Chronostratigraphy and new vertebrate sites from the upper Maastrichtian of Huesca (Spain), and their relation with the K/Pg boundary

Accep ed Manusc ip Ch onos a ig aphy and new e eb a e si es om he uppe Maas ich ian o Huesca (Spain), and hei ela ion wi h he K/Pg bounda y E. Pué olas-Pascual, I. A enillas, J.A. A z, P. Cal ín, L. Ezque o, C. Ga cía-Vicen e, M. Pé ez-Pueyo, E.M. Sánchez-Mo eno, J.J. Villalaín, J.I. Canudo PII: S0195-6671(17)30487-1 DOI: 10.1016/j.c e es.2018.02.016 Re e ence: YCRES 3817 To appea in: C e aceous Resea ch Recei ed Da e: 9 No embe 2017 Re ised Da e: 24 Janua y 2018 Accep ed Da e: 22 Feb ua y 2018 Please ci e his a icle as: Pué olas-Pascual, E., A enillas, I., A z, J.A., Cal ín, P., Ezque o, L., Ga cía- Vicen e, C., Pé ez-Pueyo, M., Sánchez-Mo eno, E.M., Villalaín, J.J., Canudo, J.I., Ch onos a ig aphy and new e eb a e si es om he uppe Maas ich ian o Huesca (Spain), and hei ela ion wi h he K/ Pg bounda y, C e aceous Resea ch (2018), doi: 10.1016/j.c e es.2018.02.016. This is a PDF ile o an unedi ed manusc ip ha has been accep ed o publica ion. As a se ice o ou cus ome s we a e p o iding his ea ly e sion o he manusc ip . The manusc ip will unde go copyedi ing, ypese ing, and e iew o he esul ing p oo be o e i is published in i s inal o m. Please no e ha du ing he p oduc ion p ocess e o s may be disco e ed which could a ec he con en , and all legal disclaime s ha apply o he jou nal pe ain. MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT Ch onos a ig aphy and new e eb a e si es om he uppe Maas ich ian o Huesca (Spain), and hei ela ion wi h he K/Pg bounda y E. Pué olas-Pascual 1,4 , I. A enillas 5 , J.A. A z 5 , P. Cal ín 2 , L. Ezque o 3 , C. Ga cía-Vicen e 5 , M. Pé ez- Pueyo 1 , E.M. Sánchez-Mo eno 2 , J.J. Villalaín 2 , J.I. Canudo 1 1 G upo A agosau us-IUCA, Á ea de Paleon ología, Dp o. Ciencias de la Tie a, Uni e sidad de Za agoza, 50009 Za agoza, Spain edua do.pue [email protected]; [email p o ec ed]; manuppuey[email p o ec ed] 2 Dp o. de Física, Facul ad de Ciencias, Uni e sidad de Bu gos, 09001 Bu gos, Spain cal inballes e @gmail.com; [email p o ec ed]; ill[email p o ec ed] 3 G upo GEOTRANSFER, Á ea de Es a ig a ía, Dp o. Ciencias de la Tie a, Uni e sidad de Za agoza, 50009 Za agoza, Spain lope@uniza .es 4 Faculdade de Ciências e Tecnologia- GeoBioTec, Uni e sidade No a de Lisboa, 2829-526 Mon e de Capa ica, Po ugal pue olas@ c .unl.p 5 IUCA, Á ea de Paleon ología, Dp o. Ciencias de la Tie a, Uni e sidad de Za agoza, 50009 Za agoza, Spain [email p o ec ed]; josea z@uniza .es; ch is ianga cia icen [email protected] Abs ac : The ansi ional-con inen al acies o he T emp Fo ma ion wi hin he Sou h-Py enean Cen al Uni (Spain) con ain one o he bes con inen al e eb a e eco ds o he Uppe C e aceous in Eu ope. This Py enean a ea is he e o e an excep ional place o s udy he ex inc ion o con inen al e eb a es ac oss he C e aceous/Paleogene (K/Pg) bounda y, being one o he ew places in Eu ope ha has a ela i ely con inuous eco d anging om he uppe Campanian o lowe Eocene. The Se aduy a ea, loca ed on he no hwes lank o he T emp syncline, has seen he disco e y o abundan e eb a e emains in ecen yea s, highligh s being he p esence o had osau id dinosau s and eusuchian c ocodylomo phs. Ne e heless, al hough hese deposi s ha e been p o isionally assigned a Maas ich ian age, hey ha e no p e iously been da ed wi h absolu e o ela i e me hods. This pape p esen s a de ailed s a ig aphic, magne os a ig aphic and bios a ig aphic s udy o he i s ime in his a ea, making i possible o assign mos e eb a e si es om he Se aduy a ea a la e Maas ich ian age, speci ically wi hin pola i y ch on C29 . These esul s con i m ha he e eb a e si es om Se aduy a e among he mos mode n o he Uppe C e aceous in Eu ope, being e y close o he K/Pg bounda y. MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT Key wo ds: K/Pg bounda y; Maas ich ian; magne os a ig aphy; bios a ig aphy; e eb a es; Py enees. In oduc ion Recognizing he K/Pg (C e aceous/Paleogene) bounda y in con inen al deposi s is a complica ed ask due o se e al biases ha a ec he con inen al eco d (Ba e e al., 2009; Bu le e al., 2011; Smi h e al., 2001; Mannion e al., 2011; Smi h and McGowan, 2011; Upchu ch e al., 2011). E en so, g ea e o s ha e been made in ecen yea s o de ec he con inen al K/Pg bounda y and asce ain i s ela ion wi h aunal and lo al ex inc ions, especially in No h Ame ica (e.g., Fas o sky and Sheehan, 2005; A chibald e al., 2010; B usa e e al., 2015; and e e ences he ein), bu also in Eu ope (Canudo e al., 2016; and e e ences he ein) and Asia (Jiang e al., 2011; and e e ences he ein). In he Eu opean scena io, he g ea es di icul y in knowing how he e eb a e aunas we e a ec ed by he K/Pg ex inc ion e en is as a esul o he agmen a y na u e o he con inen al geological eco d du ing he La e C e aceous and ea ly Paleogene. None heless, majo ad ances ha e been made in he las ew yea s, and new ou c ops, mainly in Romania, F ance and Spain, a e being disco e ed and da ings ca ied ou (Pué olas-Pascual e al., 2016; and e e ences he ein). In Eas e n Eu ope (Romania), he Maas ich ian con inen al e eb a e assemblages ha e been examined and da ed by bios a ig aphy, magne os a ig aphy and adioiso opic echniques (An onescu e al., 1983; Van I e beeck e al., 2005; Cod ea e al., 2010, 2012; Panaio u and Panaio u, 2010; Boja e al., 2011; Panaio u e al., 2011; V emi e al., 2014; Csiki- MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT Sa a e al., 2015). Howe e , he e eb a e si es a e loca ed in excessi ely b oad age anges, he co ela ion be ween he di e en si es emains p oblema ic and mo e accu a e da ings a e equi ed (Bu e au and Le Loeu , 1991; Ghee b an e al., 1999; Cod ea e al., 2012; V emi e al., 2014). Simila conce ns occu wi h espec o he Uppe C e aceous and lowe Paleogene o F ance. Apa om he P o ence a ea, whe e se e al bios a ig aphic, magne os a ig aphic, chemos a ig aphic and sedimen ological s udies wi h a good ch onos a ig aphic con ol ha e been ca ied ou (Cojan e al., 2003; Cojan and Mo eau, 2006), mos o he no he n Py enees s ill lack accu a e da ings o co ela ions (Bu e au and Le Loeu , 1991; Lau en e al., 2002). The e o e, despi e he abundan Maas ich ian e eb a e ossil eco d eco e ed om sou he n F ance, only limi ed bios a ig aphic da a (Bessié e e al., 1980, 1989; Bilo e, 1985; Ga cia and Vianey-Liaud, 2001; Ma y, 2001) and one new magne os a ig aphic s udy (Fonde illa e al., 2016b) a e a ailable, and u he s udies and co ela ions a e s ill necessa y (Dina ès-Tu ell e al., 2014). The con inen al e eb a e eco d o he uppe mos C e aceous o Spain is one o he mos comple e and mos s udied in Eu ope (e.g., Company and Szen esi, 2012; O ega e al., 2015; Pe eda-Sube biola e al., 2015; Canudo e al., 2016). Mos o hese e eb a e si es a e loca ed wi hin he T emp Basin, in he Py enees o A agon and Ca alonia (Spain), speci ically in he Maas ich ian ansi ional and con inen al acies o he T emp Fo ma ion. The T emp Fo ma ion has been exhaus i ely p ospec ed and s udied, p o iding abundan new e eb a e ossil emains including dinosau s, c ocodylomo phs, es udines, mammals, ishes, amphibians and squama es (e.g., López-Ma ínez e al., 1999, 2001; Peláez-Campomanes e al., 2000; Rie a e al., 2009; Pe eda-Sube biola e al., 2009; Blain e al., 2010; C uzado-Caballe o e al., 2010, MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT 2013, 2015; Pué olas e al., 2011; Ma mi e al., 2012, 2016; Vila e al., 2012, 2013, 2015; Mo eno-Azanza e al., 2014; Pué olas-Pascual e al., 2014, 2016; Sellés e al., 2014a, 2014b, 2016; Blanco e al., 2014, 2015a, 2015b, 2016, 2017; Company e al., 2015; To ices e al., 2015; Canudo e