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Re-evaluation of the age of some dinosaur localities from the southern Pyrenees by means of megaloolithid oospecies.

1. Introduction

Areas of the present-day Pyrenees (southern France and north-eastern Spain) contain localities ranging from the late Campanian to the latest Maastrichtian, providing a productive succession of dinosaur fossil remains from the latest Cretaceous (Buffetaut and Le Loeuff, 1991; Lopez-Martinez et al., 2001; Laurent et al., 2002; Riera et al., 2009). In the last century more than two hundred localities have been discovered in the southern Pyrenees. Mr. Bartolome Castell made the first isolated findings in 1927 in the vicinity of Tremp (Marin and Bataller, 1929); these were followed by further prospects and geological field characterizations that yielded additional material from localities such as "Orcau", "Suterrana" and "Bastus" (Lapparent and Aguirre, 1956; Lapparent, 1958). In the early 1980s and 1990s more localities were described, including significant bone, egg and track localities such as Els Nerets, Sant Roma d'Abella, Fumanya and Fontllonga (Casanovas-Cladellas and Santafe-Llopis, 1993; Casanovas-Cladellas et al., 1985, 1993; Le Loeuff and Martinez, 1997; Lopez-Martinez et al., 1998). Since then, various authors have compiled the fossil localities containing dinosaurs (Lopez-Martinez, 2003; Vila et al., 2006; Suner et al., 2008, Riera et al., 2009), and at present the total number of localities is about 220. The stratigraphic age for these continental dinosaur-bearing localities has traditionally been provided by stratigraphic correlation with marine series (Ardevol et al., 2000) and magnetostratigraphy (Galbrun et al., 1993), together with the use of various biostratigraphic markers (charophytes, palynomorphs, rudists; Feist and Colombo, 1983; Medus et al., 1992; Riveline et al., 1996; Vicens et al., 2004). In the last decade, the profusion of new discoveries and the advances in magnetostratigraphy and biostratigraphy in the southern Pyrenees (Lopez-Martinez et al., 2001; Oms et al., 2007; Pereda-Suberbiola et al., 2009; Riera et al., 2009; Prieto-Marquez et al., 2013; Vila et al., 2012, 2013) have allowed a more precise and robust correlation for these localities. Of special interest is the calibration of the succession of megaloolithid oospecies with the magnetostratigraphic scale (Garcia and Vianey-Liaud, 2001; Vila et al., 2011).

The aim of the present work is to re-evaluate the age of 20 historically significant localities by means of the use of oospecies and other biochronostratigraphic criteria and to provide age and parataxonomic data for 23 localities that yield new megaloolithid egg material. Further, we provide a review of the nomenclature and probable synonymies of 18 of the dinosaur localities in the southern Pyrenees.

2. Geological setting

The southern Pyrenees (NE Iberian Peninsula; Fig. 1) contain a 3000-m-thick succession of sedimentary rocks encompassing the transition from shallow marine carbonate platforms to fully continental environments (Riera et al., 2009 and references therein). Two main geological units of latest Cretaceous age (late Campanian to late Maastrichtian) are recognized in the area: the Aren Sandstone Fm. and the Tremp Fm. The Aren Sandstone Fm. is composed of sandstones deposited in beach, barrier-island and deltaic systems; it passes gradually to the Tremp Fm. by a diachronic interfingering of strata (Diaz-Molina, 1987). The Tremp Formation has been divided informally into four lithologic units (Rosell et al., 2001): 1) the "grey unit", 2) the "lower red unit", 3) the "Vallcebre Limestones and laterally equivalent strata", and 4) the "upper red unit".

The localities discussed in the present work are situated in distinct areas of the provinces of Barcelona and Lleida; from east to west these are: the Vallcebre, Coll de Nargo, Tremp and Ager synclines. In these areas, the deposits of late Campanian to latest Maastrichtian age have yielded a diverse and abundant tetrapod fossil record that comprises dinosaurs (sauropods, theropods, ornithopods and ankylosaurs; LopezMartinez et al., 2001; Riera et al., 2009; Pereda-Suberbiola et al., 2009; Cruzado-Caballero et al., 2010; Vila et al., 2012; Prieto-Marquez et al., 2013; Torices et al., in press), pterosaurs (Dalla Vecchia et al., 2013), crocodilians (Puertolas et al., 2011; Puertolas-Pascual et al., 2014), turtles (Marmi et al., 2012), and amphibians and squamate reptiles (Blain et al., 2010).

