,ln )Jq_tiJ Rivista Italianl di Paleontolosia e Str,rtisrafia volume 1 10 no. I 2 pls. YY'- ' -' April 2004

INNER \THORLS OF GREGORYCERAS (, PELTOCERATINAE) AS THE FIRST OCCURRENCE OF THE GENUS IN MEXICO

ANA BERTHA VILLASENOR . FEDERICO OLORIZ't{ ISABEL LOPEZ-PALOMINO

Recei.",ed Octobu 20, 2002; accepted Ooober 1, 2003

Key 'utords: Anrmonites, Gregoryceras, Upper , Oxford- siltstones. Some of the claystone horizons show calcar- ian, Mexico. eous concretions. The sampling has allowed the recos- nition of faunal assemblages characterized by ammo- Abstract. Tivo nuclei of Gregotlceras are described as the first record of the genus in Mexico. The occurrence in Mexico of the ubi- nites and bivalves. quitous Tethyan genus Gregop,ceras is interpreted as related to l nr;rjor Arnong the ammonite renlelins, the occurrence of flooding episode that affected ncritic shelves in the southern North- Gregorceras sp. in the beds 20 and 42 (Fig. 1) is of spe- American platc. cial relevance (Villasenor et alr.2002).

Riassurtto. Vengono descritti duc nuclei di Gr-egorl,ceras conre l;r prima testinronianza di tale qenere in Messico. Il ritrovamento in Mcs- Material sico dell'ubiquitario senere tetideo Cró,gor),ceras è messo in rcl,rz-ione,rd un principale episodio di annegamento, che ha interessato le pi;rttafornre Thrce inconrplete specimens have be en collected, two of which neritiche nclla parte nrerìdionale della placca Nordamericlna. sathered fronr bed 42 seenr to be nuclei. These inner moulds, IGM 6090 and IGM 6091 (Pl. l, fic. 2 - 5), l1 to l5 mm in sìze, do not show su- ture lines and therefore it is difficult to conclude if they represent nu- lntroduction clei, juveniles, or even snrall adults. The snrlll reported specinrens show the morpholorical ch,rr,rcters of the genus ,rnd rve favor their interpre- The research program on marine Upper Jurassic tation ls nuclei. Thev correspond to nroderarelv evolute shells, of 34- 367" involution. The u,horl-section is rounded and slightlv wider than rocks in Mexico is beins carried out since 1985, in a high. Thc ventrrl r('sion is u'ide, slightlv convex and shows a subtle proiect with the collaboration of the University of Mex- smoothing ,rt the nrid-r'entcr line. ico [UNAM] and the University of Granada, Spain. In Thc unrbilicus ìs deep and bordered b1'a near-vertical umbili- this context, the precise recognìtion of the biostratigraph- cal wall. The unrbilicrl edge is widely rounded. The shell smaller than ic distribution of ammonite faunas, eathered bed-by-bed 6 mm is snrooth sincc the absence of sculpture cannot be releted to poor preservation. At shell size bigger thln 6 mm the sculpture is from several Upper Jurassic type-sections in the north- nrade of ribs. These ribs do not reach the unrbilic;rl uall and some of eastern pan of Mexico. is in progress. thenr nrav be connected on the periumbilicel c.dge. to deternrine crcs- Recently, a bed-by-bed sampling on the type sec- cent-like couplet o[ ribs, but no nodes. On the flanks, the ribbing tion of the Santiago Formation has been made in Cen- is tvpicalll'rursiredirte shos.ing e u'ide inilexion (-lO to 48 degrees) tral-East Mexico. The studied section is located in the close to the mid-flank. The bisser of thc studied tu'o nuclei (IGM southeastern part of the State of San Luis Potosí, near 6090) shou's ln incre;rsins nunrber of defective bifurc;rtions torvards the ldor,rl cnd oî thc preserved inner mould. This feature is typìcal the town of Tamín on the left bank of Rio Moctezum:r for nearlv cor.nplete.rdult shell, but clnnot bc interpreted as snrlll- (Fig. 1). The lowermost 22 rn of the section are nlade size but inconrplete adult individual becluse no rÌttenu:ltion of sculp- of scarce linrestone beds and dominant claystones and ture exists (Pl. I, fig. l).