al., 2016). In addi ion o he eco d o he e eb a es hemsel es, he e is a ela i ely con inuous geological eco d anging om he Maas ich ian o he end o he Thane ian (López-Ma ínez e al., 2006), which is p obably he bes da ed and co ela ed in Eu ope o his ime in e al and which may con ain he C e aceous/Paleogene bounda y. This makes he sou he n Py enees and he T emp Basin one o he bes a eas in he wo ld o s udying e eb a e associa ions ac oss he K/Pg bounda y, allowing compa isons wi h he ex inc ion pa e ns epo ed om o he pa s o he wo ld (B usa e e al., 2015; Csiki-Sa a e al., 2015; Canudo e al., 2016; Pué olas-Pascual e al., 2016). E e since he 1980s, he e o e, a g ea e o has been pu in o da ing he ossil e eb a e si es and sea ching o he K/Pg bounda y wi hin he ansi ional and con inen al deposi s o his sec o o he Py enees. Ou s anding in his con ex a e wo ks on he bios a ig aphy o udis s (Vicens e al., 2004), cha ophy es and palynomo phs (Feis and Colombo, 1983; Médus e al., 1988; Galb un e al., 1993; López-Ma ínez e al., 2001; Villalba- B e a and Ma ín-Closas, 2011, 2013; Villalba-B e a e al., 2012; Vicen e e al., 2015), o amini e s (López-Ma ínez e al., 2001; Díez-Canseco e al., 2014), on eggshells (Vila e al., 2011; Sellés e al., 2013; Sellés and Vila, 2015), magne os a ig aphy (Galb un e al., 1993; Oms e al., 2007; Pe eda-Sube biola e al., 2009; Vila e al., 2011, 2012; Canudo e al., 2016; Fonde illa e al., 2016a) and dinosau occu ences (Rie a e al., 2009; Vila e al., 2016). MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT The Se aduy a ea, loca ed in he A agonese no hwes e n b anch o he T emp Basin, has been p ospec ed by he A agosau us-IUCA esea ch g oup o he Uni e si y o Za agoza o o e 10 yea s. This has esul ed in he disco e y o a ound 40 new paleon ological si es wi h hund eds o e eb a e emains. These indings include impo an specimens such as he holo ype o he eusuchian c ocodylomo ph Aga esuchus subjunipe us (Pué olas-Pascual, Canudo and Mo eno-Azanza, 2014) and he smalles had osau id known in Eu ope o da e (Company e al., 2015), p obably a new dwa axon. Despi e he impo ance and po en ial o hese e eb a e si es, a ch onos a ig aphic amewo k o he Se aduy sec o has no ye been p o ided. Se aduy is loca ed be ween o he a eas wi h e eb a e si es such as Campo o he wes and A én o he eas , co esponding o he no hwes e n-mos b anch o he T emp Fo ma ion wi hin he T emp Basin. In hese nea by sec o s (Campo and A én, Huesca), p e ious magne os a ig aphic s udies ha e s a ed ha he e eb a e si es o he T emp Fo ma ion in hese a eas lie wi hin magne ic pola i y ch ons C30n and C29 , being la e Maas ich ian in age (Pe eda-Sube biola e al., 2009; Canudo e al., 2016). On he basis o wo ks o magne os a ig aphy (Fonde illa e al., 2016a) and bios a ig aphy (Díez-Canseco e al., 2014) on he mo e eas wa d-lying Isona sec o o he T emp syncline, howe e , some au ho s ha e de ec ed he possible p esence o impo an hia uses in some a eas o he T emp Basin. These gaps e eal ha mos o he succession and e eb a e con en in ha a ea co ela es o he ea ly Maas ich ian (mos ly ch on C31 ), sugges ing an olde age (Fonde illa e al., 2016a) o many e eb a e si es han p e iously hough (Vila e al., 2012). In acco dance wi h hese new da ings (Pe eda-Sube biola e al., 2009; Díez-Canseco e al., 2014; Canudo e al., 2016; Fonde illa e al., 2016a), he ch onos a ig aphic s udy o he a eas o Campo-Se aduy-A én may hus acqui e g ea e ele ance, gi en ha he MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT lowe pa o he T emp Fo ma ion exposed in Campo and A én con ains he only con inen al eco d o ch on C30n in he whole T emp syncline (Fonde illa e al., 2016a). To es all hese hypo heses, we he e desc ibe o he i s ime he magne os a ig aphy, bios a ig aphy and a p elimina y s udy o he ossil e eb a e assemblage o he Se aduy a ea. Acco ding o ou bios a ig aphic and magne os a ig aphic esul s, mos o he “lowe ed uni ” o he T emp Fo ma ion up o he op would be included wi hin magne ic pola i y ch on C29 , and mos o he e eb a e si es would he e o e ha e a la e Maas ich ian age, being loca ed e y close o he K/Pg bounda y. Un o una ely, he paleomagne ic and bios a ig aphic da a om he lowe hal o he s udied sec ions a e no conclusi e enough o gi e a speci ic age o e eal he p esence o hia uses in his a ea. Geog aphical and geological con ex The s udied a ea is loca ed in he A agonese pa o he T emp Basin wi hin he Py enean ange (Se aduy a ea, Huesca, Spain) (Fig. 1A, B). The Py enees a e a 430-km-long eas -wes -o ien ed con inen al collisional old-and- h us bel , loca ed in he no heas e n Ibe ian Peninsula be ween F ance and Spain (Fig. 1A); hey o med as he esul o he oblique collision and comp essi e episodes be ween he Ibe ian mic opla e and he Eu opean pla e. This p ocess ook place du ing he Alpine o ogeny, om La e C e aceous un il ea ly Miocene imes (Ga ido- Megías and Ríos, 1972; Puigde ab egas and Souque , 1986; Muñoz, 1992; A dè ol e al., 2000; Sibue e al., 2004; Teixell, 2004). The T emp Basin is loca ed wi hin he Sou h-Py enean Cen al Uni o SPCU (Ségu e , 1972), which co esponds wi h he cen al sec o o he Sou he n Py enees (be ween he Nogue as h us aul in he no h and he Sie as Ma ginales on al h us in he sou h) (Fig. 1A). Se e al syn-sedimen a y synclines (Age , T emp, Coll de Na go and MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT Vallceb e) associa ed wi h he emplacemen o sou h- e ging h us -shee s de eloped du ing he La e C e aceous, ac ing as di e en sub-basins (Fonde illa e al., 2016a; Oms e al., 2016). The ossil emains s udied he e a e om he A én and T emp Fo ma ions (Mey e al., 1968) wi hin he T emp Basin and he no he n lank o he eas -wes -o ien ed T emp syncline. The T emp Fo ma ion, which o ms pa o wha is in o mally known as he “Ga umnian” acies (Leyme ie, 1862), was deposi ed du ing he Uppe C e aceous–Paleocene, when he Py enees Basin was comple ely illed by coas al and con inen al deposi s due o he end-C e aceous ma ine eg ession (Rosell e al., 2001), ep esen ing he las in illing episode o he Sou h-Py enean Basin (Mey e al., 1968; López-Ma ínez e al., 1999; Oms e al., 2016). The SPCU has an ex ension o 5000 km 2 , o which he T emp Fo ma ion is es ima ed o encompass abou 1000 km 2 (López-Ma ínez e al., 2006). This o ma ion c ops ou in he cen al and wes e n pa o he SPCU, eaching a hickness o abou 900 m in he depocen e nea he locali y o T emp (López-Ma ínez e al., 1999). In he no he n a eas, such as he T emp syncline, he bo om o he T emp Fo ma ion is unde lain by and la e ally in e digi a ed wi h uppe Campanian–Maas ich ian mixed-pla o m ma ine deposi s ha co espond o he beach, ba ie - island and del aic sands ones o he A én Fo ma ion (Fig. 1C) (A dè ol e al., 2000). In he sou he n ou c ops, such as he Àge syncline, he A én Fo ma ion is eplaced by mo e calca eous deposi s co esponding wi h he limes ones Les Se es Fo ma ion (Souque , 1967; López- Ma ínez e al., 2006). Abo e, he T emp Fo ma ion is o e lain by Ile dian (lowe Eocene) ma ine sedimen s o he al eoline limes ones Cadí Fo ma ion, o ma ly deposi s la e ally equi alen o he Figols G oup (Fonnesu, 1984; Eichensee and Lu e bache , 1992; López- Ma ínez e al., 1999, 2006). MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT The sedimen a y succession o he T emp Fo ma ion can be linked wi h wo main s ages o comp essi e ec onics (Puigde àb egas and Souque , 1986). The i s s age occu ed du ing he la e San onian–la e Maas ich ian and was cha ac e ized by ac i e ec onics, pa icula ly in ense du ing he la e San onian and Campanian, which caused he in e sion o he p e ious Mesozoic i s uc u es and he de elopmen o a o eland basin. Du ing his