3. Methods and materials

We have reviewed the names of 18 south-Pyrenean localities and standardized with the current toponymy, correcting previous misspellings and providing name equivalences. When possible, the criterion followed for assigning the name to a locality has been its primary citation in the literature. Many localities have received different names after their first being reported and in cases where the secondary name has been cited more frequently in the literature, we have chosen to maintain the more extensively used name. If, after the review and name standardization, the locality name coincides with a previous naming we have added numerals in order to avoid misunderstandings. Regarding terminology, we used "locality" to refer a discrete area or stratigraphic section where fossils occur and which may or may not include various sites. A "site" refers to a discrete fossil-bearing level within a locality.

The new eggshell fragments described in the present study (Appendix 1) were collected from both classic and unreported localities in the southern Pyrenees (Fig. 1). These fossil remains were treated with potassium hydroxide (KOH) and/or sodium hexametaphosphate (SHMP) 30%, and cleaned in an ultrasonic bath (Val et al., 2014). Samples were preliminarily analyzed using the Leica[R] M60 binocular, and structural characters (e.g. shell thickness, ornamental pattern, pore aperture width) were measured using Leica[R] Application Suite 2.8.1 software. Several samples were prepared as standard thin sections (30 [micro]m-thick) while others were examined and photographed using the electronic microscope ESEM Quanta 200 FEI, XTE 325/D8395, in the Department of Scientific-Technical Services of the Universitat de Barcelona. Eggshells were identified in terms of parataxonomy following Mikhailov (1997).

The studied material is housed in the collection of the Institut Catala de Paleontologia (IPS-58959, 58960, 58963 to 58965, 58967, 58968, 58975, 58982, 58987, 58992, 59132 to 59135, 59137, 82173 to 82180, 82182 to 82227, 82230 to 82270).

Institutional abbreviation: IPS, Institut de Paleontologia de Sabadell.

4. Results

4.1. Historical review of the nomenclature and synonymies of some dinosaur localities

Since the earliest dinosaur discoveries in the southern Pyrenees various authors have published data on particular localities, sometimes generating a number of different names or spellings for the same locality. It is out of the scope of the present work to review the nomenclature of all the southern Pyrenean localities, but in order to avoid further misunderstandings in some of the cases, we here discuss and standardize their name and synonymy (Table 1).

Several dinosaur remains were recovered in the area around the villages of Talarn, Suterranya and Orcau (Tremp Basin). Lapparent and Aguirre (1956) reported various fossil localities in the basal levels of the "grey unit" of the Tremp Formation in this area (Pereda-Suberbiola and Ruiz-Omenaca, 2012). The Presa de San Antonio locality reported by Lapparent and Aguirre (1956, p. 379) is situated near the village of Talarn and is probably equivalent to Sant Antoni-2 referred to Ardevol et al. (1995), which was later named Presa de Tremp by Vila et al. (2012). The locality may correspond to that described by Bartolome Castell in 1927 (Marin and Bataller, 1929). Southwards, the Norets locality referred to Pereda-Suberbiola et al. (2003) is equivalent to the Point 2 of Lapparent and Aguirre (1956, p. 380). The Els Nerets locality, originally referred to Casanovas-Cladellas et al. (1985), is equivalent to Vilamitjana-1 (V-1) of Ardevol et al. (1995). It is also worth noting that the eggshell locality Vicari referred to by Moratalla (1993, 1998) and Lopez-Martinez (2000) may be equivalent to the Vicari-4 locality of Torices Hernandez (2002) and Torices et al. (2004, 2012).

In the vicinity of the village of Suterranya, up to four localities were distinguished. The first locality, here named "Suterranya-Cami de Montesquiu" (Point 3 of Lapparent and Aguirre, 1956, p. 380), was discovered by Walter Georg Kuhne in 1954 and yielded several dinosaur bones of indeterminate affinity. The second locality, here named "Suterranya-Mina de lignit" (Point 4 of Lapparent and Aguirre, 1956, p. 380), was discovered by Josep Montane and yielded mainly fragmentary dinosaur bones currently housed in the Museo Nacional de Ciencias Naturales (MNCN) in Madrid. After conversations with Josep Montane and revision of the MNCN collections, we could claim that this latter locality probably yielded the ankylosaur bones reported by Escaso et al. (2010) as well as other fragmentary bones of sauropod affinity. In addition, Moratalla (1993) reported dinosaur eggshells from the Suterranya locality (originally spelled "Suterrana"); finally, Ardevol et al. (1995, 1999) reported the egg locality of Suterranya-1, which is a synonym of the L'Abeller locality reported by Prieto-Marquez et al. (2000) and "Suterrana" of Torices et al. (in press); this also yielded an isolated theropod tooth.

