Magnetostratigraphy of the Maastrichtian continental record in the Upper Aude Valley (northern Pyrenees, France): Placing age constraints on the succession of dinosaur-bearing sites

Cretaceous Research xxx (2015) 1e16

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Magnetostratigraphy of the Maastrichtian continental record in the Upper Aude Valley (northern Pyrenees, France): Placing age constraints on the succession of dinosaur-bearing sites
s-Turell b, Bernat Vila c, Jean Le Loeuff d, Rita Estrada a, Víctor Fondevilla a, *, Jaume Dinare a
e Oriol Oms , Angel Galobart a
noma de Barcelona, Carrer de l'Eix central, E-08193 Cerdanyola del Vall
Departament de Geologia (Estratigrafia), Facultat de Ci
encies, Universitat Auto es, Barcelona, Catalonia, Spain b Istituto Nazionale di Geofisica e Vulcanologia, Via di Vigna Murata, 605, I-00143 Rome, Italy c Grupo AragosauruseIUCA, Paleontología, Facultad de Ciencias Universidad de Zaragoza, E-50009 Zaragoza, Spain d Mus ee des Dinosaures, 11260 Esp eraza, France e
de Paleontologia Miquel Crusafont, Carrer Escola Industrial, 23, E-08201, Sabadell, Barcelona, Catalonia, Spain Institut Catala

a r t i c l e i n f o

a b s t r a c t

Article history: Received 22 April 2015 Received in revised form 6 August 2015 Accepted in revised form 19 August 2015 Available online xxx

The first detailed stratigraphic succession of the Upper Cretaceous continental record from the Upper Aude Valley (southern France) is presented together with a magnetostratigraphic study. The combined rieures Fm (Lower Red Marls), constratigraphy and magnetostratigraphy of the Marnes rouges infe strained by biochronological markers such as charophyte occurrence and revised dinosaur eggshells, results in a succession of fluvial red beds dated from chron C32n to the top of chron C31r. It implies an earliest Maastrichtian age close to the C32n.1n-C31r reversal for the majority of the dinosaur sites including Bellevue. In contrast, the upper Maastrichtian is likely represented by a short interval within the lacustrine-palustrine Calcaires et argiles de Vignevieille Fm (Vignevieille Limestones), or it might even not be recorded. The proposed age indicates that the marine to continental transition, as a result of the Late Cretaceous transgression, took place earlier in the north Pyrenean basin than in the southern area. © 2015 Elsevier Ltd. All rights reserved.

Keywords: Dinosaurs Magnetostratigraphy Maastrichtian Bellevue Pyrenees France

1. Introduction The Late Cretaceous vertebrate successions of southwestern Europe are key to an assessment of the diversity, dynamics and changes displayed by the last dinosaur faunas of the European archipelago in the last few million years before the CretaceousPaleogene mass extinction event. The ancient Ibero-Armorican island encompassed the current areas of Portugal, central and northeastern Spain and southern France and produced a wealth of fossil localities (Csiki et al., 2015). The geographic region of southern France, ranging from the Arc Basin (Provence) to the Haute ze Garonne (north Pyrenean area) and including the Villeveyrac-Me (Languedoc area) (Fig. 1A), has provided a rich dinosaur fossil record in its Upper Cretaceous continental deposits. While many

* Corresponding author. E-mail address: [email protected] (V. Fondevilla).

stratigraphic, magnetostratigraphic and sedimentological works have been carried out in Provence (Cojan, 1993; Cojan, Moreau, & Stott, 2000; Cojan & Moreau, 2006; Westphal & Durand, 1990), the northeastern Pyrenean record lacks such studies, except for the seminal works of Bilotte (1978, 1985) and Bilotte, Tambareau, and Villatte (1983). In spite of the fact that many reference dinosaur localities have been reported in this area (Bilotte, Duranthon, Clottes, & Raynaud, 1985; Buffetaut, 2005; Buffetaut & Le Loeuff, 1989, 1997; Buffetaut et al., 1989, 1997), only limited age constraints have been provided using terrestrial biochronological
re, Tambareau, & Villatte, 1980, 1989; Bilotte, 1985; markers (Bessie Garcia & Vianey-Liaud, 2001; Marty, 2001). In consequence, these poorly dated vertebrate localities in the northern Pyrenees remain in an unclear chronostratigraphic position between the upper Campanian and the lower and upper Maastrichtian. As a result, the vertebrate remains from these sites are difficult to compare and correlate with those from other regions of the Ibero-Armorican domain, such as the southern Pyrenees.

http://dx.doi.org/10.1016/j.cretres.2015.08.009 0195-6671/© 2015 Elsevier Ltd. All rights reserved.

Please cite this article in press as: Fondevilla, V., et al., Magnetostratigraphy of the Maastrichtian continental record in the Upper Aude Valley (northern Pyrenees, France): Placing age constraints on the succession of dinosaur-bearing sites, Cretaceous Research (2015), http://dx.doi.org/ 10.1016/j.cretres.2015.08.009

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Fig. 1. A, Upper Cretaceous continental deposits of southwestern Europe. Modified from Bilotte (1978), Garcia and Vianey-Liaud (2001) and Oms et al. (2007). B, Geological setting of the Aude Valley area. NPFT, North Pyrenean frontal thrust; SPFT, Sub-Pyrenean frontal thrust. Modified from Bilotte (1978) and Marty and Meyer (2006).

In the southern Pyrenees, recent studies of the magnetostratigraphy (Canudo et al., 2015; Oms et al., 2007; PeredaSuberbiola et al., 2009; Vila et al., 2012) and biostratigraphy (Vila et al., 2011; Villalba-Breva & Martín-Closas, 2012; Villalba-Breva, ndez-Marro n, 2012; ViceMartín-Closas, Marmi, Gomez, & Ferna nte, Martín-Closas, Arz, & Oms, 2015) have resulted in a well-

documented chronostratigraphic framework for the Maastrichtian terrestrial record. The need has thus become apparent to combine lithostratigraphy, magnetostratigraphy and biostratigraphy (based on dinosaur eggshells and charophytes) to study the continental deposits of the Upper Aude Valley in order to integrate the northern Pyrenean record with that of the southern foothills. The classic and

Please cite this article in press as: Fondevilla, V., et al., Magnetostratigraphy of the Maastrichtian continental record in the Upper Aude Valley (northern Pyrenees, France): Placing age constraints on the succession of dinosaur-bearing sites, Cretaceous Research (2015), http://dx.doi.org/ 10.1016/j.cretres.2015.08.009