Instituto de Geología, Dep:rrtlnrento cle Paleontologí1, Univcrsid;rd N;rcion,rl Autónonrl de México, Ciud;rcl Universitaria 0.15 10, Coyoacín, México, D. F. Mexico. E-nr;ril: ;rn;ib(a scrvidor.unanr.rnx; nictehlT3(r vahoo.cor.r.r.r.r.rx Facultad de Ciencias, Depart;rnrento de Estratigrafía v P;ileontologíe, Universidad de Granad;r, Fuentenueva s/n, 18002, Granade, Spain. E-n-r;ril: foloriz(ru usr.es 254 A. B. \:illaseúor, F. Okjríz Ei I. L(tpaz-Palonino

Vith respect to Gregowceras (Gregoryceras) chon- gi Gvgi & Hillebrandt (1991) rnd close forn.rs reported from Chile (Pl. 1, fig. 6,7),rhe differences,rre related to the level rt which the con.rplete ribs end. In the Chileirn spe cirnens, the inner extrenle of tlie ribs re2lchcs the unr- bilical iv,rll, but it is ir.nportant to note that sonre speci- r"nens of this spe cies sl.rorv shorter ribs. Morphological sin-ril;rrity rrlso exists n,ith the ubiquitous European species trdnsaersaritnrl, which is al- so known from Chile accordins to Gygi & Hillebrandt ( 199 I ), in p:rrticul;rr throush the rounded umbilical edee and the irrched bent of the ribs (Pl. 1, fig. 11, l4). Horv- - 43Jq - 42ae) Gregoryceras sp. ever, the Mexicen specin"rens differ in their less infl:rted depressed whorl section; as far as the whorl section of @ - 4OeJE nuclei in trans'oersttriuzzr is known frorlr the specintens -'-..--,--..] studied by Gygi (1977) (Pl. l, fig. 10, 14). Conrparison ',-'-_.-,-,-] 4OdJ e) - of inner rvhorls is also possible with the Er.rropean spe- z - -.-.-.-.- - 4OcJE cies GregorTceras (Grego:ceras) riazi.In fact, the beein- f- - 4Ob.9 ning of ribs occr,rrs at equivrrlent shell diameters, but, in - 4OaJq this c:rse, r.r.rorphological diff-erence has beer.r identified in - 4l)a9 the ;rbsence of periurrbilical nodes in the Mexican speci- mens, in their less coarse ribs, in their lcss flat and less conversent flanks, and ,rlso in their r.nore globose shell - 35Jq 1Pl. l. iis. S. e. 12, ll). On the basis of the conrparative :rn;rlysis, morpho- logic sir.ìrilarities and differences between the Mexican specinrens ,rnd the previously nlentioned species fron.t Chile and Europe have been est,rblished. However, the - 3()9 snr,rll siz-e of the specin.rens found in the Santi:rgo For- m:.rtion at T:rmín impedes any conclusive interpretltion -3E2e of con-specific rel:rtionships with sonre of then-r. There- - a3J2 Gregoryceras sp. -',2J91",. fore, u,s f,rvor the inte rpretirtion of the Mexicirn material - aoJc) - ,rnalyzed as GregotTcerds sp.

Remarks on biochronology and palaeobiogeography f.-j Claystone and siltstone Claystone with concretions i=B According to Gygi 8c Hillebrandt (1991), who ffiffi Limestone r. r 'r -'-"------@) necora of ammonites sunrnr,rriz-ed their proposrrl for identific;rtion of Grego- r Record of bivalves ucerds irt the spccies level, the followine morphological fe :ltLlrcs arc of interest for biostrrtigraphy: 1) the rib ,rp- liig. I - Geogrrphic loc.rtiot.t of thc'stutlicrl ,ìrcl ,tncl strutigrrphic sec- ?O tion of thc los'emrost prrt of tìre Santi.tqo Fonrltion rt thc pearance at less th;rn nrnr; l) the bending of ribs by tvpe rre.ì close to Tlnr,in, Srn I-uis Potosí St:rte. Notc'strrrti- anqles greater than 25 degrees; and 3) the simple ;lrcuate- gr,rphic ìrorizons cont,rinins Gregrnl,ccr,ts sp. in the studicd sh,rpe of ribs. All these fcatures point to :ln asL' equiv:r- t\'fic sectlor.t. lent to the errrli, tansversariun.r Chron (non bas,rÌ Ante- ccdens subchron) in Europe. Thercfore, we prelir.r.rinary cnr isrige :rn cqrrii.rlent rqe I-or thc Mcxicrrrr spt'cirttcns collectcd fron.r the lorver part of the tvpe section for the The rrbove Taxonomic remarks Santi,rgo Forrt:rtion ;rt Tanriir.r, San Luis Potosí. described Gregoryceras inrply the first rcport of this ge- The smtrll size of our specimens makes difficult nus in Mexico. and confirrl the rvide distribution of the the right identification at the species level. Morpholoui- genLls in the Anrcricas. Previous citltions of this genr.rs cal similaritv exists with the inner whorls of the spccies rvere made by Gygi & Hillebrandt (1991) in the north- Grego:ceras (Gregoryceras) chongi Gvgi 8r Hillebrrndt, ern part of the Cordillera de Don.revko in Chile, :rnd bv G r ego w c e ra s (G re go w c era s) t ran s'u er s arieriz ( Q u e n s te dt ), Young & Olóriz (1993) in the northern rim of the Gulf and Gregorycerds (Gregorytcerds) riazi (De Grossouvre). of Mexico. In pirlireobiogeogrirphic terms, the comprìrxtive Taphonon\, of Oxfordian ammonites frottt Southern Spain 251