At least five dinosaur localities have been reported in the vicinity of the village of Orcau. Lapparent and Aguirre (1956) distinguished four localities with dinosaur remains, the most important being that of "Orcau" discovered by W.G. Kuhne in 1954 and containing various sauropod bones. Llompart et al. (1984) described dinosaur tracks east of the village of Orcau. Ardevol et al. (1995) renamed the bone and track localities as Orcau-1 and Orcau-2, respectively. Riera et al. (2009) and Vila et al. (2013) followed the same procedure for naming the localities of Orcau-3 (bones) and Orcau-4 (tracks), respectively. Finally, Moratalla (1993, 1998) reported dinosaur eggshells from a locality called Orcau.

At the southern margin of the Tremp Basin, Brinkmann (1984) described dinosaur remains in the Moror locality (misspelled "Moro"); some years later, Vianey-Liaud and Lopez-Martinez (1997) and subsequent works (Lopez-Martinez, 2000; Bravo and Marugan-Lobon, 2013) referred to an egg locality as "Moro" (we here correct the misspelling and change the name to Moror-1 for the sake of distinction). Lopez-Martinez and Vicens (2012) also described the Serrat Pedregos egg locality, which is equivalent to the Cellers-2 locality referred to by the same authors some years before (Lopez-Martinez, 2000). The latter author also reported the Montrebei eggshell and tooth locality (misspelled "Montrebey" by Torices Hernandez, 2002).

In the Ager syncline, various authors reported dinosaur remains from at least three localities with similar names. The first locality reported in the literature was "Fontllonga" (Casanovas-Cladellas and Santafe-Llopis, 1993), which is the same locality later reported by Vianey-Liaud and LopezMartinez (1997) and subsequent works as Fontllonga-6. In the same year Moratalla (1993) gave the name "Fontllonga" (later named "Fontllonga L"; Moratalla, 1998) to a locality with dinosaur eggshells. Another locality in the area, also named "Fontllonga" (Casanovas et al., 1999) and yielding a hadrosaur dentary, is a synonym of "Fontllonga R" of LopezMartinez (2003).

In the Coll de Nargo syncline, Lopez-Martinez (2000) reported "several superposed levels with scattered clutches" in the area of "Santa Eularia", which probably correspond to the sites of Santa Eulalia-1 to -4 (we have corrected the misspelling), sampled in the present work (Appendix 1).

4.2. Parataxonomical study

The parataxonomical study of dinosaur eggshells collected in new and classical localities of the southern Pyrenees allows them to be attributed to four megaloolithid oospecies: Megaloolithus aureliensis, M. siruguei, M. mamillare and M. baghensis (see Appendix 2 for further data and discussion). The studied sample includes 2738 eggshell fragments from in situ complete eggs, eggshell accumulations (egg debris) and scattered eggshells (Appendix 1). Most eggshells occur in overbank deposits associated with fluvial or lagoonal systems, in grey to reddish mudstones and marls, or in limestones associated with lacustrine settings.

5. Discussion

5.1 Review and update of the age of some dinosaur localities

Some authors have argued that dinosaur eggshells can be used as biochronological markers in continental deposits (Garcia and Vianey-Liaud, 2001). This idea, originally proposed for the Aix Basin in Provence (France), has been successfully exported to the southern Pyrenees regions of Vallcebre and Coll de Nargo (Vila et al., 2011; Selles et al., 2013). The oospecies assemblages (properly named oozones, see Selles et al., 2013) are calibrated with magnetochrons and indicate restricted temporal ranges for the megaloolithid oospecies (Fig. 2). For instance, the dominance of M. siruguei (together with the oospecies M. aureliensis and Cairanoolithus) mainly indicates a late Campanian to early Maastrichtian age; its replacement with Megaloolithus mamillare occurs around the C31r-C31n polarity change; and the appearance of M. mamillare and/or M. baghensis (formerly M. pseudomamillare, Vianey-Liaud et al., 2003) indicates a late Maastrichtian age (Garcia and Vianey-Liaud, 2001; Vila et al., 2011; Selles et al., 2013).