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^teau area (Beetschen, renowned egg sites from the Rennes-le-Cha Dughi, & Sirugue, 1977; Beetschen, 1985; Breton, Fourier, & , 1986; Caillaud, 1968; Cousin, Breton, Fournier, & Watte, Watte 1989) and other reference vertebrate sites near the village of Campagne-sur-Aude, such as the Bellevue locality, have yielded an important fossil record (Bilotte et al., 1985; Buffetaut & Le Loeuff, 1989; Buffetaut et al., 1989; Clottes & Raynaud, 1983). This includes the titanosaurid Ampelosaurus atacis, rhabdodontid ornithopods, nodosaurid ankylosaurs, and non-avian theropods (Le Loeuff, 1995; Le Loeuff, 2005). Such a dinosaur association exemplifies the dominant assemblage of the late Campanian-early Maastrichtian of the Ibero-Armorican domain (Le Loeuff, Buffetaut, & Martin, 1994a), which was later replaced by the hadrosauroid-dominated assemblages of the late Maastrichtian. It is also similar to those from other Campanian-Maastrichtian sites, ~o including localities in Provence and the Iberian Peninsula (e.g. Lan and Lo Hueco; Pereda-Suberbiola, Astibia, Murelaga, Elorza, &  mez-Alday, 2000; Ortega et al., 2008). Thus, our combined Go geological and paleontological study will permit us to incorporate this rich dinosaur area into an Ibero-Armorican chronostratigraphic framework and compare it with other calibrated sections of the southern Pyrenees and Provence. 2. Geological setting The Pyrenees are an Alpine fold-and-thrust belt formed by the collision between the Iberian and the European plates. This generated a system of thrusts that affected a Hercynian basement and pre- and syntectonic sedimentary covers deposited from the Late Cretaceous mainly to the Oligocene. These foreland basins were developed in the north and the south of the mountain belt (known as north and south Pyrenean zones in France and Spain, respectively). However, the northern Pyrenean thrust system ~ oz, generated less cortical shortening than the southern one (Mun 1992). The northern belt of the Pyrenees is composed of thick Mesozoic marine successions, accumulated in a subsiding rift or transtensional basin until the Late Cretaceous, when the Alpine orogenesis began. Then, the compressive stage inverted the extensional structures and generated thrust systems, triggering the development of E-W-oriented elongate foreland basins connected to the Atlantic (Teixell, 2004 and references therein). The Upper Aude Valley is located in the eastern sector of a narrow foreland basin known as the Sub-Pyrenean area. It is compressed between the North Pyrenean frontal thrust (towards the south) and the SubPyrenean frontal thrust (to the north), separating this area from the northern Aquitaine basin (Fig. 1B). To the east, the SubPyrenean area is limited by a Paleozoic structural high known as the Mouthoumet Massif. Internally, the Sub-Pyrenean area is structured by several anticlines and synclines that follow a WNWESE orientation. To be specific, the studied area is located on the
re et al., southern side of the Couiza-Arques syncline (Fig. 1B) (Bessie 1989; Bilotte, 1990); both thrust systems and folds were developed during the early Thanetian (Tambareau, Crochet, Villatte, & Deramond, 1995). In the Oligocene, normal faults following an NEeSW orientation were developed in the Upper Aude Valley and
res during an extensive phase (Bessie
re et al., the nearby Corbie 1989). In the Upper Aude Valley, continental sedimentation started in the latest Campanian around the Mouthoumet Massif, following a regression that lasted until the Danian. In the studied area, these
s d’Alet (Alet continental deposits are represented by the fluvial Gre Sandstone) and the overlying red beds of the Marnes rouges rieures Fm (Lower Red Marls). The Alet Sandstone in turn infe overlies the marine Marnes de Sougraigne Fm (Sougraigne Marls),

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which is of late Santonian age. Consequently, a Campanian age is attributed to the Alet Sandstone on the basis of its stratigraphic position (Bilotte et al., 1983; Bilotte, 1985). The Lower Red Marls are divided into four members, which are from base to top (Bilotte, 1978, 1985; Bilotte et al., 1983): a) Marnes rouges de Campagne
s des Estous (Estous Sandstone), c) (Red Marls of Campagne), b) Gre Marnes rouges de la Maurine (Red Marls of Maurine) and d) Poudingue Fleuri (Fleuri Conglomerate) (Fig. 2). The Red Marls of Campagne provide a late Campanian palynofloral assemblage (Bilotte, 1985), so the remaining members have been considered Maastrichtian on the basis of their stratigraphic position, dinosaur eggshell biostratigraphy (Garcia & Vianey-Liaud, 2001; Le Loeuff,
re et al., 1980, 1989; 2005) and charophyte occurrence (Bessie Marty, 2001; Marty & Meyer, 2006). Regarding the Fleuri Conglomerate, an end-Maastrichtian age was suggested by Oms et al. (2007), based on its correlation with the south Pyrenean Reptile Sandstone in the Vallcebre syncline (which is located in chron C29r) and the Galante Conglomerate from the Arc Basin of Provence (also located within C29r, according to Westphal &
re et al. (1980, 1989) and Marty (2001) reDurand, 1990). Bessie ported Maastrichtian charophytes from the lower levels of the overlying palustrine limestones, known as the Calcaires et argiles de Vignevieille Fm (Vignevieille Limestones). Its upper levels, however, have yielded Danian charophytes, so the CretaceousPaleogene (K-Pg) transition has been placed within these limestones. In the north Pyrenean area, the iridium anomaly, which records the exact position of the K-Pg boundary, has been determined in the shallow marine deposits of the Larcan Quarry (Haute Garonne) (Rocchia, Robin, Tambareau, Villatte, & Bilotte, 1998) and
s, Fondecave-Wallez, in other marine units farther west (Peyberne
ne, Robin, & Rocchia, 1998). Eiche The Alet Sandstone and the Lower Red Marls become younger to the west due to facies migration in the regressive context, gradually passing diachronically to coastal and marine units in the west (Fig. 3) (Bilotte et al., 1983). Thus, the units evolve to the
s de Labarre (Labarre Sandstone, in Arie
getransitional-coastal Gre Plantaurel), the Calcaire de Nankin and the Marnes d’Auzas Fms (Nankin Limestone and Auzas Marls, respectively, in the Petites ne es-Haute Garonne). This latter unit represents sediments Pyre deposited in paralic to continental conditions during the late Maastrichtian (Bilotte, 1985; Laurent, Bilotte, & Le Loeuff, 2002; Bilotte & Andreu, 2006). The marine lateral equivalents of the continental units are represented by the Marnes de Plagne Fm
ge-Plantaurel) and the Marnes de Saint (Plagne Marls, in Arie Martory and Saint Loup Fms (Saint Martory and Saint Loup Marls, ne es-Haute Garonne; Bilotte et al., respectively, in the Petites Pyre 1983). 3. Material and methods In order to integrate the Upper Aude Valley fossil localities within a chronostratigraphic framework, we measured nine stratigraphic sections at localities with dinosaur fossil sites together with others that exhibit good rock exposures. Their location is given in Fig. 2. Amongst them, we selected the localities of Brezilhou, Bellevue and East Campagne-sur-Aude (Sections I, II and IV, respectively) to perform the magnetostratigraphic sampling due to their continuous succession and better outcrop conditions. Additionally, these localities are located near the River Aude relatively close to one another (up to two km), making their paleomagnetic signal more reliable and easier to compare. The laterally continuous Estous Sandstone, Fleuri Conglomerate and Vignevieille Limestones proved useful to correlate all the sections, resulting in a 130-mthick composite section (Fig. 4). Unfortunately, we did not study the Alet Sandstone and the lower meters of the Red Marls of Campagne