pe iod, he basin was illed by mainly siliciclas ic deposi s which p og essi ely passed o ma ine acies owa ds he wes , whe e he open sea was loca ed. The second s age occu ed du ing he la e Maas ich ian–ea ly Eocene, his being a pe iod o smoo h ec onics and almos uni o m subsidence ep esen ed p edominan ly by ca bona e-ma l deposi s. The T emp Fo ma ion has been di ided in o di e en local uni s by se e al au ho s. Cue as (1992), and la e Pujal e and Schmi z (2005), di ided he se ies in o i e o ma ions and ou membe s, ele a ing he T emp Fo ma ion o he ca ego y o g oup. Howe e , he classi ica ion o he T emp Fo ma ion as a g oup is no widesp ead wi hin he li e a u e and some au ho s indica e ha he bounda ies be ween he o ma ions o his g oup can be con using (Rie a e al., 2009). Fo his eason, Galb un e al. (1993) and Rosell e al. (2001) di ided he T emp Fo ma ion in o in o mal uni s wi h a wide egional ank. The co espondence be ween he di e en uni s o each au ho is as ollows (Fig. 2): “g ey uni ” o “G ey Ga umnian” o Rosell e al. (2001) (Posa Fo ma ion acco ding o Cue as, 1992; Uni 1 acco ding o Galb un e al., 1993); “lowe ed uni ” o “Lowe Red Ga umnian” o Rosell e al. (2001) (Conques Fo ma ion and Tala n Fo ma ion acco ding o Cue as, 1992; Uni 2 acco ding o Galb un e al., 1993); “Vallceb e limes ones” and la e al equi alen s o Rosell e al. (2001) (Su e anya Fo ma ion and S . Sal ado de Toló Fo ma ion acco ding o Cue as, 1992; Uni 3 acco ding o Galb un e al., 1993); “uppe ed uni ” o “Uppe Red Ga umnian” o Rosell e al. (2001) (Espluga eda MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT On he o he hand, he co ela ion o he “lowe ed uni ” is based on i e cha ac e is ic packages o sands ones, wo o which can be ecognized in he whole a ea and pe mi he co ela ion be ween he La a and Ba anco Se aduy p o iles (Fig. 3A, B). Th oughou he whole a ea, he i s co ela ion le el (L1 in Figs. 3 and 4) p esen s nume ous dinosau ichni es a he base and is always associa ed wi h a pu ple-g eyish muds one (Fig. 3A). In he wes e n a ea, L1 comp ises a abula body o b ownish ine- o medium-g ained sands ones wi h pa allel lamina ion ha g ade upwa ds o massi e sands ones wi h bio u ba ion on he op. The unde lying abula body o g ey-pu ple muds ones exhibi s oo bio u ba ion, ege al emains and ossil e eb a es. Towa ds he sou heas , L1 passes la e ally in o a hick body o g eyish medium- o coa se-g ained sands ones, which is composed o abula and channeled s a a. These beds a e in e inge ed wi h pu ple bio u ba ed muds ones and show had osau id dinosau acks a he base (Fig. 3A). The second le el (L2 in Figs. 3 and 4) comp ises ine- o medium-g ained massi e sands ones wi h in ense pedogeniza ion and bio u ba ion in he wes e n zone; L2 shows a nega i e-upwa ds end. L2 co esponds o g eyish sands ones ha a e ine- o e y coa se- g ained wi h ough c oss-bedding, c oss-lamina ion and ipples. This le el displays sha p a ia ions in hickness be ween he La a and Isábena Ri e ou c ops. Towa ds he eas , in he Ba anco Se aduy ou c op, he a ia ion in hickness con inues o inc ease, and L2 o ms a g oup o s a a (~7 m hick) wi h a coa sening-upwa ds end loca ed 5 m abo e L1. The p esence o had osau id ichni es is also common a he base. Le el 3 is a cm- hick bed o b own ine-g ained massi e sands ones wi h sca ce la e al con inui y. L3 is loca ed nea o he Isábena Ri e ou c op (Fig. 3B) and is no obse ed in o he a eas. MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT The ou h co ela ion le el, which is up o one me e hick, is cons i u ed by b own medium-g ained sands ones wi h pa allel lamina ion ha passes e ically in o c oss-lamina ion. L4 p esen s a cha ac e is ic small slump (~20 cm hick) a he base and massi e bio u ba ed sands ones (~15 cm hick) in he uppe pa . This le el is easily ecognizable in bo h wes e n and eas e n a eas, om he Isábena Ri e ou c op o nea Ba anco Se aduy (Fig. 3B). The i h le el (L5 in Figs. 3 and 4A) is a ining-upwa d bed composed o 20 cm o g eyish mic oconglome a es wi h sligh channel geome y a he base and 60 cm o ine-g ained massi e sands ones wi h bio u ba ion. In he wes e n a ea, om he La a o Isábena Ri e ou c ops (Figs. 4A, B), his le el p esen s good la e al con inui y (Fig. 3B) and shows spec acula had osau id dinosau ichni es a i s base. On he basis o he e ical a angemen o he guide le els, ch onos a ig aphic e inemen is possible o he s udied sedimen s. Compa ison o he hickness o he “lowe ed uni ” be ween he La a and Ba anco Se aduy p o iles shows clea a ia ions, he succession being hicke in he la e (Fig. 3A). Acco dingly, he a e age sedimen a ion a e o he Ba anco Se aduy succession was sligh ly highe han o he La a sec ion. Consequen ly, he co ela ion esul s also allow us o cons ain he e ical posi ion o paleon ological si es. This new co ela ion e eals ha he mos ecen C e aceous e eb a e emains co espond o dinosau acks and bones in L5 ( he Camino de Rin 2 si e) nea he La a sec ion, which is loca ed ca. 15 m lowe han he Danian “Vallceb e limes ones” (Fig. 3A). Sedimen ological in e p e a ion Cha ac e iza ion o he sedimen a y en i onmen s equi es exhaus i e sedimen ological analysis in o de o es ablish and in e p e co ec ly he di e en acies associa ions no s udied MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT in his wo k. E en so, he Se aduy a ea deposi s may co espond wi h he e olu ion om coas al o con inen al en i onmen s, an in e p e a ion p e iously p oposed by se e al au ho s o he T emp Basin (e.g., Eichensee , 1987; Díaz-Molina, 1987; Cue as, 1992; Rosell e al., 2001; López Ma ínez e al., 2006; Díaz-Molina e al., 2007; Oms e al., 2007, 2016; Rie a e al., 2009; Villalba-B e a e al., 2012; Díez-Canseco e al., 2014; Canudo e al., 2016; Fonde illa e al., 2016a). The sedimen a y ea u es, especially he sedimen a y s uc u es and g ain-size dis ibu ion, indica e ha he A én Fo ma ion in his a ea co esponds wi h a ba ie -island o del aic en i onmen . The p esence o m-scale coa sening-upwa ds sequences, acies associa ions and s acking pa e ns in he s udied in e al a e simila o hose desc ibed by Na a e e e al. (2013) o ba ie -island and washo e an deposi s in e bedded wi hin mud la lagoonal deposi s. The T emp Fo ma ion mainly ep esen s e es ial en i onmen s, bu he p esence o plank onic o amini e s (see below) indica es con inuous en ances o ma ine wa e in o he mo e p o ec ed a eas. Thus, he lowe mos subuni (“g ey uni ”) has been in e p e ed as a ansi ional ma ine- o-con inen al en i onmen connec ing idal sys ems wi h he ba ie island. The “lowe ed uni ” is p edominan ly composed o eddish-b ownish and g eyish-da kish muds ones ep esen a i e o back-ba ie mud la s, whe eas he b ownish abula and e osi e sands ones ep esen lu ial channels and hei o e bank deposi s in he loodplains. Thus, equen wa e - le el oscilla ions and cyclic looding o he mud la a ea can be in e ed om he sedimen a y ea u es. Se e al cha ac e is ics, such as mo ling, oxide haloes and c us s, esul ed om he mig a ion and di e en ial accumula ion o i on, also indica ing common wa e - able oscilla ions. In his con ex , he eddish colo s o he muds ones sugges equen subae ial exposu e, p obably in low wa e -le el e en s. The e ical bio u ba ion and ca bona e p ecipi a ion in he MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT aces indica e he exis ence o ege a ion wi h oo pene a ion in sea ch o he wa e le el du ing d y pe iods. Da kish and g eyish muds ones we e deposi ed unde anoxic condi ions ha a o ed he p ese a ion o o ganic ma e . This acies indica es he occu ence o high wa e - le el pe iods, in which he mud la a eas we e looded. Isola ed and anas omosed channeled sands one bodies wi h eshwa