widity rounded incomptere :"":1,1"j ,:"rilÎ|" umbilical edge bifurcation t\'"-..

rl/."':::;:"" F. 6*- * shell \__..\ "v beginning ot ribs 5 at6mm Jt ",3_, * 'fclf 5

1 't, 1'"Î,,,

PI-ATE 1 Fig.l.5.Gregot1,ge7,15sp.fronlthestudiedscctìon;I.L;rterll.rdorallncl\'entralViewsofIGM6O9O'shorvingthenlostsignific.ìntmorphologicl characters (X2). Fig. 2-5. Left- ,rnd right-sidc vieu's of the clescribed snr;rll inner nroulcls (iGM 6090 and IGM 6091, respectivelv) (Xl). Fig. 6. Re- production of inner rvhorls of Cregorl,cer,ts (GregoD'6gv'755 cf . chongi G;'ui Ei Hillebrendt ( 1 99 I , pl. 3. f ig. 9e). Fir:. Z. Reproduction ol Crego:cer ts (Crego4tceras) chongi Gvgi Ei Hillebnndt (1991, pl. 5, fig.-{a). Fir. 8-lO. Axial sections of iGM 6090,;rnd the nuclej of Europern Gregorlceras (Gregon,6gyú51riaziln,Jt).Lt|IsLrersLtTiun c[.tr,nstvrslriunz(Quenstedt).lsillustrltedinGvgic\Hillebrrndt(l99l,pl.Z,fiu.,+a).Fig.l2'ReproductionofChilernspecinlen cer.ts (Gregon'ccrLrs) riúzi (Dc Crossouvre) .rs illustrerecl in Gvgi c\ Hillebrandt (1991, pl. 3, fig. 8a). Fìs. 13. Reproduction of thc rxirrl section of Fiuropean Gregotltcer"ts (Gregoryceras) rini (De Grossor-rvre) rccordìng to G,vgi (1922, pl. 10, fir. 7). Fir. l.+. Reproduction oi the axial section ol L.urope:rn specirncn ol G rego4, cert s of Europeln species for conrparison. All figures in rìiìtllrel size orheru'ise indìclteci.

analysis made 21t small size cannot be conclusive due to tion of phenotype traits identified in ubiquitous species present limitirtions to the precise knorvledse of the mor- adapted to epioceanic and epicontinental environments. pholoeic vari:rbility during the e:rrly ontosenv in species On this basis, the occurrence of nuclei of Gregoryceras of genus Gregoryceras. showing such ;i mosaic morphology in epicontinental shelf However, the studied specimens show the combina- deposits from Mexico cannot be surprising (Fìe. 2), and 252 A. B. Villasefior, F. Olóriz &. L López-PaLomino