In the present work, the use of oozones based on megaloolithid oospecies enables the re-evaluation of the age of several localities (Fig. 2A). In the eastern part of the Tremp Basin, the Biscarri egg locality yielded the oospecies Megaloolithus siruguei and was originally referred to the late Campanian (Lopez-Martinez et al., 2000). This oospecies is represented in Oozone 1, which is of late Campanian-early Maastrichtian age (Garcia and Vianey-Liaud, 2001), but also on its own in Oozone 2, of early Maastrichtian age (Garcia and Vianey-Liaud, 2001; Vila et al., 2011). The new magnetostratigraphic correlations of lower Tremp Formation units in the Tremp Basin (Vila et al., 2012) indicate that the basal part of the formation is probably early Maastrichtian in age and thus the Biscarri locality (together with the Barranc de la Fonguera locality; Appendix 1) would be early Maastrichtian in age.

At the north-western margin of the Tremp Basin the age of the localities of Basturs, Orcau-1, Els Nerets, Orcau, Llabusta, Vicari, Compuertas, Suterranya, Suterranya-1, and Abella has been re-evaluated (Appendix 1). Basturs (originally spelled "Bastus" by Lapparent, 1958) was the first egg locality to be discovered in the southern Pyrenees. This nesting area refers to various egg levels (the egg-bearing sites of Basturs-1, Basturs- 2, Basturs-4, and Basturs-5; Ardevol et al., 1999) and was considered early Danian by Lapparent (1958), Maastrichtian by Moratalla (1993), "upper Rognacian" by Vianey-Liaud and Lopez-Martinez (1997), and late Campanian by Lopez-Martinez (2000) and Diaz-Molina et al. (2007). Moratalla (1993, 1998) and Vianey-Liaud and Lopez-Martinez (1997) reported the oospecies M. mamillare in the Basturs-1 and Basturs-2 sites, and therefore they are here considered late Maastrichtian in age.

At the end of 1954 W.G. Kuhne discovered the bone locality of Orcau-1 (originally named "Orcau" by Lapparent and Aguirre, 1956). These authors and Casanovas et al. (1987) referred the age of the locality to the Maastrichtian. Casanovas-Cladellas et al. (1985) reported the locality of Els Nerets (Tremp, Lleida), which was considered Maastrichtian in age. Lopez-Martinez (2000) dated it as late Campanian, and most recently Dalla Vecchia et al. (2014) have suggested a late Maastrichtian age on the basis of the stratigraphic correlation of Riera et al. (2009) but with no further biostratigraphic constraints. Recent prospects and excavations in the Orcau-1 and Els Nerets localities indicate that they bear Megaloolithus baghensis and M. mamillare eggshells, respectively (Appendices 1 and 2), hence supporting a late Maastrichtian age.

Moratalla (1993, 1998) reported the oospecies Megaloolithus cf. mamillare and/or M. trempii in the localities of Orcau, Llabusta, Vicari, Compuertas, Fontllonga L and Suterranya, and Vianey-Liaud and Lopez-Martinez (1997) identified the oospecies M. pseudomamillare and M. mamillare oospecies in the Suterranya-1 and Abella localities, respectively. Moratalla (1993) considered the eggshell localities of Orcau, Llabusta, Vicari, Compuertas and Suterranya to be Maastrichtian in age, whereas Lopez-Martinez (2000) suggested a latest Campanian age for these as well as for the Suterranya-1 and Abella localities. In the case of the Vicari locality, Torices et al. (2012) indicated that the Vicari section encompasses the latest Campanian but also the early Maastrichtian. The authors also located the Vicari-4 locality, previously referred to the late Campanian but with no further information on the parataxonomic affinity of the eggshells (Torices Hernandez, 2002; Torices et al. 2004; in press). Vicari-4 is geographically near to where Moratalla (1993) reported the Vicari locality, being located in the same ravine, and we consider that, if not the same site, they may both have a similar stratigraphic position and age. The present work maintains that M. pseudomamillare and M. trempii must be synonymised with M. baghensis and M. cf mamillare with M. mamillare (Appendix 2); therefore, the occurrence of these oospecies in the abovementioned localities (Orcau, Llabusta, Vicari, Compuertas, Fontllonga L, Suterranya, Suterranya-1 and Abella) indicates a late Maastrichtian age. At the south-western margin of the Tremp Basin, we concur with Prieto-Marquez et al. (2013) that the Moror locality (the "Moro" locality reported by Brinkmann, 1984, 1988) is probably late Maastrichtian in age given the diachronous deposition of the "grey unit", which took place in a regressive context, and given that it becomes younger westward. Likewise, the Moror-1 locality would be of similar age. However, the latter locality yielded what were purported to be M. petralta eggshells and was dated as "early Rognacian" (Vianey-Liaud and Lopez-Martinez, 1997). Pending the revision of this eggshell material, the age of this locality remains controversial. The Montrebei locality, which was originally regarded as early Maastrichtian (Lopez-Martinez, 2000) and later as late Campanian-early Maastrichtian in age (Torices et al., in press), may also be of late Maastrichtian age. Unfortunately, there are no published data on the parataxonomic affinity of the dinosaurs eggshells recovered in this locality (Lopez-Martinez, 2000), and the charophyte content (only Clavator brachycerus) has an ambiguous and wide stratigraphic range (Villalba-Breva and Martin-Closas, 2012).

