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. Pe eda o hei help in
he ield wo k; and J. La añaga and G. Ma ín o he disco e y o he La a paleon ological
si es. Many hanks o P. C uzado o he help ul commen s. Rupe Glasgow edi ed he ex in
English.
Re e ences
Ál a ez-Sie a, M.A., A ibas, M.E., A dé ol, L., Ci is, J., Daams, R., K auss, S.,
López-Ma ínez, N., De La Peña, A., Sole , R., Vianey-Liaud, M., 1994. El lími e C e ácico-
Te cia io en la sección de Fon llonga (cuenca de Age , p o incia de Lé ida), in: Jaca (Spain), II
Cong eso Del G upo Español Del Te cia io, Comunicaciones, 23–26.
And ade, M.B., Be ini, R.J., 2008. Mo phology o he den al ca inae in Ma iliasuchus
ama ali (C ocodylomo pha, No osuchia) and he pa e n o oo h se a ion among basal
Mesoeuc ocodylia. A qui os do Museu Nacional ( io de Janei o) 63–82.
An onescu, E., Lupu, D., Lupu, M., 1983. Co ela ion palinologique du C é acé e minal
du sud-es des Mon s Me ali e i e des Dep essions de Haţeg e de Rusca Mon ană. Anua ul
Ins i u ului de Geologie şi Geo izică 59, 71–77.
MANUS CRIP T
ACCEP TED
ACCEPTED MANUSCRIPT
Csiki-Sa a, Z., Bu e au , E., Ősi, A., Pe eda-Sube biola, X., B usa e, S.L., 2015.
Island li e in he C e aceous- aunal composi ion, biogeog aphy, e olu ion, and ex inc ion o
land-li ing e eb a es on he La e C e aceous Eu opean a chipelago. ZooKeys 1.
doi:h ps://doi.o g/10.3897/zookeys.469.8439
Cue as, J.L., 1992. Es a ig a ía del “ga umniense” de la Conca de T emp: P epi ineo
de Lé ida. Ac a geológica hispánica 27, 95–108.
Cushman, J.A., 1926. The o amini e a o he Velasco shale o he Tampico
embaymen . Bulle in o he Ame ican Associa ion o Pe oleum Geologis s 10(6), 581–612.
Cushman, J.A., 1928. Fo amini e es du S ampien du Bassin de Pa is. Bulle in de la
Socie e des Sciences de Seine-e -Oise 9, 47–63.
Cushman, J.A., 1933. Some new o amini e al gene a. Con ibu ions om he Cushman
Labo a o y o Fo amini e al Resea ch 9(2), 32–38.
Cushman, J.A., 1938. C e aceous species o Gümbelina and ela ed gene a.
Con ibu ions om he Cushman Labo a o y o Fo amini e al Resea ch 14(1), 2–28.
Dalbiez, F., 1955. The genus Globo uncana in Tunisia. Mic opaleon ology 1(2), 161–
171.
de Lappa en de B oin, F., Mu elaga, X., 1996. Une nou elle aune de chéloniens dans
le C é acé supé ieu eu opéen. Comp es endus de l’Académie des sciences. Sé ie 2. Sciences de
la e e e des planè es 323, 729–735.

MANUS CRIP T
ACCEP TED
ACCEPTED MANUSCRIPT
Díaz-Molina, M., 1987. Sedimen ación sin ec ónica asociada a una subida ela i a del
ni el del ma du an e el C e ácico supe io (Fm. T emp, p o incia de Lé ida). Es udios
Geológicos 43, 69–93.
Díaz-Molina, M., Kälin, O., Beni o-Mo eno, M.I., López-Ma ínez, N., Vicens, E.,
2007. Deposi ional se ing and ea ly diagenesis o he dinosau eggshell-bea ing A en Fm a
Bas us, La e Campanian, sou h-cen al Py enees. Sedimen a y geology 199, 205–221.
Díez-Canseco, D., A z, J.A., Beni o, M.I., Díaz-Molina, M., A enillas, I., 2014. Tidal
in luence in edbeds: A palaeoen i onmen al and bioch onos a ig aphic econs uc ion o he
Lowe T emp Fo ma ion (Sou h-Cen al Py enees, Spain) a ound he C e aceous/Paleogene
bounda y. Sedimen a y Geology 312, 31–49. doi:h ps://doi.o g/10.1016/j.sedgeo.2014.06.008
Dina es-Tu ell, J., Fonde illa, V., Oms, O., Le Loeu , J., Vila, B., Es ada, R., Rie a,
V., 2014. Magne os a ig aphic da ing o he Maas ich ian e es ial successions om he
Py enees: S a e o he a and new da a om Aude (F ance). J. Ma mi, O. Oms, B. Vila,A.