Please cite this article in press as: Fondevilla, V., et al., Magnetostratigraphy of the Maastrichtian continental record in the Upper Aude Valley (northern Pyrenees, France): Placing age constraints on the succession of dinosaur-bearing sites, Cretaceous Research (2015), http://dx.doi.org/ 10.1016/j.cretres.2015.08.009

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raza, with the location of the studied sections. Modified from Crochet et al. (1989). Sections: I, Brezilhou; II, Bellevue; III, Fig. 2. Geological map of the Upper Aude Valley at Espe ^teau; VIII, Roque Fumade; IX, La Valdieu. Campagne-sur-Aude; IV and V, East Campagne-sur-Aude; VI, Gourg de l'Encantado; VII, Rennes-le-Cha

Fig. 3. Schematic EeW cross section of north Pyrenean Upper Cretaceous deposits. Modified from Laurent, Le Loeuff, Bilotte, Buffetaut, and Odin (2001).

due to the lack of good exposure. We also found the same problem in the middle part of the Red Marls of Maurine (meters 67 to 91). The rest of the sections represent short stratigraphic successions, often with thick covered intervals. Small oriented blocks were generally collected from reddish soft mudstones and sandstones for paleomagnetic analysis. The emphasis was on sampling the most fine-grained lithology available. We sampled 77 sites from Brezilhou, Bellevue and East Campagne-sur-Aude. Additionally, we sampled three sites in a short interval of 4 m in the Roque Fumade section. Altogether, the sites include a total of 195 samples and encompass about 130 m of stratigraphic succession. A thick accumulation of colluvium along

the middle interval results in poor outcrop conditions that prevented sampling for about 25 m (Fig. 4). All samples were oriented in situ with a magnetic compass, and standard specimens were subsequently cut in the laboratory for paleomagnetic analysis. Natural remanent magnetization (NRM) and remanence through demagnetization were measured on a 2G Enterprises DC SQUID high-resolution pass-through cryogenic magnetometer (manufacturer noise level of 10
12 Am2) operated in a shielded room at the Istituto Nazionale di Geofisica e Vulcanologia in Rome, Italy. A Pyrox oven in the shielded room was used for thermal demagnetizations, and alternating field (AF) demagnetization was performed with three orthogonal coils installed in line with the cryogenic

Please cite this article in press as: Fondevilla, V., et al., Magnetostratigraphy of the Maastrichtian continental record in the Upper Aude Valley (northern Pyrenees, France): Placing age constraints on the succession of dinosaur-bearing sites, Cretaceous Research (2015), http://dx.doi.org/ 10.1016/j.cretres.2015.08.009

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Fig. 4. Sections and composite section of the studied area with paleomagnetic ChRM declination, inclination, polarity zones and VGP latitude. Crosses indicate “class 3” specimens (no data), open symbols denote “class 2” specimens (unreliable directions), and closed symbols mark “class 1” specimens (reliable directions). Magnetostratigraphic interpretation includes normal polarity (black), reverse polarity (white) and undefined polarity (grey) bands. Half-width bands indicate “class 1” intervals represented by data from a single stratigraphic level. Sections: I, Brezilhou (triangles for the paleomagnetic data); II, Bellevue (circles); IV, East Campagne-sur-Aude (squares); VIII, Roque Fumade (diamonds).

Please cite this article in press as: Fondevilla, V., et al., Magnetostratigraphy of the Maastrichtian continental record in the Upper Aude Valley (northern Pyrenees, France): Placing age constraints on the succession of dinosaur-bearing sites, Cretaceous Research (2015), http://dx.doi.org/ 10.1016/j.cretres.2015.08.009

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magnetometer. Progressive stepwise AF demagnetization was routinely used and applied after a single heating step to 150  C. AF demagnetization included 14 steps (4, 8, 13, 17, 21, 25, 30, 35, 40, 45, 50, 60, 80, 100 mT). After the AF demagnetization protocol, stepwise thermal demagnetization was continued up to temperatures of 600  C or above. Characteristic remanent magnetizations (ChRM) were computed by least-squares fitting (Kirschvink, 1980) on the orthogonal demagnetization plots (Zijderveld, 1967) using the software package Paldir (Utrecht University). The ChRM declination and inclination were used to derive the latitude of the virtual geomagnetic pole (VGP) of each sample. This parameter was taken as an indicator of the original magnetic polarity, normal polarity being indicated by positive VGP latitudes and reverse polarity by negative VGP latitudes. We resampled the eggshell material from several sites: Bellevue, formerly known as C3 after Buffetaut et al. (1989) (nine eggshell fragments, MDE-D-375 to MDE-D-378); La Valdieu (six eggshell fragments, MDE-D-387); Rennes-le-Ch^ ateau, known as F3 after Beetschen (1985) and Founbit after Cousin and Breton (2000) (33 eggshell fragments, MDE-D-382 to MDE-D-385); Arques (thin raza (two eggshell fragsections and resin stubs, MDE-D-396); Ve ments, MDE-D-395); Campagne-sur-Aude, equivalent to C1 of Buffetaut et al. (1989) (eggshell fragments in a block and one thin section; MDE-D-394); and four levels at Roque Fumade, equivalent to the F4 site of Beetschen (1985) (Roque Fumade levels 1, 2, 3, and 5; fourteen eggshell fragments, MDE-D-389, 390, 391, 393). In addition, we analyzed eggshell material from two new sites (Les Boudous: four eggshells, MDE-D-388; Gourg de l’Encantado: six eggshells, MDE-D-386), as well as from the new level 4 in the Roque Fumade section (Roque Fumade level 4; four eggshell fragments, MDE-D-392). The eggshell fragments were collected from eggs that were in situ in two levels and scattered in the sediment in eleven levels. All eggshells were first observed under a Nikon SMZ-10 binocular microscope and a Leica MZ16A stereomicroscope, in order to describe the inner and outer eggshell surfaces and the radial view, and to take measurements. Eight fragments were used to prepare thin slides following standard petrographical techniques; these were photographed under a Leica DM 2500P polarized light microscope at 1.25 and 5 magnification. Some fresh broken fragments were photographed under a Quanta 200 FEI XTE 325/ D8395 environmental scanning electron microscope (ESEM). The egg taxa were identified in terms of parataxonomy following Mikhailov (1997). The studied material is housed in the collection e des Dinosaures, Espe raza (Aude, France). of the Muse 4. Results 4.1. Lithostratigraphy With the aim of establishing the sedimentary context of the studied area, we identified ten different lithofacies in the Upper Aude Valley (Fig. 4). To allow comparison between the different Pyrenean basins, we followed similar descriptive criteria to those used in Oms et al. (2007) and Riera, Oms, Gaete, and Galobart (2009), such as lithology, color, texture, sedimentary structures, paleontological content if identified, and the architecture of the sedimentary bodies. The recognized lithofacies are summarized in Supplementary Information 1. The association of ochre-brown, orange, red and purple mudstones and siltstones here identified exhibits features of continental paleoenvironments, such as edaphic processes characterized by the occurrence of carbonate nodules, broad plant activity visible as root mottling due to subaerial exposure, and the presence of the continental trace fossil Spirographites (see Mayoral & Calzada, 1998). These fine sediments appear linked to meandering and braided