In the Ager syncline, the Perauba-Figuerola locality represents the only south-Pyrenean locality with rhabdodontid remains (Llompart and Krauss, 1982) and yielded the oospecies Megaloolithus aureliensis (Appendix 1). Torices et al. (in press) reported dinosaur eggshells at the Figuerola-2 locality, but no further taxonomic assignation is provided by the authors. Figuerola-2 was originally regarded as early Maastrichtian (Torices Hernandez, 2002) and later as late Campanian (Lopez-Martinez, 2003; Torices et al. 2004; in press). The two localities are geographically adjacent to one another (about 25 m apart) and stratigraphically almost equivalent. They are located in the lower part of the "Figuerola de Meia" unit, the regional equivalent of the "lower red unit" of the Tremp Formation. This continental unit contains a charophyte assemblage of Maastrichtian age (Septorella brachycera, S. ultima, Peckichara sertulata, and Maedleriella sp.; Fig. 2B) and overlies the La Ma?ana Fm., which is considered late Campanian in age on the basis of its charophyte assemblages (Villalba-Breva and Martin-Closas, 2012). Thus, the recent data on charophyte biostratigraphy indicate that both localities are most probably early Maastrichtian in age. The new age assignment is consistent with the magnetostratigraphical data of Galbrun et al. (1993), who correlated the base of the "Figuerola de Meia" unit with the C32r magnetochron (currently early Maastrichtian in age after Gradstein et al., 2012).

Lopez-Martinez (2003) reported dinosaur eggshells from the Blancafort locality although she did not provide any further parataxonomic assignment. Recently, we have collected several eggshells, which have been attributed to Megaloolithus aureliensis (Appendices 1 and 2). The samples come from a lacustrine limestone on top of the La Ma?ana Fm. (late Campanian in age according to charophyte assemblages; Villalba-Breva and Martin-Closas, 2012). The occurrence of the oospecies M. aureliensis oospecies in Blancafort is consistent with this age assignment since this oospecies ranges temporally from the late Campanian to the early Maastrichtian (Garcia and Vianey-Liaud, 2001).

In the Coll de Nargo syncline most of the multiple egg-bearing sites (Appendix 1) in the localities of Sallent, El Codo, Pinyes Est, Santa Eulalia, La Teuleria, Cal Fontanet, and Els Encerris yielded a single oospecies, Megaloolithus siruguei (Appendix 2). The occurrence of this single oospecies for more than 100 m of stratigraphic section seems to indicate that most of these localities fall within the oozone 2 (in accordance with Selles et al., 2013), which is mainly characteristic of the early Maastrichtian (Fig. 2A, B). In the same region the El Mirador locality yielded three megaloolithid oospecies (Cairanoolithus roussetensis, Megaloolithus siruguei and M. aureliensis; Selles et al., 2013). This cooccurrence originally seemed to be characteristic of the late Campanian Oozone 1 of Garcia and Vianey-Liaud (2001) but with the new age calibration of Gradstein et al. (2012) the assemblage extends within the early Maastrichtian (C32n.1n). In the case of El Mirador locality, Selles et al. (2013) assumed that it probably represents the last co-occurrence of these ootaxa in the Coll de Nargo area, and therefore the site may fall on the upper part of the C32n.1n and be early Maastrichtian in age. Finally, as noted by Vila et al. (2011) and Selles et al. (2013), the egg-bearing sites of the upper part of the Coll Nargo and Vallcebre sections contain the oospecies M. siruguei right in contact with M. mamillare or M. baghensis (sites of Pinyes Est-5, Santa Eulalia-3, El Codo-41, E-MUN06, J-BAR01; Appendix 1). This probably indicates that these sites fall around the C31r-C31n reversal, which is late Maastrichtian in age according to the recent age calibrations of Gradstein et al. (2012).