Galoba , R. Es ada, & J. Dina es-Tu ell (Eds.), Paleon ologia i E olució, mem. especial 68.
Donze, P., Po haul , B., Thomel, G., Villou eys, O. de, 1970. Le Sénonien in é ieu de
Puge -Thénie s (Alpes-Ma i imes) e sa mic o aune. Geobios 3, 81–82.
d'O bigny, A., 1839. Fo amini e es. In: de la Sag a, R. (Ed.), His oi e physique e
na u elle de l'Ile de Cuba. A. Be and, Pa is, F ance, 224.
Eh enbe g, C.G., 1839. Ube die Bildung de K eide elsen und des K eideme gels du ch
unsich ba e O ganismen. physikalische abhandlungen de Koniglichen akademie de
wissenscha en zu Be lin, 59–147.
MANUS CRIP T
ACCEP TED
ACCEPTED MANUSCRIPT
Eichensee , H., 1987. Facies geology o la e Maas ich ian o Ea ly Eocene coas al and
shallow ma ine sedimen s, emp-g aus basin, no heas e n spain. A b. Mus. Geol. Pal. Uni .
Tübingen 237.
Eichensee , H., Lu e bache , H., 1992. The ma ine paleogene o he emp egion (NE
Spain)-deposi ional sequences, acies his o y, bios a ig aphy and con olling ac o s. Facies 27,
119–151. doi:10.1007/BF02536808
El Nagga , Z.R., 1966. S a ig aphy and Plank onic Fo amini e a o he Uppe
C e aceous-Lowe Te ia y: Bulle in o he B i ish Museum (Na u al His o y). Geology,
Supplemen , 2, 291.
E ans, S.E., Milne , A.R., 1993. F ogs and Salamande s om he Uppe Ju assic
Mo ison Fo ma ion (Qua y Nine, Como Blu ) o No h Ame ica. Jou nal o Ve eb a e
Paleon ology 13, 24–30.
Fas o sky, D.E., Sheehan, P.M., 2005. The ex inc ion o he dinosau s in No h
Ame ica. Gsa Today 15, 4–10.
Feis , M., Colombo, F., 1983. La limi e C e ace-Te iai e dans Ie no d-es de l’Espagne,
du poin de ue des cha ophy es. Ceol. medil 10, 303–326.
Fonde illa, V., Dina ès-Tu ell, J., Oms, O., 2016a. The ch onos a ig aphic amewo k
o he Sou h-Py enean Maas ich ian succession eapp aised: Implica ions o basin de elopmen
and end-C e aceous dinosau aunal u no e . Sedimen a y Geology 337, 55–68.
doi:10.1016/j.sedgeo.2016.03.006
MANUS CRIP T
ACCEP TED
ACCEPTED MANUSCRIPT
Fonde illa, V., Dina ès-Tu ell, J., Vila, B., Le Loeu , J., Es ada, R., Oms, O.,
Galoba , À., 2016b. Magne os a ig aphy o he Maas ich ian con inen al eco d in he Uppe
Aude Valley (no he n Py enees, F ance): Placing age cons ain s on he succession o dinosau -
bea ing si es. C e aceous Resea ch 57, 457–472. doi:10.1016/j.c e es.2015.08.009
Fonnesu, F., 1984. Es a ig a ía ísica y análisis de acies de la secuencia de Figols en e
el ío Nogue a Palla esa e Iscles:(p o s. de Lé ida y Huesca). Uni e sidad Au ónoma,
Depa amen o de Es a ig a ía y Geología His ó ica.
Galb un, B., Feis , M., Colombo, F., Rocchia, R., Tamba eau, Y., 1993.