sandstone bodies, implying a fluvio-alluvial setting with river deposits and their associated floodplains or fine overbank deposits, both widely colonized by vegetation. Thus, this scenario is consistent with the previous studies of the zone (Bilotte, 1978, 1985; Bilotte et al., 1983). Also, we found a great similarity between the lithofacies of the Upper Aude Valley, specially the mudstones and siltstones of the Red Marls of Campagne and Maurine, and the red bed facies described by Riera et al. (2009). This implies that the paleoenvironment of the Upper Aude Valley did not differ so much from the near-marine alluvial-deltaic paleoenvironment argued for the south Pyrenean ‘lower red unit’ of the Tremp Fm (Díez-Canseco, Arz, Benito, Díaz-Molina, & Arenillas, 2014; Oms et al., 2007; Riera et al., 2009; Rosell, Linares, & Llompart, 2001; Vila et al., 2013). At the top of the succession, the Vignevieille Limestones represent palustrine and lacustrine facies. This unit, which displays karstification, varies greatly in thickness, from about 5 m in the western sector to more than fifteen in the Roque Fumade section, further north. 4.2. Magnetostratigraphy The NRM intensity of the demagnetized specimens generally ranged from 0.3 to 2.5 mA/m, occasionally with lower intensities in certain limited grayish lithologies. Specimens from 15 sites provided scattered demagnetization data, preventing the computation of any magnetization components. These samples are qualified as “class 3” samples (Fig. 5I). In general, the intensity of the samples dropped noticeably after the first heating step at 150  C. Subsequent stepwise AF demagnetization up to 100 mT generally did not produce substantial changes, suggesting hematite as the main magnetic mineral carrier (in addition possibly to goethite). Above these demagnetizing fields, stepwise thermal demagnetization was resumed up to about 600  C, at which point samples generally appear fully demagnetized. A few samples, however, demagnetize at lower temperatures of around 450-500  C (Fig. 5A and D) or require higher temperatures (Fig. 5F). Thermal demagnetization trajectories above the AF demagnetization usually trend toward the origin of the demagnetization diagram and define the characteristic remanent magnetization (ChRM). The ChRM components are either northerly and moderately steeply downward oriented (Fig. 5AeC) or southerly and upward oriented (Fig. 5DeF) in bedding-corrected coordinates (“class 1” samples). For a number of samples the demagnetization data are scattered, and no linear trajectory trending to the origin can be unambiguously defined. We consider these samples to be unreliable and qualify them as “class 2” (Fig. 5GeH), implying that they cannot be used to derive the magnetostratigraphy. Instead, the “class 1” samples, which display a clear ChRM trajectory including several demagnetization steps trending towards the origin, are interpreted as dual-polarity primary magnetization directions and are used to derive the local magnetostratigraphy (Fig. 4). The subhorizontal attitude of the sampled strata prevents a fold test from being performed. Hence, it remains a possibility that some of the “class 1” directions might represent a secondary magnetization (see discussion below). In any case, the presence of a dual-polarity ChRM component that does not seem to be controlled by lithology with means that are almost perfectly antipodal (Fig. 6) is an indication of the primary nature of the isolated components. The results indicate that the lower 46 m of the studied composite section are consistently of normal polarity with a predominance of “class 1” samples (N1 in Fig. 4). Within this N1 magnetozone, the interval 39e43 m includes numerous “class 3” samples but also an isolated “class 1” sample at about meter 40 (Fig. 5D). This is depicted as a half-width strip in the polarity column (Fig. 4). Above N1 and up to meter 66 “class 1” samples

Please cite this article in press as: Fondevilla, V., et al., Magnetostratigraphy of the Maastrichtian continental record in the Upper Aude Valley (northern Pyrenees, France): Placing age constraints on the succession of dinosaur-bearing sites, Cretaceous Research (2015), http://dx.doi.org/ 10.1016/j.cretres.2015.08.009

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Fig. 5. Representative “class 1” (AeF), “class 2” (GeH) and “class 3” (I) orthogonal demagnetization diagrams in geographic coordinates for the studied sections in the Aude region. The stratigraphic position in meters, the natural remanent magnetization (NRM) intensity, and some demagnetization steps are indicated. Open and closed symbols indicate projections onto the upper and lower hemisphere respectively. The computed ChRM direction is shown by a solid thick grey line.

depict a reverse magnetozone (R1, Fig. 4) although an isolated normal sample is present at about meter 59. The upper part of the succession is predominantly of reverse polarity (R2, Fig. 4), including two small intervals of isolated “class 1” normal samples. The uppermost part above meter 122, straddling the Vignevieille Limestone, contains a rapid succession of samples of either normal or reverse polarity, some of them represented by single samples, so this interval is considered ambiguous and depicted grey in Fig. 4.

4.3. Dinosaur eggshells Historically, dinosaur eggs and eggshells have been found in several localities within the department of Aude. The first reports correspond to the works of Plaziat (1961), Caillaud (1968), Beetschen et al. (1977) and Beetschen (1985), who described  de Roqueeggshell fragments from the localities of Saint Andre ^teau, longue, Albas I, Albas II, F1 (AeB), F2 (AeB), Rennes-le-Cha
s. Subsequent Roque Fumade (F4 and F5), Les Labadous and Grane

Please cite this article in press as: Fondevilla, V., et al., Magnetostratigraphy of the Maastrichtian continental record in the Upper Aude Valley (northern Pyrenees, France): Placing age constraints on the succession of dinosaur-bearing sites, Cretaceous Research (2015), http://dx.doi.org/ 10.1016/j.cretres.2015.08.009

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Fig. 6. Stereographic projection of the “class 1” computed components in geographic coordinates (in situ) (strata are subhorizontal). Open and closed symbols indicate projections onto the upper and lower hemisphere respectively. Mean direction and statistics are given. N ¼ number of samples; Dec ¼ declination; Inc ¼ inclination; k ¼ Fisher's statistical parameter; a95 ¼ semiangle of the 95% cone of confidence.