e cha ophy es (see below) indica e he exis ence o low-ene gy, meande ing lu ial channels. Tabula , poo ly so ed sands one bodies and bio u ba ion aces illed wi h sands in muds ones e eal sha p looding e en s ela ed o high-ene gy wa e discha ges. These loods occu ed as a consequence o he o e low in he lu ial channels and he loodplain. The “Vallceb e limes ones” ep esen he es ablishmen o an ex ensi e eshwa e lake. Ma e ial and me hods In o de o ensu e he eplicabili y o his esea ch, all he paleon ological ma e ial igu ed in his s udy, including he e eb a e emains and o amini e s, is p ope ly labeled wi h MPZ abb e ia ions (Museo Paleon ológico de la Uni e sidad de Za agoza) and housed in he Museo de Ciencias Na u ales de la Uni e sidad de Za agoza (Za agoza, Spain). The me hodology applied in his s udy (magne os a ig aphy and bios a ig aphy) is de ailed and explained in he co esponding sec ion. Magne os a ig aphy Paleomagne ic sampling and labo a o y p ocedu es 115 le els we e sampled as pa o he Se aduy magne os a ig aphic s udy, 21 om he A én Fo ma ion (SB) and 94 om he T emp Fo ma ion (SR). Bo h magne os a ig aphic MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT p o iles we e gene a ed in he icini y o he Ba anco Se aduy (BS) s a ig aphic sec ion. SB consis s o 6 le els o blue-g ey ma ls om he Campo Membe , and 14 le els o sands ones wi h one le el o g ey ma ls om he A én Fo ma ion. In he SR p o ile, 17 le els co espond o he “g ey uni ” o ma ls and calca eni es o he T emp Fo ma ion. Ano he 73 le els a e de ined as an al e na ion o ed, g ey and e sicolo muds ones, wi h some le els o sands ones in he so- called “lowe ed uni ”. Finally, 4 le els o he Paleocene “Vallceb e limes ones” we e sampled a he op o he SR sec ion (Fig. 5). The comple e SB p o ile was sampled wi h a po able gas-powe ed and wa e -cooled d ill and di ec ly o ien ed in he ield wi h a magne ic compass and an inclinome e , p o iding om 1 o 3 samples pe le el, each di isible in o 1 o 3 s anda d-sized specimens. In he SR p o ile, 62 le els we e d illed wi h a po able elec ical wa e -cooled d ill, hand samples (blocks) we e aken om 29 le els and in 3 le els bo h d illed and hand samples we e collec ed. These we e o ien ed in si u wi h a magne ic compass. Hand samples we e collec ed because o how easily b oken up (being disagg ega ed) he ines ma e ials co esponding o he “lowe ed uni ” we e. They we e consolida ed wi h sodium silica e dissol ed in dis illed wa e o y o make he consolida o pe cola e o he in e io o each piece. Once ha dened, abou 3 cubes pe block we e sec ioned wi h a disc cu e , main aining he ace pe pendicula o he s ike line and pa allel o he dip line, bo h o ien ed in he ield, as he ma ke o he paleomagne ic analysis. The sampled le els in he SB p o ile we e es ablished each 1 m om he SB01 o SB06 ma ls, e e y 2–4 m a he beginning o he sands ones, and e e y 6 m a e wa d, due o he homogenei y o he ma e ials. The sampled SR p o ile le els we e sepa a ed by 1 m whene e possible. In o al a sequence o 173 m was sampled. MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT The mal (Th) and al e na ing ield (A ) demagne iza ions we e ca ied ou in he paleomagne ic labo a o y o he Uni e si y o Bu gos, using a 755 supe conduc ing magne ome e (2G) wi h an al e na ing ield induc o demagne ize sys em ( o au oma ic A ), a TD48-DC (ASC) o en and a LDA3 (Agico) al e na ing ield demagne ize ( o manual A ). A o al o 198 samples (1 o 3 samples pe le el) we e demagne ized wi h di e en s epwise empe a u es and applied al e na ing ields acco ding o he sample li hology. 145 o hese we e Th-demagne ized, hea ing up o 400–575°C o ma ls, sands ones, calca eni es and limes ones, and up o 475–675°C o muds ones (se e al samples o all li hologies we e hea ed up o 675°C in o de o check hei magne ic beha io s), 39 wi h au oma ic A , 14 wi h manual A ying o imp o e he accu acy o he me hod, and an A p o ocol wi h an ini ial he mal s ep o 130°C o dele e he pa o he signal ca ied by goe hi e (all A up o 100 mT). P incipal componen analysis (PCA) and g ea ci cle (GC) analysis we e pe o med wi h Remaso 3.0 so wa e (Chadima and H ouda, 2006). Vi ual Geomagne ic Poles (VGPs) we e calcula ed, h ough he isola ed paleomagne ic di ec ions conside ed p ima y. In cases whe e o e lapping p e en s he isola ion o s able paleomagne ic componen s, GCs we e calcula ed. Toge he wi h he s a ig aphic column, he VGP la i udes ob ained om PCA paleomagne ic di ec ions a e symbolized wi h a poin , whe eas o he p ima y componen s e i ied by GC a ba occupies he s a us co esponding o no mal o in e se la i ude (-90° o 0° o 0° o 90°) (Fig. 5). In addi ion, ock-magne ic measu emen s we e ca ied ou a he Uni e si y o Bu gos wi h a a iable ield ansla ion balance (VFTB). Powde ed whole- ock specimens om 14 ep esen a i e samples om all li hologies we e submi ed o expe imen s on IRM acquisi ion and back ield cu es, hys e esis loops and s ong ield magne iza ion e sus empe a u e (Ms-T) MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT cu es. Analysis o hese measu emen s was pe o med wi h RockMagAnalyze 1.0 so wa e (Leonha d , 2006). Paleomagne ic beha io The na u al emanen magne iza ion (NRM) beha io was analyzed sepa a ely in acco dance wi h he li hology because o he big a ia ions among li hologies. Sands ones and g een/ iole muds ones in gene al show low NRM in ensi ies (0.1–0.5 mA/m) and a he e ogeneous paleomagne ic beha io , being paleomagne ically uns able (i is no possible o isola e a eliable paleomagne ic componen ). In ma ls and ed beds wo paleomagne ic componen s can be iden i ied on he basis o he unblocking empe a u e anges and he cohe ence o he di ec ional da a. Blue-g ey ma ls These ocks appea below he A én Fo ma ion (Campo Membe ), a he bo om o he T emp Fo ma ion (“g ey uni ”) and in e cala ed wi h con inen al sedimen s o he T emp Fo ma ion (“lowe ed uni ”) (Fig. 5). Ma ls o he Campo Membe (samples SB01-SB06 and SB11) show homogeneous paleomagne ic beha io , wi h NRM in ensi ies be ween 1.1 and 1.78 mA/m, and display wo di e en componen s in he mal (Th) demagne iza ion. A low- empe a u e componen MB (unblocking empe a u es be ween 250/300–450 ºC) wi h a no hwa ds di ec ion and posi i e inclina ions (Fig. 6A) is isola ed in all samples. This componen does no go o he o igin in some samples (Fig. 6B, C), going sys ema ically o he sou he n quad an wi h nega i e inclina ion (Fig. 6B). G ea ci cle analysis (Fig. 6E) allows us o in e his high- empe a u e componen (up o 500 ºC) wi h nega i e inclina ion (componen MA), bu his canno be isola ed MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT because o spu ious componen o ma ion du ing hea ing. Al e na ing ield (A ) demagne iza ion diag ams show only one ecognizable componen (6/15–40/60 mT), which co esponds wi h componen MB (Fig. 6A) wi h a sligh o e lapping wi h MA when his appea s, as can be obse ed in he equal-a ea demagne iza ion diag am (Fig. 6B), bu i is no ewo hy ha he o e lapped componen is almos ending o he o igin, p e en ing ecogni ion o he p esence o wo componen s. The NRM in ensi y is lowe in he ma ls o he T emp Fo ma ion (SR samples), a ound 0.4–0.5 mA/m. A low- empe a u e componen (250–400 ºC) is ecognizable, and wi h some excep ions i goes o he o igin, mainly in he basal ma ls (SR01-SR17); his componen shows he same beha io as he al eady desc ibed MB componen o he SB samples. Howe e , a ew samples (SR-01A, SR26-1) show an o e lapping o componen s ei he (i) in he de ini ion o g ea ci cles (Fig. 6E), indica ing he p esence o a high componen (Fig. 6C), o (ii) by a clus e - end showing a sou h declina ion and nega i e inclina ion (Fig. 6D). The e o e, mos samples