Magne os a ig aphy and bios a ig aphy o C e aceous-Te ia y con inen al deposi s, Age
Basin, P o ince o Le ida, Spain. Palaeogeog aphy, Palaeoclima ology, Palaeoecology 102, 41–
52. doi:10.1016/0031-0182(93)90004-3
Ga cia, G., Vianey-Liaud, M., 2001. Dinosau eggshells as bioch onological ma ke s in
Uppe C e aceous con inen al deposi s. Palaeogeog aphy, Palaeoclima ology, Palaeoecology
169, 153–164. doi:10.1016/S0031-0182(01)00215-2
Ga ido-Megías, A., Ríos, L., 1972. Sín esis geológica del Secunda io y Te cia io en e
los íos Cinca y Seg e (Pi ineo cen al de la e ien e su pi enaica, p o incias de Huesca y
Lé ida). Bole ín Geológico y Mine o de España 83, 1–47.
Ghee b an , E., Cod ea, V., Hosu, A., 1999. Décou e e de e éb és dans les Calcai es
de Rona (Thané ien ou Spa nacien), T ansyl anie, Roumanie: les plus anciens mammi e es
cénozoiques d’Eu ope O ien ale. Bi khäuse .
MANUS CRIP T
ACCEP TED
ACCEPTED MANUSCRIPT
Gómez-G as, D., Roigé, M., Fonde illa, V., Oms, O., Boya, S., Remacha, E., 2016.
P o enance cons ain s on he T emp Fo ma ion paleogeog aphy (sou he n Py enees): Eb o
Massi VS Py enees sou ces. C e aceous Resea ch 57, 414–427.
doi:10.1016/j.c e es.2015.09.010
G ads ein, F.M., Ogg, J.G., Schmi z, M., Ogg, G., 2012. The geologic ime scale 2012.
else ie .
Jackson, M., Swanson-Hysell, N.L., 2012. Rock magne ism o emagne ized ca bona e
ocks: ano he look. Geological Socie y, London, Special Publica ions 371, 229.
doi:10.1144/SP371.3
Jiang, X., Liu, Y., Ji, S., Zhang, X., Xu, L., Jia, S., Lü, J., Yuan, C., Li, M., 2011.
Dinosau -bea ing s a a and K/T bounda y in he Luanchuan-Tan ou Basin o wes e n Henan
P o ince, China. Science China Ea h Sciences 54, 1149. doi:10.1007/s11430-011-4186-1
Juá ez, M.T., Ose e, M.L., Meléndez, G., Lange eis, C.G., Zijde eld, J.D.A., 1994.
Ox o dian magne os a ig aphy o he Aguilón and Tosos sec ions (Ibe ian Range, Spain) and
e idence o a p e-Oligocene o e p in . Physics o he Ea h and Plane a y In e io s 85, 195–211.
doi:10.1016/0031-9201(94)90017-5
Lau en , Y., Bilo e, M., Le Loeu , J., 2002. La e Maas ich ian con inen al e eb a es
om sou hwes e n F ance: co ela ion wi h ma ine auna. Palaeogeog aphy, Palaeoclima ology,
Palaeoecology 187, 121–135. doi:10.1016/S0031-0182(02)00512-6
Leonha d , R., 2006. Analyzing ock magne ic measu emen s: The RockMagAnalyze
1.0 so wa e. Compu e s & Geosciences 32, 1420–1431. doi:10.1016/j.cageo.2006.01.006
MANUS CRIP T
ACCEP TED
ACCEPTED MANUSCRIPT
Leyme ie, A., 1862. Ape çu géognos ique des Pe i es Py énées e pa iculiè emen de la
mon agne d’Ausseing. Bull. Soc. Geol. F . 19, 1091–1096.
Liebau, A., 1973. El Maas ich iense laguna (Ga umniense) de Isona. Lib o-Guıa, XIII
Coloquio Eu opeo Mic opaleon ologıa 87–112.
Llompa , C., 1979. Yacimien o de huellas de pisadas de ep il en el C e ácico
p epi enaico. Ac a geológica hispánica 14, 333–336.
López-Ma inez, N., A de ol, L., A ibas, M.E., Ci is, J., Gonzalez-Delgado, A., 1998.
The geological eco d in non-ma ine en i onmen s a ound he K/T bounda y (T emp Fo ma ion,
Spain). Bulle in de la Socie e Geologique de F ance 169, 11.
López-Ma ínez, N., A ibas, M.E., Robado , A., Vicens, E., A dè ol, L., 2006. Los
ca bona os danienses (Unidad 3) de la FM T emp (Pi ineos Su -Cen ales) : paleogeog a ía y
elación con el lími e C e ácico-Te cia io. Re is a de la Sociedad Geológica de España 19, 233–
255.