work confirmed the fossil richness of dinosaur eggs in the area ^teau, and many excavations around the village of Rennes-le-Cha were conducted at this locality (Breton et al., 1986; Cousin et al., 1989). Buffetaut et al. (1989) reported eggs and eggshell fragments in the Campagne-sur-Aude, C2 and E1 sites. From the 1990s onwards various authors (Cousin, 1997a, 1997b; Garcia, 1998; Cousin & Breton, 2000; Vianey-Liaud & Garcia, 2000; Garcia & Vianey-Liaud, 2001) have provided further data from new localities raza, La Valdieu, Ruisseau de (Le Linas, Bellevue, Arques, Auriac, Ve
de) and reviewed the material from previously known sites la Pine ^teau, Roque Fumade, Campagne-sur-Aude, Grane
s, (Rennes-le-Cha  de Roquelongue, Les Labadous). In the present Albas, Saint Andre study we have reviewed the eggshell material collected in some of the studied sections of the Upper Aude Valley (see methods). Two main oofamilies are present in the Upper Aude Valley: Megas & Galobart, 2015. loolithidae Zhao, 1979 and Cairanoolithidae Selle The former is typically associated with titanosaurid sauropods (Chiappe, Salgado, & Coria, 2001) and is represented in the area by the oospecies Megaloolithus aureliensis, Megaloolithus siruguei, Megaloolithus mamillare and M. baghensis (formerly M. pseudomamillare); the latter is represented in the area by the oospecies Cairanoolithus dughii and Cairanoolithus roussetensis and s & has been tentatively attributed to nodosaurid ankylosaurs (Selle Galobart, 2015). We also provided material from two new sites (Les Boudous and Gourg de l’Encantado) and one new eggshell level at the Roque Fumade section (level 4) (Fig. 7). Ten eggshell fragments

have been collected from the site of Les Boudous (four samples) and Gourg de l’Encantado (six samples), in the lower part of the Red Marls of Maurine. These have an eggshell thickness ranging from 1.56 to 2.23 mm (Les Boudous: 2.02 mme2.23 mm, mean value 2.13 mm; Gourg de l’Encantado: 1.56 mme2.02 mm, mean value 1.86 mm) and a smooth outer surface with a very few sinuous ridges. Radial sections show columnar shell units, some of them with straight edges partially fused between adjacent units (Fig. 7A and A0 ). The growth lines are slightly arched or almost horizontal. Respiratory channels are straight and slender and are found at the edges of the shell units. All these parataxonomical characters allow the material to be referred to the oospecies Cairanoolithus dughii Vianey-Liaud, Mallan, Buscail, & Montgelard, 1994. Four eggshell fragments have been collected from level 4 at the Roque Fumade site, in the upper part of the Red Marls of Maurine. These eggshell fragments are thin (thickness ranging from 1.52 mm to 2.17 mm; mean value: 1.89 mm) and show an external surface covered with rounded nodes that are sometimes fused forming short ridges. Radial thin sections show small spherulitic fan-shaped units (Fig. 7B) that are higher than wide (H/W ratio ~2). Their boundaries are distinct, with no fused units. The growth lines appear arched from the base to the top of the units (Fig. 7B’). The respiratory system is of the tubocanaliculate morphotype, with more or less straight channels. The abovementioned characteristics allow us to refer the eggshells to the oospecies Megaloolithus mamillare Vianey-Liaud et al. 1994. This differs from the oospecies M. siruguei Vianey-Liaud et al. 1994 in the lack of transversal channels and in the size proportions of the shell units (wider than high in M. mamillare), and from M. baghensis Vianey-Liaud, Hirsch, Sahni, & Sig, 1997 in the surface ornamentation and in the pattern of growth lines in the shell units. 5. Discussion 5.1. Correlation with the Geologic Time Scale Polarity reversals and biochronological markers make it possible to correlate the obtained magnetozones in the Upper Aude Valley with the Standard Geologic Time Scale (GTS) (Ogg & Hinnov, 2012). The N1 magnetozone (from the base of the section up to meter 46) includes the oological association of Cairanoolithus roussetensis and Cairanoolithus dughii, whereas the R1 magnetozone (meters 46 to 67) includes C. dughii only at its very base (Fig. 8). According to Garcia and Vianey-Liaud (2001), both Cairanoolithus oospecies occur in the Rousset-La Cairanne section (Arc Basin, Provence) in chron C32n according to the magnetostratigraphic data from this section (Westphal & Durand, 1990). Consequently, as a first approach, the N1-R1 transition can be correlated either with the C32n.2n-C32n.1r or the C32n.1n-C31r reversal (Figs. 4 and 8). Considering again the eggshell biostratigraphy from the Rousset-La Cairanne section, C. dughii disappeared before C32n.1n, but data
res) (Galbrun, 1997; Garcia & Vianeyfrom the Albas section (Corbie Liaud, 2001) indicate that this oospecies may also be present in that chron. Hence, without a more conclusive biochronological marker, our preferred interpretation correlates the N1-R1 reversal at about meter 46 with C32n.1n-C31r, and N1 is ascribed entirely to C32n.1n (Fig. 8). The possibility that the short interval of reverse polarity at about 40 m represents C32n.1r and hence that N1 also includes C32n.2n appears less likely. Conversely, the alternative correlation of the N1-R1 reversal to C32n.2n-C32n.1r would also be feasible, but less parsimonious. According to Ogg and Hinnov (2012), the CampanianMaastrichtian boundary is placed close to the reversal between C32n.2n and C32n.1r. Hence, its position in the Upper Aude Valley

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Fig. 7. New eggshell material from the Upper Aude Valley localities. A, SEM image of Cairanoolithus dughii eggshell from the Gourg de l’Encantado site (MDE-D-386); A′, radial thin section under transmitted light microscope of the same sample. B, SEM image of Megaloolithus mamillare eggshell from level 4 of the Roque Fumade site (MDE-D-392), arrow indicates pore channel; B′, radial thin section under transmitted light microscope of the same sample; arrow indicates growth lines. Scale bars in A and B equal to 200 mm.