Fig.2 -Oxfordian paleogeographic mtr '.l"nt,',1 frnnr Srnith et al. ('l995) showing the distri- bution of the genus Gregoryc Germany- Switzerland erds. D:ltabase in the Appen- H: Hungary dix. l: ltaly In: lndia M: Madagascar Mx: Mexico P: Poland R: Russia Eq.ator S: Spain - -\-r- î Tunisia US: USA

l_l rueritic environments Oceanic/ Epioceanic envtronments

presumably relates to transgressive pulses in the area (see and for more variable trends related to palaeogeographic Oloriz et al. 2003 for extended trearmenr), as well as oc- restructuring in the Americas (Riccardr et al. 1992;Mar- curred in other neritic environments far from European zolf 2002). The Early to Middle Oxfordian was also the shelves and their epioceanic fringes (e.e. India: Krishna time for major palaeogeographic change in peripheral ar- et al. 2000). eas of the central north-Atlantic and the Gulf of Mexico Discontinuous records of late-Early to Middle Basin and related areas (Emery & Uchupi 1984; Jansa Oxfordian ammonite assenlblages, registered in more 1986; Salvador 1987, 1991; Michaud tr Fourcade 1989; or less calcareous horizons intercalated between mainly Olóriz et al. 2003). terrigenous sediments elsewhere in South America, ac- cord with this interpretation (Gygi & Hillebran& 1991 for northern Chile, includinu Gregoryceras; Hillebrandt Conclusions & Gròschke 1995). In addition, ammonite assemblages The present record Gregoryceras Tamín from age-equivalent, discontinuous horizons, and con- of from opens nev/ perspectives for the future precision in the taining a variable xmount of widespread Tehyan genera biostratigraphic interpretation Mexican ammonite (or ammonites with Tethyan-like morphology), have been of assemblages known Burckhardt (1912), and rein- reported even southwards from Argentina (Hillebrandt from forces the known occurrence of Tethyan perisphinctids et al. 1992; Riccardi et al. 1992; Parent 1998), and can in Mexico during the 1912; Cal- be correlated with the stratigraphic interval showing bi- Oxfordian (Burckhar& lomon 1992;Myczynski et al. 1998). The occurrence of valve-rich assemblases interpreted as resulted from trans- the genus Gregoryceras in Mexico was favored by the ma- gressive pulses in these resions (Damborenea 1996). A jor Oxfordian flooding affecting epicontinental shelves similar but slightly younger Oxfordian case, includine all over the proto-Atlantic. Gregoryceras, has been recently reported from lndia by Krishna et al. (1995, 2002). All this well adapts the widespread context of discontinuous record of an.rmonite assemblages in the Aclenotledgenterrs. This research s'as supported by CONACyT Americas throughout the Lower to Middle Oxfordian Prolect 27555T fronr Mexico, and the EMMI Group of the University of Gran;rde (RNM lZ8, Junta de Andalucia; Project BTE2OO24A29), (Imlay 1984, for North America; Riccardi er al. 1992, f or Spain. The ;ruthors are indebted to C. D'Arpa (Universitv of Palcrmo) South America), which corresponds to the tinre for nra- rnd A. Gal:icz (Eótvos [-. Universitr', Budapest) for valuable sugges- jor transgression in North America (Taylor et ;rl. 1984), tions m,rde on ln e;rrly draft of this paper. Tapbonoml, of Oxfordian ammonites from Southern Spain 253

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Appendix

(Synthetic bibliography for the dat:r base used to consider records of the genus Gregotl,cerasas depicted in Fig. 3): [Al] Algeria: Atrops 6i Benest (1984) [B] Bulgaria: Sapunov (1976);Gvgi (1977) [Ch] Chile (Cordillera de Donrevko): Gvgi & Hillebrandt (1991) IF-G-S] Frarrce, Germant', Su,itz-erland: GySi (1977) [Hl Hungarv: l"ózy &: Meléndez (1996) [I] Italy (Sicily): Gemmellaro (1874, 1827, 1882); Sarti (1984, rlso for reference to classic papers); D'Arp;r & Meléndez (2002) IIn] India: Krishna et al. (1995,2002) [M] Madauascar: Collignon (1959); Gvgi (1977) [Mx] Mexico (Central-East): Vilhsenor et al. (2002) [P] Poland (Pieniny Klippen Belt): Gvgi (1977); Malinou'ska (1988) [R] Russia (Russian Plate): Arkell (1956); Rogov (2001). [S] Spain (Southern Spain, Subbetic): Sequeiros (197.1); Caracuel ct rl. (2000). (Southern Spain, Prebetic): Glrcía-Hcrníndez et al. (1979): Olóriz ct al. (1999). (Iberian Range): Arkell (1956); Meléndez., (1989). (Balearic Islands): Fallot (1922)r Olóriz et al. (1996). [T] Tunisia: Gygi (1977) [US] United States of America (Northern rirn of the Gulf of Mexico): Young E: Olóriz (1993)