López-Ma ínez, N., Fe nández-Ma ón, M.T., Valle, M.F., 1999. The succession o
e eb a es and plan s ac oss he C e aceous-Te ia y bounda y in he T emp Fo ma ion, Age
alley (sou h-cen al Py enees, Spain). Geobios 32, 617–627.
López-Ma ınez, N., Canudo, J.I., A dè ol, L., Sube biola, X.P., O ue-E xeba ia, X.,
Cuenca-Bescós, G., Ruiz-Omeñaca, J.I., Mu elaga, X., Feis , M., 2001. New dinosau si es
co ela ed wi h Uppe Maas ich ian pelagic deposi s in he Spanish Py enees: implica ions o
he dinosau ex inc ion pa e n in Eu ope. C e aceous Resea ch 22, 41–61.
doi:10.1006/c es.2000.0236

MANUS CRIP T
ACCEP TED
ACCEPTED MANUSCRIPT
Mannion, P.D., Upchu ch, P., Ca ano, M.T., Ba e , P.M., 2011. Tes ing he e ec o
he ock eco d on di e si y: a mul idisciplina y app oach o elucida ing he gene ic ichness o
sau opodomo ph dinosau s h ough ime. Biological Re iews 86, 157–181. doi:10.1111/j.1469-
185X.2010.00139.x
Ma mi, J., Blanco, A., Fonde illa, V., Dalla Vecchia, F.M., Sellés, A.G., Vicen e, A.,
Ma ín-Closas, C., Oms, O., Galoba , À., 2016. The Molí del Ba ó-1 si e, a di e se ossil
assemblage om he uppe mos Maas ich ian o he sou he n Py enees (no h-eas e n Ibe ia).
C e aceous Resea ch 57, 519–539. doi:10.1016/j.c e es.2015.06.016
Ma mi, J., Luján, Á.H., Rie a, V., Gae e, R., Oms, O., Galoba , À., 2012. The younges
species o Polys e non: A new bo h emydid u le om he uppe mos Maas ich ian o he
sou he n Py enees. C e aceous Resea ch 35, 133–142. doi:10.1016/j.c e es.2011.12.004
Ma y, D., 2001. Sedimen ology, Paleoecology and S a ig aphy o he Palus ine Facies
Rognacien a he K/T-bounda y in Sou he n F ance. Geologisch-Paläon ologisches Ins i u .
Mas ie a, A., Ullas e, J., 1983. Essai de syn hèse s a ig aphique des couches
con inen ales de la in du C é acé des Py enées ca alanes (NE de lEspagne). Géologie
Médi e anéenne 20, 283–290.
Médus, J., Feis , M., Rocchia, R., Ba en, D.J., Bocle , D., Colombo, F., Tamba eau, Y.,
Villa e, J., 1988. P ospec s o ecogni ion o he palynological C e aceous Te ia y bounda y
and an i idium anomaly in nonma ine acies o he eas e n Spanish Py enees: a p elimina y
epo . Newsle e s on S a ig aphy 18, 123–138. doi:10.1127/nos/18/1988/123
MANUS CRIP T
ACCEP TED
ACCEPTED MANUSCRIPT
Mey, P.H.W., Nag egaal, P.J.C., Robe i, K.J., Ha e el , J.J.A., 1968.
Li hos a ig aphic subdi ision o pos -He cynian deposi s in he sou h-cen al Py enees, Spain.
Leidse Geologische Mededelingen 41, 221–228.
Mo eno-Azanza, M., Bauluz, B., Canudo, J.I., Pué olas-Pascual, E., Sellés, A.G., 2014.
A e-e alua ion o a . Megalooli hidae eggshell agmen s om he uppe mos C e aceous o he
Py enees and implica ions o c ocodylomo ph eggshell s uc u e. His o ical Biology 26, 195–
205. doi:10.1080/08912963.2013.786067
Muñoz, J.A., 1992. E olu ion o a con inen al collision bel : ECORS-Py enees c us al
balanced c oss-sec ion. Th us ec onics 235–246.
Mu elaga, X., Canudo, J.I., 2005. Desc ipción de los es os de quelonios del
Maas ich iense supe io de A en y Se aduy (Huesca). Geogace a 38, 51–54.