should be located below the sampled section, towards the middle or the base of the Red Marls of Campagne or even within the underlying Alet Sandstone. The lack of good exposures around meter 67 to 91 results in a thick gap with no paleomagnetic data. Even though the paleomagnetic signal obtained between meters 91 and 97 was not conclusive, we found a Megaloolithus siruguei succession overlaid by an occurrence of the oospecies Megaloolithus mamillare there. The R2 magnetozone occurs just above these eggshell sites (meters 97 to 122). Thus, we correlate this polarity interval with chron C31r on the basis of the fact that: (1) the replacement between these megaloolithid oospecies seems to occur in chron C31r (Fig. 8) (Vila et al., 2011; Tabuce et al., 2013) and this makes it plausible that the overlying polarity interval represents the same chron, in the absence of any evidence of hiatus or erosion; (2) the reported charophyte species from the lower levels of the Vignevieille Limestones (meters 126 to 130) are compatible with chron
re et al. C31r but not with the younger C29r. For instance, Bessie (1980, 1989) cited Septorella ultima, which disappeared at the beginning of the late Maastrichtian (Galbrun, Feist, Colombo, Rocchia, & Tambareau, 1993; Riveline et al., 1996). Similarly, Marty (2001) found Peckichara cf. caperata and Microchara punctata, both species belonging to non-terminal Maastrichtian deposits in other Pyrenean areas (Feist & Colombo, 1983; Vicente et al., 2015). The boundary between the lower and the upper Maastrichtian, located in the upper half of C31r at about 70Ma in GTS2012 (Ogg & Hinnov, 2012), cannot be precisely pinpointed, but the occurrence of M. mamillare seems to indicate that R2 corresponds to the upper part of chron C31r. If the alternative attribution of C29r for R2 were valid, the above-mentioned eggshell replacement would imply a hiatus below R2 that comprises the totality of C31n, C30r and C30n.

However, there is no field observation of a great depositional interruption where this hypothetical hiatus would be. Regarding the K-Pg boundary, Marty (2001) and Marty and Meyer (2006) placed the transition towards the top of the Vignevieille Limestones. According to our interpretation, most of the upper Maastrichtian is extremely reduced within the Vignevieille Limestones or is not recorded in the Upper Aude Valley (hiatus). The short undetermined polarity interval found in the lower few meters of the limestones would only allow for an eventual reduced upper Maastrichtian interval. Still, and contrary to the alternative interpretation discussed above, here the tangible karstification of these limestones is in support of the hiatus scenario. In summary, the age of the studied area (from the upper part of the Red Marls of Campagne to the Vignevieille Limestones) ranges from C32n to C31r. The upper Maastrichtian (C31n to C29r) could be recorded entirely in the limestones or it could be absent. These chronostratigraphic constraints are in agreement with the original interpretations of Bilotte (1978, 1985) and Bilotte et al. (1983) and those reported on the basis of previous eggshell biostratigraphy (Garcia & Vianey-Liaud, 2001). 5.2. Integration of the Upper Aude Valley chronostratigraphy within the framework of southwestern Europe The magnetostratigraphic and sedimentological studies carried out in the last few years in the Upper Cretaceous of the southern Pyrenees have allowed a well-dated geological framework to be built for this area. Now, our results permit a general analysis of the marine to transitional-continental formations of southwestern Europe. The fluvial succession we propose for this north Pyrenean sector is in agreement with Bilotte (1978, 1985) and Bilotte et al. (1983),

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Fig. 8. Eggshell biostratigraphy, magnetostratigraphy and biochronological markers in the Upper Aude Valley composite section, and correlation with the Geomagnetic Polarity Time Scale (Ogg & Hinnov, 2012). See the preferred and alternative magnetostratigraphic correlations discussed in text. Dinosaur eggshell biostratigraphy after Garcia and Vianeys et al. (2013) and Selle s and Vila (2015). Dinosaur eggshell occurrence in the Upper Aude Valley after Garcia (1998), Vianey-Liaud and Garcia Liaud (2001), Vila et al. (2011), Selle
re et al. (1980, 1989), Marty (2001) and Marty and Meyer (2006). Sites: 1: Rennes-le(2000), Garcia and Vianey-Liaud (2001) and this study. Charophyte occurrence after Bessie raza and F5. Ch^ ateau; 2: Bellevue; 3: Gourg de l'Encantado; 4: Roque Fumade 1e5; 5: approximate location of Ve

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and exhibits similar lithofacies to those from the southern Pyrenees. However, regarding the red bed units, differences in accommodation and possibly sedimentary hiatuses during the late Maastrichtian resulted in a remarkable contrast in thickness between the northern and southern areas: up to 500 m in the Vallcebre section (Oms et al., 2007) in the south, and just 130e150 m in the Upper Aude Valley in the north. Further, our paleomagnetic data indicate that the marine to transitional-continental environments occurred a few million years earlier in the Upper Aude Valley. Thus, whereas the lagoonal setting of the ‘grey unit’ of the Tremp Fm (Nagtegaal, Vanvliet, & Brouwer, 1983; Cuevas, 1992; Rosell et al., 2001; Riera et al., 2009, 2010) dominated the southern area during the late Campanian-early Maastrichtian (Fig. 9A) (Oms et al.,
vol, Lo  pez-Martínez, & Arribas, 2007; Riera et al., 2009; Vicens, Arde 2004; Villalba-Breva & Martín-Closas, 2012), a terrestrial paleoenvironment (braided-meandering streams and their associated overbank deposits) is recorded in the Upper Aude Valley during a similar age range (Fig. 9C). Alluvial and deltaic red beds (the ‘lower red unit’ of the Tremp Fm, sensu Rosell et al., 2001) do not appear in

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the south Pyrenees until the mid-part of the early Maastrichtian
(Oms et al., 2007; Vila et al., 2012), with the exception of the Ager syncline (Fig. 9B) (Colombo & Cuevas, 1993; Galbrun et al., 1993; pez-Martínez, Arde
vol, Arribas, Civis, & Gonza lez-Delgado, Lo 1998; Villalba-Breva & Martín-Closas, 2012). However, from this moment onwards, similar red bed facies occur on both sides of the Pyrenees. These facies evolve to paralic or lagoonal paleoenvironments westwards, represented by the Auzas Marls (Haute Garonne) and the ‘grey unit’ in the northern and southern Pyrenean areas, respectively (Bilotte et al., 1983; Riera et al., 2009, 2010). Eastward from the Pyrenean range, a thick continental deposition has been reported from the upper Santonian to the Paleogene in the Arc syncline (Provence). There, during the late Campanian and the Maastrichtian, palustrine-lacustrine and fluvial environments characterize the local Rognacian (Fig. 9D) (Babinot & Durand, 1980a, 1980b; Durand, Gaviglio, Gonzales, & Monteau, 1985). In the latest Maastrichtian interval, the south Pyrenean area together with the Arc syncline exhibit a similar coarsening and regressive trend (Fig. 9A and D), culminating in the development of proximal


n/Areny, Isona, Vallcebre and Fontllonga sections (Tremp, Vallcebre and Ager Fig. 9. Paleoenvironmental succession of southwestern European regions. AeB, Campo, Are synclines, southern Pyrenees). C, Upper Aude Valley (northern Pyrenees). D, Arc Basin (Provence). Litho-bio-magnetostratigraphic data from the following references: Arc Basin after Westphal
pez-Martínez et al. (2001) and Canudo et al. (2015); Isona and Durand (1990); Vallcebre syncline after Oms et al. (2007); Ager syncline after Galbrun et al. (1993); Campo after Lo n/Areny after Pereda-Suberbiola et al. (2009). after Vila et al. (2012); Are