o SB and SR ma ls show he low- empe a u e MB componen (be ween 250/300 ºC and 350/400 ºC) wi h no mal pola i y (posi i e inclina ion owa ds he no h). This componen does no go o he o igin because o he p esence o a high- empe a u e componen (up o 350/400 ºC) wi h e e sed pola i y, which canno be isola ed because o he o ma ion o spu ious componen s bu is clea ly e idenced by analyzing he g ea ci cles. Acco ding o he NRM beha io (low coe ci i y and unblocking empe a u es be ween 150–500? ºC), and he magne ic ock p ope ies (Fig. 6F), bo h componen s a e ca ied by magne i e. The momagne ic cu es show a majo g ow h o magne ic mine als (magne i e acco ding o he Cu ie empe a u e in he cooling cu e) in ag eemen wi h ha obse ed in he NRM. MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT Red muds ones Red and o ange muds ones appea h oughou he T emp Fo ma ion, mainly in he uppe sec ion. Mos samples show low–medium NRM in ensi ies anging om 0.1 o 1.8 mA/m, bu some o hem ha e highe in ensi ies a ound 3 mA/m. Low-in ensi y samples (~0.2–0.8 mA/m) usually show a single componen (Fig. 7A) wi h unblocking empe a u es om 350/550 ºC up o 620 ºC; howe e , he end o he componen is obli e a ed because o a spu ious componen gene a ed du ing hea ing. In some samples, gene ally hose o a highe in ensi y (> 1 mA/m), componen A o e laps wi h an in e media e empe a u e componen be ween 350 ºC and 550 ºC (Fig. 7B, C); his o e lapping componen has low inclina ion and does no go o he o igin. In hese samples, componen RA can be obse ed a empe a u es up o 500 ºC (Fig. 7B). Finally, SR53 (Fig. 7D) shows high in ensi y and a single componen wi h posi i e inclina ion owa d he no h, which can be in e p e ed as componen RA acco ding o i s unblocking empe a u es (550–625 ºC). The unblocking empe a u es and high coe ci i y (Fig. 7) poin o hema i e as he ca ie o componen RA. This is in ag eemen wi h he ock magne ism expe imen s (Fig. 7E), which a e cha ac e ized by a high-coe ci i y magne ic phase wi h Cu ie empe a u es o e 600 ºC. Di e ences be ween he cooling and he hea ing in he he momagne ic cu e indica e he g ow h o magne ic mine als (p obably magne i e o maghemi e) du ing hea ing, a empe a u es abo e 600 ºC. In e p e a ion o he paleomagne ic componen s Ca bona ic ocks show he p esence o wo componen s wi h di e en unblocking empe a u es. Componen B, ca ied by magne i e, is cha ac e ized by low o in e media e MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT unblocking empe a u es (300–450 ºC) and does no go o he o igin. Se e al wo ks (e.g. Juá ez e al., 1994; Villalaín e al., 1994; Ose e e al., 2007) e idence he p esence o low o in e media e unblocking empe a u es (below 450–500 ºC) o diagene ic seconda y magne i e, and a high- empe a u e componen (abo e 450 ºC) co esponding wi h p ima y magne i e. The unblocking empe a u es o seconda y mine als a e usually lowe han hose o p ima y ones. This is because o he small size o seconda y mine als, which ange om he supe pa amagne ic o he s able single domain (see Jackson and Swanson-Hysell, 2014). This sugges s a seconda y o igin o componen B ound in he ma ls, bu does no ensu e a p ima y o igin o he high- empe a u e componen since he p esence o wo seconda y magne iza ions is also possible. The paleopole e e ence o he La e C e aceous o Ibe ia om he Lisbon Volcanics (Van de Voo and Zijde eld, 1971) co esponds o an expec ed di ec ion o he sec ion loca ion o D=1.08º and I=47.39º. In spi e o he low dip o he s udied ma e ials, he mean di ec ion o componen B is in be e ag eemen be o e han a e he bedding co ec ion (BBC and ABC espec i ely, Fig. 8B). This ac ag ees wi h a seconda y o igin o his componen . We can hus conside componen B o be a chemical emanen magne iza ion (CRM). This was acqui ed p obably du ing he ea ly diagenesis, bu a e he il ing o he se ies (no e ha he il ing is Maas ichian–ea ly Paleocene [Simó e al., 1985], sligh ly pos da ing he age o he ocks). Componen B pa ially obli e a es he high- empe a u e componen (A), which canno be calcula ed because o he g ow h o magne ic mine als du ing hea ing. Howe e , his is clea ly e idenced by he demagne iza ion g ea ci cles. Compa isons be ween he demagne iza ion g ea ci cles (Fig. 8C) calcula ed in ca bona es ( he NW–SE GC) and in ed beds ( he NE–SW GC) a e cohe en wi h he calcula ed MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT had osau oids in Eu asia (Sellés e al., 2014a). The eco e ed eggshells om Se aduy ha e been iden i ied up un il he lowe -mid pa o he "lowe ed uni " o he T emp Fo ma ion (172-i/04/ paleon ological si e). I hei assignmen o Sphe ooli hus eu opaeus is con i med, hese eggshells would he e o e be e en mo e mode n han hose eco e ed a he Pon d'O i locali y. The opods The p esence o he opods wi hin he Se aduy a ea is no e y abundan , ep esen ing be ween 1% and 4% o he iden i ied dinosau emains. These he opod bones ha e been ound om he “g ey uni ” o he middle pa o he “lowe ed uni ” o he T emp Fo ma ion, hus cons i u ing he younges eliable eco d o non-a ian he opods in he Ibe ian Peninsula and one o he younges eco ds in Eu ope. The mos impo an eco d co esponds o wo isola ed ee h ha belong o wo di e en axa, a medium–la ge o m and a small-sized he opod. The i s specimen (MPZ 2017/804; Fig. 11F) co esponds o a medium–la ge oo h wi h se a ed ca inae, which was eco e ed a he op o he "g ey uni " o he T emp Fo ma ion, e y close o he 172-i/04/e paleon ological si e. This oo h is e y simila o Mo pho ype 1 desc ibed by To ices e al. (2015) in he Spanish si es o Blasi (Huesca, uppe Maas ich ian), Mon ebei (Lleida, uppe Campanian–lowe Maas ich ian) and Laño (Bu gos, uppe Campanian–lowe Maas ich ian). Due o i s limi ed diagnos ic alue, his mo pho ype has been assigned o The opoda inde . (López-Ma ínez e al., 2001; To ices e al., 2004, 2015; Pe eda-Sube biola e al., 2015). The second specimen co esponds o a small oo h wi h smoo h ca inae, which was eco e ed a he La a 4 paleon ological si e wi hin he "lowe ed uni " o he T emp Fo ma ion. This oo h is e y simila o he ee h assigned o Coelu osau ia inde . om Blasi (Huesca, uppe Maas ich ian), Mon ebei (Lleida, uppe Campanian–lowe Maas ich ian), Laño (Bu gos, uppe Campanian–lowe Maas ich ian) and MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT Vica i 4 (Lleida, uppe Campanian) (López-Ma ínez e al., 2001; To ices e al., 2004, 2015; Pe eda-Sube biola e al., 2015). The o he emains, also eco e ed in he “g ey uni ” and he “lowe ed uni ” o he T emp Fo ma ion, co espond o a possible ce ical e eb a o an a ian he opod (C uzado- Caballe o e al., 2012) and agmen a y long bones and a e eb al agmen ha may co espond o unde e mined he opods. Ne e heless, o he p ope axonomic assigna ion o hese emains, u he de ailed s udies will be necessa y. Sau opods Among he dinosau emains, he p esence o sau opods is he sca ces , amoun ing o a ound 1% o he iden i ied emains. The mos impo an i em is a p oximal le emu agmen (MPZ 99/143; Fig. 11E) assigned o Ti anosau ia inde . (Canudo, 2001; Vila e al., 2012). This emu was eco e ed in he "g ey uni " o he T emp Fo ma ion, ep esen ing one o he younges sau opods ye documen ed in Eu asia (Canudo, 2001; Vila e al., 2012; Sellés e al., 2016). O he possible sau opod emains consis o a caudal e eb a om he Ba anco de Se aduy 4 si e (C uzado-Caballe o e al., 2012) and a p oximal agmen om a big au opodial bone om he Camino de Fo nons 1 si e. Howe e , due o he agmen a y na u e o hese bones, hei assignmen o Sau opoda s ill emains doub ul. Fo he p ope assigna ion o hese specimens, u he s udies as well as he eco e y o new emains will be necessa y. Bo h hese emains appea ed in he middle pa o he "lowe ed uni ", so i hei