Regarding the new eggshell-bearing localities reported in the present work (Appendices 1 and 2), some of them yielded M. mamillare eggshells (La Nou-1, Bergueda area, Barcelona; Tossal de Sant Roma d'Abella, and Costa de la Coma, Tremp Basin, Lleida) and M. baghensis eggshells (Sant Bartomeu, Barranc de la Boi'ga, Moli del Baro-1, Serrat del Rostiar-2, and Cami del Soldat, Tremp Basin, Lleida; L'Espinau, Ager Syncline, Lleida) and hence indicate a late Maastrichtian age. It is worth noting that some of these reports (localities of La Nou-1, Cami del Soldat and Moli del Baro-1) expand the temporal range of these two oospecies (M. mamillare and M. baghensis) into the C29r magnetochron (Fig. 2A).

5.2 Implications for dinosaur faunal turnover

Le Loeuff et al. (1994) stated that a faunal replacement occurred around the early Maastrichtian-late Maastrichtian boundary in the so-called Ibero-Armorican Domain in southwestern Europe. During this turnover a late Maastrichtian assemblage dominated by hadrosauroids replaced an early Maastrichtian fauna dominated by titanosaurian sauropods. To understand how this purported turnover took place it is pivotal to establish the age of the dinosaur-bearing localities throughout the region. The new age assignments proposed for some south-Pyrenean localities have implications mainly for the temporal distribution of certain dinosaur taxa (ankylosaurians and theropods) in the latest Cretaceous and therefore for the turnover pattern of south-western Europe (Fig. 2). In the southern basins, for example, the localities of Fontllonga-6, Biscarri, Els Nerets and Suterranya-Mina de lignit yielded ankylosaurian remains (Santafe-Llopis et al., 1997; Lopez-Martinez et al., 2000; Lopez-Martinez, 2003; Escaso et al., 2010). The present revision indicates that at least two of these localities (Els Nerets and Suterranya-Mina de lignit) have a late Maastrichtian age, and this implies that the ankylosaurian dinosaurs persisted until the early late Maastrichtian (Fig. 2A). This assertion concurs with what has been reported in the northern Pyrenees (Fig. 2C), where ankylosaurian remains have been collected in deposits of late Maastrichtian age (Laurent et al., 2002). As a whole, the Pyrenean record seems to indicate that the clade did not go extinct at the early Maastrichtian-late Maastrichtian boundary but they lasted beyond it and coexisted for some time with hadrosauroid faunas.

In terms of theropod distribution and fossil abundance, the new age assignments proposed for some localities modify the recent theropod successions proposed by Torices et al. (in press) in the southern Pyrenees. For instance, the authors situated several of the localities with theropod teeth in the late Campanian. However, the Figuerola-2 locality is here considered to be early Maastrichtian in age, whereas the localities of Vicari-4, Suterranya-1 and probably Montrebei are late Maastrichtian in age. This implies that some of the theropod taxa reported in late Campanian localities are now restricted to the late Maastrichtian (Fig. 2A). This is the case for cf. Richardoestesia sp. and "Richardoestesia-like", which now seem to be restricted exclusively to the late Maastrichtian, at least in the southern Pyrenees. Moreover, with the new distribution the theropod record in the southern Pyrenees clearly varies through the latest Cretaceous in terms of the predominant type of evidence. That is, the oological record is more diverse than the bone record prior to the early Maastrichtianlate Maastrichtian boundary (Selles et al., 2014). By contrast, the bone record (mainly teeth) is much more highly represented in deposits of late Maastrichtian age, being scarce in the late Campanian-early Maastrichtian interval. This pattern, which is probably influenced by sampling and other biases, comes to light clearly in Fig. 2C, where the theropod record is notably more concentrated in the late Maastrichtian.

Regarding sauropods, the late Maastrichtian age of some eggshell localities (e.g. Els Nerets, Basturs, Orcau, Suterranya-1) demonstrates that the egg record of this group is almost continuous and abundant throughout the entire Maastrichtian. This contradicts previous statements made by Lopez-Martinez (2003), who postulated a major abundance of egg localities in Upper Campanian and a decrease in Maastrichtian deposits. In the light of the new temporal distribution of the localities and the integration of new sites from other areas (e.g. the Coll de Nargo and Vallcebre synclines) the egg record of sauropod dinosaurs becomes notably scarce in Upper Campanian deposits but is well represented in the Maastrichtian

The new age assignments do not have significant implications for the temporal distribution of rhabdodontid and hadrosauroid dinosaurs, since the ornithopod-bearing localities discussed in the present work (Perauba-Figuerola, and Els Nerets and L'Espinau, respectively) fall within the time interval expected for each group in the region. The present study will hopefully lay the groundwork for future studies on dinosaur biostratigraphy not only in the southern Pyrenees but also in the whole Ibero-Armorican Domain.

