP). fossils These the Benemérita Universi ontological collection of UNAM,the pale within and asimilaramountis the Colección Nacionalde Paleontología (IGM), San José de Gracia have been catalogued into For now, about a hundred fossil specimensfrom Yaguarasaurinae inMexico. subfamily the of record first the represents 11194 IGM that goesbeyond thiswork. discoveryThe of effort major a represents IGM this desirable; is of 11194 identification taxonomic accurate the in which there is asmallanterior carina. Although middle and clearly compressed laterally in the tip IGM 11194are rounded inthe base, ovoid inthe smooth. Peculiarly, in cross section the teeth of to Contrary (13–16). yaguarasaurins nominal the in teeth (11) in IGM11194islower maxillary than of (extended over number twoteeth). The maxillary suture is shortasin / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín R. sorbinii G 19 hw euircmiaino IGM 11194shows a peculiar combination of Centimetric coprolites are also preserved in et al ., 2013,p. 601,figs. 2,4,and7). , teeth of IGM 11194are superficially 11194are IGM of teeth , Yaguarasaurus and http://dx.doi.org/10.18268/BSGM2020v72n1a160819 Russellosaurus - - - - /72(1)2020

or emptyalveolus. pmx, premaxilla;pms,premaxilla-maxilla suture; ua,unoccupied from theSanJosédeGraciaquarry.Abbreviations:mx,maxilla; Figure 8 Damian andLazăr, 2005). the Turonian (Riccardi, 1981; species index of (1813) that has been widely recognized as afossil species pers. observ.) the potential as representatives of (JAO, Alencáster Gloria by BUAPidentified were in the Some inoceramids specimensdeposited Coniacian (Kennedy andGale, 2015), respectively. to Turonian (Wright and Anisoceratidae, which ranges from Albian themrepresent the families Hamitidae some of ammonitesareheteromorph remarkable because that distortstheirshapeinthisplane. them show a horizontal elongation only someof San José deGraciaare pending;oncethese are invertebrates and the possible microfossilsof all the fossil studies of the moment, the formal in the San José(Figure 8).For deGraciaquarry Turonian age to the fossiliferous strata exploited above are considered,possible itis to attribute a Inoceramus labiatus and Anisoceratidae ammonites, the pelecypod Specimen IGM 11194, indeterminatedYaguarasaurinae11194, Specimen IGM Among the fossilsfrom San José de Gracia, the f h eprldsrbtoso Hamitidae the temporal distributions of If Inoceramus , andthe vertebrates described ( Mityloides et al ., 1996) and Albian to ) labiatus Artícle A160819 Schlotheim # Artícle A160819 4. Discussion In the south, Cambrian-Devonian shalesand San Joséis somewhat complex. deGraciaquarry new localitybelongtothat formation. this hence, there thatis nopossibility the rocksof theCoatzingo Formation; not matchwith that of theSanJosédoes deGraciaquarry of lithology the Zacapala Basin (Beraldi-Campesi, 2003).The low within energy in a lacustrine environment of volcanic ashandsandlayers deposited sequence of a covered with are that horizons flint intercalated limestoneswithsome iscomposedof formation thiscontinental Gracia (Figure baseof 1). The SanJosé de are exposed aroundthe town of the Coatzingo Formation of continental deposits Mexicano (2006) show that the Oligocene-Miocene alreadyMaps published byServicio Geológico accurate agefor thisdeposit. developed, it willbepossible toreacha more lacustrine travertines belonging to the Agua de Formations aswell as a Pleistocene sequence of the Pie de Vaca of lacustrine and alluvial deposits fluvial, of sequence Eocene-Oligocene a above, theTlayúa Formation; the eroded surfaceof the Balsas Group (Fries, 1960) lie on merates of units; different among these, the Paleocene-Eocene­ conglo to belonging sediments clastic Tepexi de Rodríguez show continental Tertiary parts near San Josétopographic de Gracia and lower Aptian-Cenomanian age. low The strata of marine at thickcomposed of least 300 meters sequence, which isrecognized asasequence of described the Tlayúa Formation asanincomplete San José deGracia,Pantoja Alor (1992) south of Tepexi de Rodríguez,town of a few kilometers Early Near the Cretaceous (Neocomian-Aptian). and calcareous lensesfrom the Upper Jurassic- Paleozoic strata, there are red terrigenous layers the region 1978).Above (Ortega-Gutiérrez, these the Acatlán Complex are the oldest rocksin of phyllites complex belonging to the metamorphic eie,telclgooia rmwr fBesides, the local geological frameworkof http://dx.doi.org/10.18268/BSGM2020v72n1a160819 Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Beraldi-Campesi, 2003; Applegate Beraldi-Campesi, 2003; Applegate Luna Formation are present (Pantoja-Alor, 1992; (Figure 1). conditions (Buitrón and Barceló-Duarte, 1980) fossil-lackingstrata where inshallow deposited belonging to Miahuatepec Formation, whose marinelimestones the Aptian of Gracia consists José de to theEastandNortheast highlands of Otherwise, marinedeposit. platform the hills and originally described byFries a shallow (1960)as the Morelos Formation, which was limestones of byformed the Cenomanian massive marine gray among others). however, the fossilassemblage of comparison theMexcala Formation; an outcropis of quarry observation, that to suggest the San José deGracía tempting, Itis 2000; amongothers). based onthis Hernández-Romano, 1999;Aguilera-Franco, Oaxaca (Alencáster, 1980; López-Ramos, 1983; exposed along Guerrero, Morelos, Puebla, and and hemipelagic environments, which todayis under moderate currents, low turbidity, andpelagic depocenter, the Morelos-Guerrero Basin,occurred within a marginalmarine sediments deposited Cenomanian-Maastrichtian is asequence of previouslydescribe by Fries formation (1960).This River, and belong to the Mexcala Formation, northwest20 km fromsite,this alongtheAtoyac exposed totheSanJoséare located deGraciaquarry of Puebla. region inthis observed time first the for unit San José deGraciafossiliferousstrata represent a near San José deGracia(Figure 1).Therefore, the Miahuatepec, andMorelos) previouslymapped the (Tlayúa, Cretaceous Early-middle marine limestone formations of any of part as identified be San José cannot de Gracia quarry marly strata of Pie de Vaca, fossiliferous andAguadeLuna).The previouslydocumented regionin this (Coatzingo, the Tertiary continental geological units any of the San José donot belong to de Gracia quarry to conclude that the marine strata exploited in Regarding itsgeological context,possible itis south toSanJosé hills The deGraciaare h eietr ok fTuronian age closest rocks sedimentary of The /72(1)2020 et al ., 2006; 15 15

Late Cretaceous vertebrates from the San José de Gracia quarry DISCUSSION DISCUSSION / CONCLUSIONS / Late Cretaceous vertebrates from the San José de Gracia quarry ACKNOWLEDGEMENTS / REFERENCES Acknowledgements 5. Conclusions 16 16 family Dercetidae). Today, more the recoveryof thegenus (the occurrence of et al. Guerrero alreadyreported byAlvarado-Ortega the Mexcalaand Taxo Formation in El Viejo) of Turonian sites(the Arroyo Bocas quarry Las from SanJoséwith thosefrom deGraciaquarry oto thisproject was provided by the UNAM port of improvement to this contribution. Financial sup AgustinMartinelli and Sergio allowed Bogan of observations2008. Thoughtful andsuggestions support and friendship during the fieldworks since for their thisquarry very to the workers grateful of fossilsfrom the SanJosé deGraciaquarry. We are the Aranguthy for sharingtheir knowledge of thank Messrs. Félix authors and Sebastian The Family (Figure Clupeidaeinthe country 8). the a specimenthat represents theoldestrecord of Dercetis as (such Mexico in recorded time first the for taxa previouslyknown Mexicoin (such as an unexpected diversity.includes taxa This assemblage recovered from thisnewsite represents required thisto idea.confirm The fossil vertebrate Mexcala Formation; additional geological data are Although it Puebla. of the theserockspossibleis to suggest part as of region in this time first fossiliferous Turonian rocksobserved for the marine Puebla, Mexico, constitute an outcrop of SanJosé deGraciaQuarry, marlsThe of geological unit remains asapending task. a formal these therocks belongingfinal to of recognition of the sequence that is being exploited; hence, the of problematic, given the small extension anddepth geological data intheSanJoséis deGraciaquarry / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín (2006b) shows scarceelements common , Xenyllion andYaguarasaurinae), as well as Enchodus http://dx.doi.org/10.18268/BSGM2020v72n1a160819 Enchodus andthe ), - /72(1)2020 References Agassiz, J.L.R., Agassiz, 1839,Recherches surlespoisons J.L.R., Agassiz, 1843,Recherches surlespoisons J.L.R., Agassiz, 1833,Recherches surlespoisons IN106011, andIN207314. through DGAPA-PAPIIT the grants IN225008, Alvarado-Ortega, J., Garibay-Romero, L.M., Alvarado-Ortega, J., González-Rodríguez, K.A., Alencáster, G.,del 1980,MoluscosMaastrichtiano Aguilera-Franco, N.,resolution 2000,High Neuchâtel, 296p. fossils, volume 4:Neuchâtel, Universitat Neuchâtel, Universitat Neuchâtel, 356p. fossils, volume 2(preface dated in 1843): Neuchâtel, Universitat Neuchâtel, 188p. fossils, volume 1(preface dated in 1833): 2006b, Lospecesfósilesde la Formación Vega-Vera, F.J., Centeno-García, E., Blanco-Piñón, A., González-Barba, G., Springer, 169–207. 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