assignmen o Sau opoda is con i med, hey would be mo e mode n han he emu , ex ending he p esence o sau opods o ch on C29 , as al eady seen in o he sec o s o he T emp Basin (Sellés e al., 2016). Tes udines MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT Ano he common clade, comp ising abou 5% o he emains ound in Se aduy, is Tes udines. The p esence o es udines has been ecognized om he op o he A én Fo ma ion o he las deposi s wi h e eb a es o he “lowe ed uni ” o he T emp Fo ma ion. This clade is ep esen ed en i ely by isola ed and disa icula ed pla es, in mos cases p e en ing a mo e accu a e iden i ica ion o classi ica ion in he p elimina y s udy, and ende ing u he sys ema ic s udies necessa y. Ne e heless, when he pla es a e well p ese ed, i is possible o obse e hei smoo h and b illian o namen a ion c ossed by e y ine dicho omic sulci, sugges ing highly ascula ized shell bones. This cha ac e is ic o namen a ion pa e n is widely used o ecognize bo h emydids (e.g., de Lappa en de B oin and Mu elaga, 1996; Mu elaga and Canudo, 2005; Ma mi e al., 2012), so mos o hese pla es a e assigned o Bo h emydidae inde . In addi ion, be e -p ese ed pla es allowed mo e accu a e ana omical iden i ica ion, and a le xiphiplas on and a igh mesoplas on belonging o Bo h emydidae om he Rim 2 si e a he op o he A én Fo ma ion ha e been ecognized (Mu elaga and Canudo, 2005). C ocodylomo phs Rep esen ing abou 4% o he eco e ed bone emains, c ocodylomo phs a e one o he mos ep esen a i e axa in Se aduy. This g oup o a chosau s is mainly ep esen ed by isola ed ee h, al hough some ichni es, a eusuchian e eb a, os eode m agmen s and a comple e skull ha e also been ound (Fig. 13E). All he eco e ed emains ha e been assigned o Eusuchia. These ha e a highly ex ended s a ig aphic dis ibu ion, emains being ound om he op o he A én Fo ma ion up un il he las le els wi h e eb a es be o e he K/Pg bounda y. The mos impo an axon co esponds o he comple e skull o he eusuchian c ocodylomo ph o iginally e ec ed in Se aduy wi h he name o Allodaposuchus subjunipe us MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT (Pué olas-Pascual e al., 2014) (MPZ 2012/288; Fig. 13E). This axon was la e assigned o he new genus Aga esuchus and included wi hin Allodaposuchidae (Na áez e al., 2016), a clade o endemic Eu opean eusuchian c ocodylomo phs wi h a eco d un il he K/Pg bounda y. Aga esuchus subjunipe us was eco e ed in one o he las Maas ich ian sands one s a a (Amo 3 si e), so his axon may ep esen he las and younges eco d o Allodaposuchidae be o e he K/Pg ex inc ion e en (Pué olas-Pascual e al., 2014). As ega ds isola ed ee h, a leas wo di e en mo pho ypes ha e been dis inguished. The i s mo pho ype (Fig. 13A) co esponds o a slende conical oo h o namen ed wi h well- ma ked longi udinal idges, which was ound a he op o he A én Fo ma ion (Ba anco de Ex emadu a si e). These oo h ha e been assigned o c . Tho acosau us (Pué olas-Pascual e al., 2016). This ma ine genus belonging o Ga ialoidea is ypical o he Uppe C e aceous–lowe Paleocene o Eu ope and No h Ame ica, which is consis en wi h i s p esence wi hin he shallow ma ine acies o he A én Fo ma ion. Tee h o he second mo pho ype (Fig. 13B, C) ha e been eco e ed om he “g ey uni ” up un il he las le els wi h e eb a es wi hin he “lowe ed uni ” o he T emp Fo ma ion. This mo pho ype co esponds o gene alis conical ee h wi h an o namen a ion ha a ies om smoo h o gen ly longi udinally- idged enamel. This gene alis mo phology is widely dis ibu ed wi hin C ocodylomo pha and has li le axonomic alue (e.g., P asad and B oin, 2002; Tu ne , 2006; And ade and Be ini, 2008; Buscalioni e al., 2008). Howe e , as his den al mo phology is also ypical o Allodaposuchidae, he mos common clade in Eu ope du ing he Campanian–Maas ich ian, his mo pho ype has been en a i ely assigned o c . Allodaposuchidae. Addi ionally, se e al isola ed ee h simila o hose p esen in A. subjunipe us we e eco e ed in he same si e whe e he holo ype was eco e ed, so hese ee h ha e been assigned o c . Aga esuchus subjunipe us. MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT O he bone emains co espond o os eode m agmen s and a p ocoelous do sal e eb a (Fig. 13D) ha ha e been assigned o Eusuchia inde . Fu he e idence o C ocodylomo pha is he p esence o abou i e acks (MPZ 2012/832) composed o sc a ch ma ks and one pedal imp ession loca ed on a lu ial channel deposi in he uppe mos pa o he “lowe ed uni ” o he T emp Fo ma ion (Se aduy No e si e, ch on C29 ). The sc a ch ma ks esemble Cha acichnos whe eas he pes ack has been assigned o c . C ocodylopodus (Vila e al., 2015). Amphibians Due o hei small size, amphibian emains ha e only been eco e ed by washing and sie ing echniques. In he a ea o Se aduy, one o he paleon ological si es wi h he g ea es po en ial o he s udy o mac o e eb a es and mic o e eb a es is La a 4, loca ed wi hin he "lowe ed uni " o he T emp Fo ma ion. This si e is loca ed in a da k g ey lu i e laye wi h a high o ganic con en , whe e ege al emains (wood and ambe agmen s), mac o e eb a es (dinosau s, c ocodylomo phs and es udines) and mic o e eb a es a e highly abundan . Mos o he mic o e eb a e emains a e e y agmen a y and need a mo e ho ough sys ema ic s udy o hei p ope iden i ica ion. Ne e heless, a e y p elimina y s udy o he mic opaleon ological con en allowed us o iden i y se e al emains ha may co espond wi h amphibians. The mos ou s anding emains a e se e al dis al pa s o hume i. In spi e o he low axonomic alue o he hume us (E ans and Milne , 1993), he la ge and sphe ical hume al ball shi ed la e ally and a a he long ulna epicondyle allow us en a i ely o assign hese specimens o Discoglossidae inde ., being e y MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT simila o o he hume i assigned o his clade in o he si es wi hin he T emp Basin (Blain e al., 2010; Blanco e al., 2016). Discussion Conside ing componen A as p ima y (see “Magne os a ig aphy” sec ion), he local magne ic s a ig aphy o he Se aduy sec ion can be co ela ed wi h he geomagne ic pola i y ime scale (GPTS) (G ads ein e al., 2012). Acco ding o ou esul s, he “g ey uni ” and he “lowe ed uni ” o he T emp Fo ma ion can be assigned o a e e se pola i y ch on (Fig. 5). The p esence o he plank onic o amini e P. ha iaensis in sample SR35 indica es ha his le el has a maximum age o 67.3 Ma (maximum age ange o he species), o younge i i is ewo ked. The age ange o his species is be ween 67.3 and 66.0 Ma, i s lowe mos and uppe mos occu ences being coinciden espec i ely wi h he uppe pa o C30n and he K/Pg bounda y in he middle pa o C29 . Acco ding o López-Ma ínez e al. (2006) and Díez- Canseco e al. (2014), he “Vallceb e limes ones uni ” and la e al equi alen s a e la e Danian in age. Because his bios a ig aphic in o ma ion indica es ha he K/Pg bounda y is loca ed be ween he “lowe ed uni ” and he “Vallceb e limes ones uni ”, all he e e se pola i y sec ion be ween le els SR35 and SR90 can hus only co espond wi h ch on C29 (Fig. 5). As a esul , he K/Pg bounda y can be loca ed wi hin he las 25 m o he “lowe ed uni ”, be ween he las ho izon wi h dinosau emains and he “Vallceb e limes one uni ” (Fig. 5). Co ela ing he p o ile o Ba anco Se aduy wi h La a, loca ed u he wes , he K/Pg bounda y can be loca ed wi h mo e p ecision wi hin he las 5 m o he “lowe ed uni ” (Figs. 3, 5). The e o e, he uppe sec ion o he “lowe ed uni ” is well de ined as e e se pola i y, poin ing o he C29 (Fig. 5). Howe e , some inconsis encies can be obse ed in he middle pa MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT o his o ma ion. SR37, abo e he le el ma ked by P. ha iaensis, has e e sed pola i y (indica ing C29 ), whe eas le el SR53 shows no mal pola