6. Conclusions

The present study updates the nomenclature and age of several classic and new dinosaur fossil localities in the uppermost Cretaceous (Campanian-Maastrichtian) continental deposits of the southern Pyrenees. The age of about 30 localities has been re-evaluated by means of an analysis and review of new and previous parataxonomic assignations, with some material re-assigned to known megaloolithid oospecies (M. aureliensis, M. siruguei, M. mamillare, M. baghensis). The new age assignation includes classical dinosaur-bearing localities such as Basturs, Orcau-1, Els Nerets, Suterranya-1 and Figuerola-2, among others, which have significant implications for the faunal turnover proposed in south-western Europe. According to the new data, the ankylosaurians would have survived until the late Maastrichtian in the southern Pyrenees, as reported in the northern basins. The new age assessment suggests that in the southern Pyrenees the fossil record of theropods is scarce prior to the early Maastrichtian and that their taxonomic diversity (richness) is much higher in the late Maastrichtian. Interestingly, the sauropod egg record becomes continuous throughout the entire Maastrichtian but is scarce in the Upper Campanian.

Appendix 1.-Listed samples and localities with megaloolithid oospecies
discussed in the present work.

Locality            Egg-bearing level (site)    Source      Number of
                                                material     samples

Barranc de La       Barranc de La Fonguera-1   Egg debris       7
  Fonguera          Barranc de La Fonguera-2   Egg debris       12
Barranc de                                     Eggshells        95
  la Boiga
Orcau-1                                        Eggshells        38
Els Nerets                                     Eggshells        51
Suterranya-1                                      Eggs          25
Abella                                            Eggs          5
Tossal de Sant                                    Eggs          84
  Roma d'Abella
Costa de la Coma                                  Eggs         110
Sant Bartomeu                                     Egg           6
Moli del Baro-1                                Eggshells        50
Serrat del                                     Eggshells        1
Cami del Soldat                                Eggshells        50
Els Terrers                 J-BAR01               Egg           4
Torrent de                  E-MUN06            Eggshells        1
La Nou-1                                       Eggshells       119
L'Espinau                                      Eggshells        8
Perauba-Figuerola                              Eggshells        1
Blancafort                                     Eggshells        17
Sallent                    Sallent-1              Eggs          6
                           Sallent-2              Eggs          12
                           Sallent-3           Egg debris       3
                           Sallent-4              Eggs          4
                           Sallent-5              Eggs          6
                           Sallent-6              Eggs          3
                           Sallent-7              Eggs          13
Cal Fontanet             Cal Fontanet-1           Eggs          29
                         Cal Fontanet-2           Eggs          10
                         Cal Fontanet-3           Eggs          9
                         Cal Fontanet-4           Eggs          26
Pinyes Est                Pinyes Est-1            Eggs          77
                          Pinyes Est-2            Eggs          11
                          Pinyes Est-3         Egg debris       20
                          Pinyes Est-4            Eggs          12
                          Pinyes Est-5            Eggs          23
Santa Eulalia           Santa Eulalia-1           Eggs          15
                        Santa Eulalia-2           Eggs          28
                        Santa Eulalia-3           Eggs          6
                        Santa Eulalia-4        Egg debris       4
La Teuleria              La Teuleria-1            Eggs          50
                         La Teuleria-2            Eggs          93
                         La Teuleria-3            Eggs          5
                         La Teuleria-4            Eggs         152
                         La Teuleria-5            Eggs          7
                         La Teuleria-6            Eggs          92
                         La Teuleria-7            Eggs         117
                         La Teuleria-8            Eggs          96
                         La Teuleria-9            Eggs          78
                         La Teuleria-10           Eggs          11
Els Enserris             Els Enserris-1           Eggs          9
                         Els Enserris-2           Eggs          6
                         Els Enserris-3           Eggs          30
                         Els Enserris-4           Eggs          15
                         Els Enserris-5           Eggs          13
                         Els Enserris-5           Eggs          15
                         Els Enserris-6           Eggs          13
                         Els Enserris-7           Eggs          10
                         Els Enserris-8           Eggs          20
                         Els Enserris-9           Eggs          7
                        Els Enserris-10           Eggs          67
                        Els Enserris-11           Eggs          32
El Codo                  El Codo-1                Eggs          33
                         El Codo-2                Eggs          34
                         El Codo-3                Eggs          11
                         El Codo-4                Eggs          5
                         El Codo-5                Eggs          3
                         El Codo-6                Eggs          3
                         El Codo-7                Eggs          29
                         El Codo-8                Eggs          38
                         El Codo-9                Eggs          18
                         El Codo-10               Eggs          3
                         El Codo-11               Eggs          5
                         El Codo-12               Eggs          12
                         El Codo-13               Eggs          42
                         El Codo-14               Eggs          15
                         El Codo-15               Eggs          2
                         El Codo-16               Eggs          47
                         El Codo-17               Eggs          8
                         El Codo-18               Eggs          16
                         El Codo-19               Eggs          26
                         El Codo-20               Eggs          12
                         El Codo-21               Eggs          33
                         El Codo-22               Eggs          28
                         El Codo-23               Eggs          10
                         El Codo-24               Eggs          10
                         El Codo-25               Eggs          15
                         El Codo-26               Eggs          23
                         El Codo-27               Eggs          15
                         El Codo-28               Eggs          45
                         El Codo-29               Eggs          15
                         El Codo-30               Eggs          15
                         El Codo-31               Eggs          10
                         El Codo-32               Eggs          8
                         El Codo-33               Eggs          18
                         El Codo-34               Eggs          20
                         El Codo-35               Eggs          35
                         El Codo-36               Eggs          24
                         El Codo-37               Eggs          7
                         El Codo-38               Eggs          35
                         El Codo-39               Eggs          53
                         El Codo-40               Eggs          10
                         El Codo-41               Eggs          8