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alluvial systems during C29r, located a few meters below the K-Pg boundary (Reptile Sandstone and Galante Conglomerate units, respectively) (Oms et al., 2007; Westphal & Durand, 1990). The present correlation places the Fleuri Conglomerate unit of the Upper Aude Valley within C31r, it being older than the former conglomerate and sandstone units. A major climatic change before the mass extinction event has been proposed as the origin of this modification in the floodplain paleoenvironments at the very end of the Cretaceous, since there is no tectonic event described for this
interval (Oms et al., 2007). The Ager syncline also exhibits a particular evolution (Galbrun et al., 1993), recording coarse deposits earlier, in C30n (Fig. 9B). Finally, the red bed facies shift diachronically into a palustrine-lacustrine paleoenvironment in all southwestern European areas: before the K-Pg boundary in the case of the Vignevieille Limestones in the Upper Aude Valley (Marty, 2001; Marty & Meyer, 2006), or after the boundary in the southern Pyrenees and Provence, where the lacustrine facies are recorded as the Vallcebre Limestones and laterally equivalent strata and  pezthe Vitrolles Limestone, respectively (Cojan et al., 2000; Lo
vol, 2006; Vila et al., Martínez, Arribas, Robador, Vicens, & Arde 2013 and references therein). In summary, the Upper Cretaceous of the whole Pyrenean region exhibits a gradual marine regression towards the west, starting as early as the Campanian in the north and reaching the south in the early Maastrichtian. In this latter area, alluvial and deltaic red beds were the main paleoenvironments during the late Maastrichtian, attaining a maximum regression peak that resulted in the development of coarse-grained sandstones and conglomerates very close to the K-Pg boundary, in a similar way to what occurred in the nearby Provence area. By contrast, the upper Maastrichtian of the Upper Aude Valley seems to comprise the lacustrine-palustrine environments represented by the karstified Vignevieille Limestones (i.e. very reduced) or not be deposited at all. 5.3. The dinosaur record of the Ibero-Armorican domain around the Campanian-Maastrichtian boundary The combination of nine stratigraphic sections with the paleomagnetic data results in a precise succession of dinosaur and other vertebrate fossil sites in the Upper Aude Valley near the villages of ^teau (Aude Department) Campagne-sur-Aude and Rennes-le-Cha (Fig. 10). The succession is characterized by: (1) the absence of vertebrate and eggshell remains in the Red Marls of Campagne, and their occurrence as debris deposits in the Estous Sandstone; (2) the abundance of bone and eggshell-bearing sites towards the base of the Red Marls of Maurine during C32n (early Maastrichtian), close to the reversal between C32n.1n and C31r; this corresponds to an age around 71.5 Ma for these lower sites; (3) the scarcity of fossil sites in the upper part of the latter unit, with the exception of the Roque Fumade section, where six eggshell-bearing sites are located. The paleontological content of twenty-one revised sites is summarized in Supplementary Information 2. Regarding the eggshell material, the integration of all the parataxonomic data (Garcia, 1998; Vianey-Liaud & Garcia, 2000; and present review) allows us to summarize the dinosaur oological record of the Upper Aude Valley as follows: Bellevue (M. sp., ^teau (M. siruguei, C. dughii, C. dughii, C. roussetensis), Rennes-le-Cha C. roussetensis), La Valdieu (C. dughii, C. roussetensis), Arques raza (M. baghensis), Campagne-sur-Aude (M. aureliensis), Ve (C. dughii), Gourg de l'Encantado (C. dughii), Les Boudous (C. dughii) and Roque Fumade (levels 1, 2, 3, and 5: M. siruguei; level 4: M. mamillare). Garcia and Vianey-Liaud (2001, Fig. 3) reported a
res-Aude Valley succession of dinosaur oospecies from the Corbie area; in spite of the fact that the authors did not provide the names of the eggshell-bearing sites in the section, we assume that they

correspond, from bottom to top, to the sites of Arques, Bellevue, ^teau, Roque Fumade and Ve raza because of their Rennes-le-Cha stratigraphic position and the oospecies represented there. This succession concurs with the oospecies observed in our review of these sites. As regards the dinosaur remains recovered from the sites of Bellevue, Gourg de l’Encantado, La Valdieu, Campagne-sur-Aude ^teau, the main taxonomic groups correspond and Rennes-le-Cha to titanosaurid sauropods such as Ampelosaurus atacis, the rhabdodontid Rhabdodon priscus, nodosaurid ankylosaurs and dromaeosaurid theropods (Le Loeuff, 2005; Le Loeuff et al., 1994b, 1997). These belong to the late Campanian-early Maastrichtian titanosaurid-rhabdodontid-dominated assemblage that Le Loeuff et al. (1994a) contrasted with the hadrosauroid-dominated faunal assemblage present in the late Maastrichtian of the IberoArmorican domain. Hence, these authors proposed a faunal turnover around the early-late Maastrichtian boundary, in which the former assemblage was replaced by hadrosauroids. The lack of a thick upper Maastrichtian record in the studied area could explain the complete absence of hadrosauroid remains throughout the succession. Several titanosaurids have been described for the upper Campanian-lower Maastrichtian in the localities of the IberoArmorican domain. Hence, in addition to Ampelosaurus atacis from Bellevue, another titanosaurid form is waiting to be described in Massecaps, in the northern Pyrenees (Díez Díaz, Tortosa, & Le Loeuff, 2013). In Provence, the species Atsinganosaurus velauciensis has been reported from Velaux-La Bastide Neuve (Garcia, Amico, Fournier, Thouand, & Valentin, 2010); two different morphotypes come from Fox-Amphoux (Díez Díaz et al., 2012); and other titanosaur remains belonging very likely to distinct taxa are still under study (Tortosa, Dutour, Cheylan, & Buffetaut, 2012). Likewise, the Iberian Peninsula yields a rich titanosaur record for the upper Campanian-lower Maastrichtian transition. Thus, the species Lirainosaurus astibiae has a broad distribution, being pre~ o in the Basque Country (Sanz, Powell, Le Loeuff, sent in Lan
ncia Martínez, & Pereda Suberbiola, 1999), Chera in Vale ~ aca, 2009) and Sacedo n (Company, Pereda Suberbiola, & Ruiz-Omen rez-García, 2009). In Lo Hueco (Cuenca, in Cuenca (Ortega & Pe Spain), the titanosaurid remains are still under study, but up to two different forms have been reported, one of them being regarded as close to the genus Ampelosaurus (Knoll, Ridgely, Ortega, Sanz, & Witmer, 2013; Díez Díaz, Ortega, & Sanz, 2014). Our chronostratigraphic framework for the dinosaur faunas of the Upper Aude Valley is a step towards understanding the diversity and changes that occurred during the late Campanian-early Maastrichtian interval in southwestern Europe, especially with regard to the titanosaurid assemblage. Until now, only Atsinganosaurus velauciensis and the other remains from Provence could be integrated within a chronostratigraphic framework. Thus, according to data from Tortosa et al. (2014) and Tortosa (pers. comm.), Atsinganosaurus velauciensis was found in the lower part of chron C33n, whereas the other Provence remains are located in its upper part. As our results show, these occurrences are not synchronous with Ampelosaurus atacis. Therefore, distinct ages can explain the titanosaurid faunal differences. Because the differences (or similarities) between the titanosaurid remains of the above-mentioned localities can be linked to possible differences (or coincidences) in the age of the sites, we focus on the importance of achieving good chronostratigraphic resolution in order to draw faunal comparisons between localities, and especially to attempt to establish the faunal succession. The marine-to-continental lagoonal facies of the lower Maastrichtian of the Tremp Fm have yielded very few bone sites, conferring an apparently low dinosaur diversity on this period. By