i y. The e o e, one pa o his paleomagne ic in o ma ion mus be w ong. As ega ds i s paleomagne ic p ope ies, le el SR35 has simila beha io o samples om he uppe mos le els, wi h a non-ze o ending clus e up o 500 ºC wi h e e sed pola i y. O he wise, componen RA o le el SR53 (Fig. 7D) has been de ined by i s unblocking empe a u e ange; howe e , i is possible ha his componen co esponds wi h he CRM de ined as RB. The li hology o bo h samples can also be analyzed: he paleomagne ic componen de ined in SR35 is mo e eliable han ha de ined in SR53 because he o me is sampled in ed muds ones, simila o he uppe sec ion whe e cohe en paleomagne ic componen s appea , whe eas SR53 is in a le el o shale loca ed be ween sands ones. The g ea e po osi y o sands ones could ha e a o ed chemical p ocesses in le el SR53. In he ligh o hese conside a ions, we conside ha he sec ion be ween le els SR37 and SR90 belongs o subch on C29 . The lowe sec ion o he T emp Fo ma ion also shows e e sed pola i ies (see “Magne os a ig aphy” sec ion), so a he beginning i is possible o asc ibe his o subch on C29 (Fig. 14). Howe e , Fonde illa e al. (2016a) p o ide e idence o he p esence o a majo hia us a ec ing ch ons C31n, C30 and C30n in he Isona sec ion loca ed in he eas e n sec o o he T emp syncline (Fig. 14), p obably ela ed o an ab up mig a ion o he basin depocen e . This would imply he p esence o consecu i e deposi s associa ed wi h ch ons C31 and C29 , wi h a hia us lacking mos o he uppe Maas ich ian. Ne e heless, acco ding o he magne os a ig aphic wo ks o Pe eda-Sube biola e al. (2009) on A én, and Canudo e al. (2016) on Campo, his majo hia us seems no o ha e a ec ed he mos wes e n sec o s o he T emp Basin (Fig. 14). The e o e, he sec ions o Campo and A én acqui e g ea e ele ance because MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT he “lowe ed uni ” exposed he e is he only con inen al eco d o ch on C30n in he whole T emp syncline (Fonde illa e al., 2016a). In summa y, he magne os a ig aphic da a p esen ed in his wo k show ha bo h he ma ls and ed beds o he T emp Fo ma ion in he Se aduy a ea a e o e e se pola i y. Bios a ig aphic da a ensu e ha he uppe pa o his o ma ion ( he las 60 m o he “lowe ed uni ”) belong o ch on C29 . Howe e , al hough his canno be ully con i med o he lowe pa o he T emp Fo ma ion ( he “g ey uni ” and beginning o he “lowe ed uni ”), hese uni s p obably also belong o he same ch on C29 , unless he e exis s a hia us such as ha obse ed by Fonde illa e al. (2016a) in he Isona sec ion. I should be poin ed ou ha his hia us has no been obse ed in Campo and A én (Pe eda-Sube biola e al., 2009; Canudo e al., 2016), he sec ions closes o he Se aduy a ea. Conclusions In his wo k, a ch onos a ig aphic amewo k o he e eb a e si es o he A én and T emp Fo ma ions wi hin he Se aduy sec o o he T emp Basin is p oposed o he i s ime. The join s udy o s a ig aphy, ield co ela ions, magne os a ig aphy and bios a ig aphy has allowed mos o he e eb a e si es in his a ea o be da ed o wi hin ch on C29 , making his one o he a eas wi h dinosau si es closes o he K/Pg bounda y anywhe e in Eu ope. In addi ion, a comple e aunal lis o he axa eco e ed in he Se aduy a ea is p esen ed. This shows a g ea di e si y o he opods, sau opods and had osau dinosau s, eusuchian c ocodylomo phs, es udines, amphibians and p obably p e osau s. The p esence o dinosau s (ichni es and bones) in he highes le els o he se ies has pinpoin ed he ange o he K/Pg bounda y o he las 5 m o he “lowe ed uni ” wi hin he MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT T emp Fo ma ion, below he “Vallceb e limes ones”. This sugges s a high abundance o had osau id dinosau s, eusuchian c ocodylomo phs, amphibians and es udines jus be o e he g ea ex inc ion e en o he La e C e aceous. Al hough e e y hing poin s o a la e Maas ich ian age o he s udied deposi s, he lowe hal o he sec ion un o una ely shows unclea paleomagne ic signals and inconclusi e bios a ig aphic con en , so i has been assigned an unde e mined pola i y. The magne os a ig aphic esul s also seem o indica e he p esence o a e e se pola i y ch on in he lowe hal o he sec ion, ye we do no ha e he bios a ig aphic da a o be able o assign i o a speci ic ch on (C31 , C30 o C29 ). Fo his eason, he con inui y o he lowe pa o he se ies o he p esence o possible hia uses canno be de e mined. Fu he s udies in he adjacen ou c ops loca ed be ween Se aduy and Campo (e.g. Rin o La a sec ions) and be ween Se aduy and A én (e.g. Iscles sec ion) could be c ucial o achie e mo e accu a e knowledge o he ch onos a ig aphic amewo k o he no hwes e n-mos b anch o he T emp Basin. In conclusion, hese esul s show he g ea paleon ological po en ial o he Se aduy a ea, which is one o he ew and mos impo an places in he wo ld o s udying, wi hin con inen al deposi s, he g ea ex inc ion e en which a ec ed plane Ea h a he end o he C e aceous. Acknowledgemen s This wo k was suppo ed by he Spanish Minis y o Economy and Compe i i eness (g an numbe s CGL2014-53548-P, CGL2015-64422-P and CGL2017-85038-P), co inanced by he Eu opean Regional De elopmen Fund; and by he Depa men o Educa ion and Science o he A agonese Go e nmen (g an numbe s DGA g oups H54 and E05), co inanced by he MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT Eu opean Social Fund (ESF). The paleomagne ic s udy was possible hanks o he complemen a y g an s (bene icia ies o FPU, g an numbe CGL2010-16447/BTE: B ie S ays and Tempo a y T ans e s, yea 2015) suppo ed by he Spanish Minis y o Cul u e, Educa ion and Spo s; and he Labo a o y o paleomagne ism o he Uni e si y o Bu gos (Spain). Edua do Pué olas Pascual is he ecipien o a pos doc o al g an (SFRH/BPD/116759/2016) unded by he Fundação pa a a Ciência e Tecnologia (FCT-MCTES). Special hanks o R. Sil a, V. Bu iel, M. Gallego, A. Alonso, C. Nuñez, J. Galán, J. Pa illa, J.C. Ga cía and X. 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MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT V emi , M., Bălc, R., Csiki-Sa a, Z., B usa e, S.L., Dyke, G., Naish, D., No ell, M.A., 2014. Pe eş i-A ini – An impo an bu epheme al Uppe C e aceous con inen al e eb a e si e in he sou hwes e n T ansyl anian Basin, Romania. C e aceous Resea ch 49, 13–38. doi:10.1016/j.c e es.2014.02.002 Figu e cap ions Figu e 1. Geog aphical and geological si ua ion o he Se aduy sec o (Huesca, Spain). A. loca ion o he Py enees wi hin he Ibe ian Peninsula; B. geological map o he Se aduy sec o ; he ed s a s indica e he a eas wi h he highes concen a ion o e eb a e si es; he ed ec angles indica e he magne os a ig aphic sec ions (SB, SR) s udied in his wo k; C. composi e s a ig aphic sec ion o he s udied p o iles and s a ig aphic e eb a e dis ibu ion. Figu e 2. Li hos a ig aphic subdi ision o he T emp Fo ma ion acco ding o di e en au ho s. Modi ied om Cue as (1992) and Rie a (2010). Figu e 3. A. co ela ion panel o he s a ig aphic p o iles and e eb a e si es s udied in his wo k; B. de ailed geological map o he Se aduy sec o and guide le el loca ions. Figu e 4. Landscape iews o he main a eas wi h ou c ops and paleon ological e eb a e si es in he Se aduy sec o . A. ou c op o he La a s a ig aphic sec ion loca ed on he wes side o he Isábena Ri e ; B. succession o he A én and T emp Fo ma ion in he sec o loca ed wes o he Isábena Ri e ; C. succession o he A én and T emp Fo ma ion in he Ba anco Se aduy sec o loca ed eas o he Isábena Ri e . Red s a s poin o he main a eas wi h e eb a e si es; he MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT whi e do ed lines ma k he con ac be ween di e en uni s; and he colo ed do ed lines poin o he loca ion o he di e en guide le els. Figu e 5. Li hology, paleon ological si e posi