Nag egaal, P.J.C., Van Vlie , A., B ouwe , J., 1983. Syn ec onic coas al o lap and
concu en u bidi e deposi ion: The Uppe C e aceous A en sands one in he Sou h-Cen al
Py enees, Spain. Sedimen a y Geology 34, 185–218. doi:10.1016/0037-0738(83)90086-6
Nakkady, S.E., 1950. A new o amini e al auna om he Esna Shales and Uppe
C e aceous Chalk o Egyp . Jou nal o Paleon ology, 675–692.
Na áez, I., B ochu, C.A., Escaso, F., Pé ez-Ga cía, A., O ega, F., 2016. New Spanish
La e C e aceous eusuchian e eals he synch onic and sympa ic p esence o wo
allodaposuchids. C e aceous Resea ch 65, 112–125. doi:10.1016/j.c e es.2016.04.018
Na a e e, R., Rod íguez-López, J.P., Liesa, C.L., So ia, A.R., Veloso, F. de M.L.,
2013. Changing physiog aphy o i basins as a con ol on he e olu ion o mixed siliciclas ic–
MANUS CRIP T
ACCEP TED
ACCEPTED MANUSCRIPT
ca bona e back-ba ie sys ems (Ba emian Ibe ian Basin, Spain). Sedimen a y Geology 289, 40–
61. doi:10.1016/j.sedgeo.2013.02.003
Nede b ag , A.J., 1991. La e C e aceous Bios a ig aphy and De elopmen o
He e ohelicidae (Plank ic Fo amini e a). Mic opaleon ology 37, 329–372.
h ps://doi.o g/10.2307/1485910
Ogg, J.G., Hinno , L.A., Huang, C., 2012. Chap e 27 - C e aceous, in: The Geologic
Time Scale. Else ie , Bos on, 793–853. h ps://doi.o g/10.1016/B978-0-444-59425-9.00027-5
Oms, O., Dina ès-Tu ell, J., Vicens, E., Es ada, R., Vila, B., Galoba , À., B a o, A.M.,
2007. In eg a ed s a ig aphy om he Vallceb e Basin (sou heas e n Py enees, Spain): New
insigh s on he con inen al C e aceous−Te ia y ansi ion in sou hwes Eu ope.
Palaeogeog aphy, Palaeoclima ology, Palaeoecology 255, 35–47.
doi:10.1016/j.palaeo.2007.02.039
Oms, O., Fonde illa, V., Rie a, V., Ma mi, J., Vicens, E., Es ada, R., Anadón, P., Vila,
B., Galoba , À., 2016. T ansi ional en i onmen s o he lowe Maas ich ian Sou h-Py enean
Basin (Ca alonia, Spain): The Fumanya Membe idal la . C e aceous Resea ch 57, 428–442.
doi:10.1016/j.c e es.2015.09.004
Oms, O., Pé ez-Cano, J., Fonde illa, V., Anadón, P., Ibáñez-Insa, J., Rejas, M.,
Fe nandez-Tu iel, J.L., Dina ès-Tu ell, J., Pa do, P., Es ada, R., 2014. On he sea ch o he K-
PG bounda y in he e es ial eco d in Eu ope: p elimina y geochemical da a om he Puig
Ped ós sec ion (T emp basin, Ca alonia, Spain). Abs ac book Recons uc ing he e es ial end-
C e aceous paleoen i onmen s in Eu ope (T emp, 2014). Paleon ologia i E olució Special
memoi 7, 74.
MANUS CRIP T
ACCEP TED
ACCEPTED MANUSCRIPT
O ega, F., Ba de , N., Ba oso-Ba cenilla, F., Callapez, P.M., Camb a-Moo, O.,
Gómez, V.D.-, Díaz, V.D., Domingo, L., El i a, A., Escaso, F., Ga cía-Oli a, M., Gómez, B.,
Houssaye, A., Knoll, F., Ma cos-Fe nández, F., Ma ín, M., Mocho, P., Na áez, I., Ga cía, A.P.-
, Pey o , D., Segu a, M., Se ano, H., To ices, A., Vidal, D., Sanz, J.L., 2015. The bio a o he
Uppe C e aceous si e o “Lo Hueco” (Cuenca, Spain). Jou nal o Ibe ian Geology 41.
Ose e, M.-L., Gialanella, P.-R., Gómez, J.J., Villalaín, J.J., Goy, A., Helle , F., 2007.