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Fig. 10. EeW correlation framework for the Upper Aude Valley sections and fossil sites. The location of the sections is shown in Fig. 2. The correlation lines correspond to the major lithostratigraphic units of Bilotte (1978, 1985) and Bilotte et al. (1983). Paleontological data appear in Supplementary Information 2. Arrows indicate aproximated or projected ^teau; g, La Valdieu; h, Les Boudous; i, Bellevue; j, Campagne-sur-Aude; k, C2; l, Gourg de locations. Sites: a, Arques; b, ‘Bellevue road’; c, E1; d, F1 AeB; e, F2 AeB; f, Rennes-le-Cha raza. l’Encantado; m, Les Labadous; n, Roque Fumade level 1; o, Roque Fumade level 2; p, Roque Fumade level 3; q, Roque Fumade level 5; r, Roque Fumade level 4; s, F5; t, Ve

contrast, the northern Red Marls of Maurine in the Upper Aude Valley exhibit a rich fossil record in an equivalent age, as explained above. This asymmetry in the fossil record observed between the north and south Pyrenean areas may have been strongly influenced

by paleoenvironments. However, even though very few dinosaur bone remains have been discovered in the lower Maastrichtian of the Tremp and Vallcebre synclines, the evidence for dinosaurs is widely present, often in the form of eggs and footprints.

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Accordingly, Le Loeuff and Martínez (1997) suggested that the titanosaurid assemblage found in Bellevue was the possible producer of the Fumanya trackways (Vallcebre syncline) on the basis of their age similarities. Since the south Pyrenean site is dated as C32n.1n (early Maastrichtian; Oms et al., 2007), the present magnetostratigraphy of the Upper Aude Valley succession confirms the age equivalence between the two titanosaurid sites. Moreover, whereas ichnological evidence of titanosaurids has been found in the lagoonal facies of the Vallcebre and Isona basins, no rhabdodontid remains have been reported there (Riera et al., 2009). This could be evidence of the habitat preference of this dinosaur group for continental paleoenvironments. The fact that rhabdodontid remains have been reported from the lower Maastrichtian alluvial deposits of Perauba-Figuerola (Llompart & Krauss, 1982), as well as the Upper Aude Valley, supports this idea. However, more evidence and further statistical analysis are required to substantiate these possible paleoenvironmental preferences. 6. Conclusions The comprehensive lithostratigraphic study of the Upper Cretaceous continental red beds of the Upper Aude Valley (southern France) makes it possible to correlate several sections with reference dinosaur-bearing sites, resulting in the first study of the stratigraphic and dinosaur succession performed in this area. In addition, we obtained a magnetostratigraphic scheme constrained by biochronological markers such as dinosaur eggshells and charophytes, which can be correlated with the standard GTS2012. Accordingly, the top of the Red Marls of Campagne, the Estous Sandstone and the lower part of the Red Marls of Maurine belong to chron C32n, probably the uppermost part of C32n.1n, whereas the rest of the latter unit can be correlated with chron C31r. Overlying this, the karstified Vignevieille Limestones record the K-Pg transition. In light of these results, the following concluding remarks can be made:  The end-Cretaceous continental record of the studied area is mostly restricted to the lower Maastrichtian (from C32n to C31r). The upper Maastrichtian is represented by a short interval along the Vignevieille Limestones (a few meters) or is completely absent.  A marine to continental transition took place a few million years earlier in the northern Pyrenean basin than in the southern part. During chron C32n, the Upper Aude Valley records fluvial environments, whereas the southern Tremp and Vallcebre synclines display marine-to-lagoonal settings during the coeval interval.  The Fleuri Conglomerate, here correlated with C31r, is not equivalent to other coarse units described in southwestern Europe (e.g. the Reptile Sandstone of the Vallcebre syncline and the Galante Conglomerate of Provence, both dated as C29r), as has previously been discussed.  The majority of the dinosaur fossil sites in the studied area are found in the lower levels of the succession. Their dinosaur faunal content is represented mainly by titanosaurids and rhabdodontids, and to a lesser extent, nodosaurid ankylosaurs and theropods.  The age of the Bellevue site is constrained between 71.5 Ma and 72 Ma. In light of the present data, the preferred age correlation would be 71.5 Ma (top of C32n.1n). The integrated stratigraphy presented in this paper is a substantial addition to our knowledge of the temporal distribution of the last dinosaur faunas that inhabited the European archipelago, especially those of the early Maastrichtian.

Acknowledgments This paper is a contribution to the projects CGL2011-30069-C0201, 02/BTE and CGL2010-16447, funded by the Ministerio de Economía y Competitividad of Spain. V. Fondevilla acknowledges support from the Ministerio de Economía y Competitividad, (FPI grant, BES-2012-052366). B. Vila acknowledges support from the  n (Subprograma Juan de la Cierva Ministerio de Ciencia e Innovacio (MICINN-JDC) 2011). We thank A. Blanco for his field assistance during the campaign of October 2013. We are grateful to Dr. Emilio Pueyo, Dr. Thierry Tortosa and the editor (Dr. Eduardo Koutsoukos) who provided useful comments to improve the manuscript. Thanks are due to Rupert Glasgow for his English grammar corrections. References lien, Be gudien, Rognacien, Babinot, J. F., & Durand, J. P. (1980a). Valdonien, Fuve tages français et leurs stratotypes. Orle ans, France. Bureau de Vitrollien. Les e Recherche G eologique et Mini
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Appendix A. Supplementary data Supplementary data related to this article can be found at http://dx.doi.org/10. 1016/j.cretres.2015.08.009.

Please cite this article in press as: Fondevilla, V., et al., Magnetostratigraphy of the Maastrichtian continental record in the Upper Aude Valley (northern Pyrenees, France): Placing age constraints on the succession of dinosaur-bearing sites, Cretaceous Research (2015), http://dx.doi.org/ 10.1016/j.cretres.2015.08.009