ions, and he p oposed magne os a ig aphy om he Ba anco Se aduy sec ion. VGP la i ude logs along he A én Fo ma ion (SB) and T emp Fo ma ion (SR) p o iles a e shown by ci cles (black o whi e) when he pola i y has been calcula ed om paleomagne ic di ec ions ob ained by p incipal componen analysis (PCA) o by ba s (whi e) when i has been ob ained by g ea ci cle analysis (GCA). Figu e 6. A o D. demagne iza ion diag ams, in geog aphic coo dina es, showing he paleomagne ic beha io s in ep esen a i e samples o ca bona ic ocks; A o B. samples om he SB and SR sec ions showing an o e lap be ween wo componen s wi h opposi e di ec ion; D. sample o he SR sec ion wi h a no mal pola i y componen a low empe a u es and a dispe sed high empe a u e clus e ; E. equal a ea p ojec ion o he MB componen and he demagne iza ion g ea ci cles calcula ed in hese ocks, be o e and a e bedding co ec ion (BBC and ABC espec i ely). No e he pa h om no mal pola i y o e e se pola i y ollowed by all samples; F. ep esen a i e ock magne ic expe imen s o his li hology. F om le o igh , acquisi ion o he iso he mal emanen magne iza ion (IRM), non-co ec ed and co ec ed hys e esis loop, and he momagne ic cu e. The low coe ci i y may indica e he p esence o magne i e as he main magne ic mine al, as can be obse ed in he IRM and hys e esis loops; he he momagne ic cu e shows an impo an g ow h o magne i e up o 400ºC. MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT Figu e 7. A o D. demagne iza ion diag ams showing, in geog aphic coo dina es, he paleomagne ic beha io s in ep esen a i e samples o ed beds; A. componen s RA and RB wi h he same pola i y; B and C. componen RB does no go o he o igin and a non-ze o end clus e e eals componen RA wi h e e sed pola i y; in SR78-2B, RA is pa ially demagne ized be o e he g ow h o he magne ic mine al du ing hea ing (up o 620ºC); D. RA and RB wi h no mal pola i y; E. ep esen a i e ock magne ic expe imen s o his li hology. F om le o igh , IRM, hys e esis loop and he momagne ic cu e. The high coe ci i y obse ed in he IRM and in he hys e esis loop and he p esence o a magne ic phase wi h Cu ie empe a u es abo e 620ºC indica e he p esence o hema i e as he main magne ic phase. The highe magne iza ion o he cooling cu e indica es he g ow h o magne i e du ing hea ing. Figu e 8. Equal-a ea p ojec ion o componen A (A) and componen B (B) wi h hei espec i e Fishe means (Fishe , 1953), be o e and a e bedding co ec ion (BBC and ABC espec i ely); C. calcula ed demagne iza ion g ea ci cles (GC) and mean di ec ion o bo h componen s; no e ha bo h componen s o e lap wi h he GC mean in e sec ion, componen A being almos coinciden wi h i s mean. n: numbe o samples. NW-SW GCs co espond wi h ca bona es and NE-SW GCs wi h ed beds. Figu e 9. Some ep esen a i e plank onic and ben hic o amini e species iden i ied in he Ba anco Se aduy sec ion. F om le o igh and om op o bo om: Lae ihe e ohelix glab ans (MPZ 2018/25), He e ohelix globulosa (MPZ 2018/23), Guembeli ia blowi (MPZ 2018/28), Hedbe gella land ini (MPZ 2018/24), Ven ilab ella egge i (MPZ 2018/22), Pseudoguembelina ha iaensis (MPZ 2018/21), Globige inelloides p ae olu us (MPZ 2018/27), Globo uncana MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT bulloides (MPZ 2018/26), Con uso uncana o nica a (MPZ 2018/20), Fa usella washi ensis (MPZ 2018/18), Gy oidinoides sp. (MPZ 2018/19). Figu e 10. Ve eb ae o Had osau idae inde . om Se aduy. A. caudal (ju enile MPZ 2017/796 om AM2 si e); B. caudal (adul MPZ 2017/797 om LAR3 si e); C. caudal (ju enile MPZ 2017/798 om LAR3 si e); D. ce ical (adul ? MPZ 2017/799 om LAR3 si e). In pos e io /an e io , do sal, la e al and en al iews espec i ely. Scale ba = 3cm. Figu e 11. Dinosau emains om Se aduy. A. co acoid o Had osau idae inde . (MPZ 2017/800 om LAR2 si e) in medial and la e al iews; B. phalanx o Had osau idae inde . (MPZ 2017/801 om LAR3 si e) in do sal, la e al and en al iews; C. ibia (dis al agmen ) o Had osau idae inde . (MPZ 2017/802 om LAR3 si e) in an e io , pos e io and dis al iews; D. emu (p oximal agmen ) o Had osau idae inde . (MPZ 2017/803 om LAR3 si e) in an e io , la e al, pos e io , medial and p oximal iews; E. emu (p oximal agmen ) o Ti anosau ia inde . (MPZ 99/143 om emu si e) in pos e io iew; F. ee h o The opoda inde . (MPZ 2017/804 om 172-i/04/e si e) in lingual and labial iew (whi e boxes show he de ailed den icles). Scale ba wi hou numbe = 3cm. Figu e 12. Dinosau ichni es o Had osau opodus inde . om Se aduy (Ped egal si e). Figu e 13. C ocodylomo ph (Eusuchia) emains om Se aduy. A. ee h o c . Tho acosau us (MPZ 2017/806 om EXT si e); B. ee h o c . Allodaposuchidae (MPZ 2017/807 om DL3 si e); C. ee h o c . Allodaposuchidae (MPZ 2017/808 om PED si e); D. do sal e eb a o MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT Eusuchia inde . (MPZ 2017/805 om LAR3B si e); E. skull holo ype o Aga esuchus subjunipe us (MPZ 2012/288) ( om AM3 si e). Figu e 14. Ch onos a ig aphic amewo k wi h indica ion o li hos a ig aphy, paleomagne ic da a, co ela ion and he K/Pg ansi ion in he main Maas ich ian Sou h Py enean con inen al sec ions wi h magne os a ig aphic da a. Table 1. Ve eb a e aunal lis o Se aduy (Huesca, Spain), uppe Maas ich ian. MANUS CRIP T ACCEP TED ACCEPTED MANUSCRIPT Si e Abb e ia ion Taxa Si e Abb e ia ion Taxa 172-i/04/a 172-i/04/a Had osau idae inde . Camino Rin 2 CRIN2 Had osau idae? inde . The opoda? inde . P e osau ia? inde . 172-i/04/b 172-i/04/b Dinosau ia inde . Dolo 1 DL1 Had osau idae? inde . 172-i/04/c 172-i/04/c Dinosau ia inde . Dolo 2 DL2 Dinosau ia inde . A ialae? inde . Bo h emydidae inde . 172-i/04/d 172-i/04/d Dinosau ia inde . Dolo 3 DL3 Had osau idae inde . A ialae? inde . Bo h emydidae inde . c . Allodaposuchidae 172-i/04/e 172-i/04/e Had osau idae inde . The opoda inde . Ba anco Ex emadu a EXT Dinosau ia inde . Had osau idae inde . Bo h emydidae inde . c . Tho acosau us 172-i/04/ 172-i/04/ Had osau idae inde . Fo nons 1 F1 Dinosau ia inde . Had osau idae inde . Amo 1 AM1 Dinosau ia inde . Had osau idae inde . Bo h emydidae inde . Fo nons 2 F2 Dinosau ia inde . Had osau idae? inde . Amo 2 AM2 Had osau idae inde . Bo h emydidae inde . Fo nons 3 F3 Dinosau ia inde . The opoda? inde . Amo 3 AM3 Dinosau ia inde . Had osau idae? inde . Bo h emydidae inde . Aga esuchus subjunipe us La a 1 LAR1 Ve eb a a inde . Eusuchia inde . Ba anco Se aduy 1 BS1 Dinosau ia inde . Had osau idae inde . La a 2 LAR2 Dinosau ia inde . O ni hopoda inde . Had osau idae inde . Bo h emydidae inde . c . Allodaposuchidae Ba anco Se aduy 2 BS2 Dinosau ia inde . Had osau idae inde . La a 3 LAR3 Dinosau ia inde . Had osau idae inde . Eusuchia inde . Ba anco Se aduy 3 BS3 Ve eb a a inde . La a 4 LAR4 Had osau idae inde . Coelu osau ia inde . Bo h emydidae inde . c . Allodaposuchidae Discoglossidae inde . Ba anco Se aduy 4 BS4 Dinosau ia inde . Had osau idae inde . Sau opoda inde . Bo h emydidae inde . Eusuchia inde . La a 5 LAR5 Had osau idae inde . Ba anco Se aduy 5 BS5 Had osau idae inde . Bo h emydidae inde . La a 6 LAR6 Dinosau ia inde . Bo h emydidae inde . Camino Fo nons 1 CF1 Dinosau ia inde . Had osau idae inde . The opoda? inde . Sau opoda? inde . Os eich hyes inde . Bo h emydidae inde . Ped egal PED c . Allodaposuchidae Camino Fo nons 2 CF2 Had osau idae inde . c . Allodaposuchidae Rin 1 y 2 RIN1-2 Dinosau ia inde . Colo COL Dinosau ia inde . Had osau idae inde . Bo h emydidae inde . San C is obal SCRI Dinosau ia inde . Had osau idae inde . Camino Rin 1 CRIN1 The opoda? inde . Sie a de Sis 1 SIS1 Dinosau ia inde . Had osau idae inde . Bo h emydidae inde . Sie a de Sis 2 SIS2 Dinosau ia inde . Had osau idae inde . 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