Magne os a ig aphy o Ea ly–Middle Toa cian expanded sec ions om he Ibe ian Range
(cen al Spain). Ea h and Plane a y Science Le e s 259, 319–332.
doi:10.1016/j.epsl.2007.04.048
Panaio u, A.G., Ciobăne e, D., Panaio u, C.G., Panaio u, C.E., Csiki, Z., 2011. New
palaeomagne ic da a om he Haţeg Basin, Romania, in: Abs ac Book, 8 h Romanian
Symposium o Paleon ology, Bucha es (Romania). Ed. A s Docendi, Bucha es , 84–85.
Panaio u, C.G., Panaio u, C.E., 2010. Palaeomagne ism o he Uppe C e aceous
Sânpe u Fo ma ion (Haţeg Basin, Sou h Ca pa hians). Palaeogeog aphy, Palaeoclima ology,
Palaeoecology 293, 343–352. doi:10.1016/j.palaeo.2009.11.017
Peláez-Campomanes, P., López-Ma ínez, N., Ál a ez-Sie a, M.A., Daams, R., 2000.
The Ea lies Mammal o he Eu opean Paleocene: The Mul i ube cula e Hainina. Jou nal o
Paleon ology 74, 701–711.
Pe eda-Sube biola, X., Canudo, J.I., C uzado-Caballe o, P., Ba co, J.L., López-
Ma ínez, N., Oms, O., Ruiz-Omeñaca, J.I., 2009. The las had osau id dinosau s o Eu ope: A
new lambeosau ine om he Uppe mos C e aceous o A en (Huesca, Spain). Comp es Rendus
Pale ol 8, 559–572. doi:10.1016/j.c p .2009.05.002
MANUS CRIP T
ACCEP TED
ACCEPTED MANUSCRIPT
Vila, B., Sellés, A.G., B usa e, S.L., 2016. Di e si y and aunal changes in he la es
C e aceous dinosau communi ies o sou hwes e n Eu ope. C e aceous Resea ch 57, 552–564.
doi:10.1016/j.c e es.2015.07.003
Villalaín, J.J., Ose e, M.L., Vegas, R., Ga cía-Dueñas, V., Helle , F., 1994. Widesp ead
Neogene emagne iza ion in Ju assic limes ones o he Sou h-Ibe ian palaeoma gin (Wes e n
Be ics, Gib al a A c). Physics o he Ea h and Plane a y In e io s 85, 15–33. doi:10.1016/0031-
9201(94)90005-1
Villalba-B e a, S., Ma ín-Closas, C., 2013. Uppe C e aceous paleogeog aphy o he
Cen al Sou he n Py enean Basins (Ca alonia, Spain) om mic o acies analysis and cha ophy e
bios a ig aphy. Facies 59, 319–345. doi:10.1007/s10347-012-0317-1
Villalba-B e a, S., Ma ín-Closas, C., 2011. A cha acean hallus wi h a ached
gy ogoni es and associa ed ossil cha ophy es om he Maas ich ian o he Eas e n Py enees
(Ca alonia, Spain). Jou nal o Phycology 47, 131–143. doi:10.1111/j.1529-8817.2010.00947.x
Villalba-B e a, S., Ma ín-Closas, C., Ma mi, J., Gomez, B., Fe nández-Ma ón, M.T.,
2012. Pea - o ming plan s in he Maas ich ian coals o he Eas e n Py enees. Geologica Ac a: an
in e na ional ea h science jou nal 10. doi:h p://dx.doi.o g/10.1344/105.000001711
Vogle , J., 1941. Obe -Ju a und K eide on Misol. Bei age zu Geologie on
Niede landisch-Indian 4, 243–293.
Voo wijk, G.H., 1937. Fo amini e a om he uppe C e aceous o Habana, Cuba. Akad.
We ., Ams e dam, Sec . Sci. 40, 190–198.

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 .
MANUS CRIP T
ACCEP TED
ACCEPTED MANUSCRIPT
MANUS CRIP T
ACCEP TED
ACCEPTED MANUSCRIPT
MANUS CRIP T
ACCEP TED
ACCEPTED MANUSCRIPT
MANUS CRIP T
ACCEP TED
ACCEPTED MANUSCRIPT

MANUS CRIP T
ACCEP TED
ACCEPTED MANUSCRIPT
MANUS CRIP T
ACCEP TED
ACCEPTED MANUSCRIPT
MANUS CRIP T
ACCEP TED
ACCEPTED MANUSCRIPT
MANUS CRIP T
ACCEP TED
ACCEPTED MANUSCRIPT