Locality            Egg-bearing level (site)      Eggshell
                                               thickness (mm)

Barranc de La       Barranc de La Fonguera-1     2.67-2.73
  Fonguera          Barranc de La Fonguera-2     2.17-2.39
Barranc de                                       1.33-1.53
  la Boiga
Orcau-1                                          1.50-1.54
Els Nerets                                       1.64-1.77
Suterranya-1                                     1.76-1.78
Abella                                              1.85
Tossal de Sant                                   1.54-1.62
  Roma d'Abella
Costa de la Coma                                 1.56-1.64
Sant Bartomeu                                    1.47-1.88
Moli del Baro-1                                  1.87-1.93
Serrat del                                          1.82
Cami del Soldat                                  1.12-1.32
Els Terrers                 J-BAR01               0.7-0.9
Torrent de                  E-MUN06                 0.75
La Nou-1                                         1.51-1.59
L'Espinau                                        1.25-1.85
Perauba-Figuerola                                   1.57
Blancafort                                       1.50-1.61
Sallent                    Sallent-1             2.78-2.95
                           Sallent-2             2.93-3.22
                           Sallent-3             2.92-2.95
                           Sallent-4             2.80-3.10
                           Sallent-5             2.11-2.34
                           Sallent-6             2.79-2.93
                           Sallent-7             2.57-2.65
Cal Fontanet             Cal Fontanet-1          3.14-3.66
                         Cal Fontanet-2          3.33-3.61
                         Cal Fontanet-3          3.34-3.76
                         Cal Fontanet-4          3.45-3.65
Pinyes Est                Pinyes Est-1           2.51-2.59
                          Pinyes Est-2           2.73-2.87
                          Pinyes Est-3           2.37-2.79
                          Pinyes Est-4           2.54-2.72
                          Pinyes Est-5           1.98-2.07
Santa Eulalia           Santa Eulalia-1          2.53-2.92
                        Santa Eulalia-2          2.48-2.73
                        Santa Eulalia-3          1.97-2.19
                        Santa Eulalia-4          2.63-2.77
La Teuleria              La Teuleria-1           2.29-2.49
                         La Teuleria-2           3.08-3.16
                         La Teuleria-3           2.57-2.66
                         La Teuleria-4           2.86-3.01
                         La Teuleria-5           2.93-3.22
                         La Teuleria-6           2.48-2.73
                         La Teuleria-7           2.31-2.45
                         La Teuleria-8           2.78-2.95
                         La Teuleria-9           2.52-2.65
                         La Teuleria-10          2.71-2.75
Els Enserris             Els Enserris-1          2.68-2.82
                         Els Enserris-2          2.88-2.93
                         Els Enserris-3          2.66-3.04
                         Els Enserris-4          2.76-2.94
                         Els Enserris-5          2.85-2.87
                         Els Enserris-5          3.21-3.33
                         Els Enserris-6          3.04-3.09
                         Els Enserris-7          2.11-2.81
                         Els Enserris-8          2.43-2.72
                         Els Enserris-9          2.83-2.99
                        Els Enserris-10          2.63-2.67
                        Els Enserris-11          2.78-2.92
El Codo                  El Codo-1               2.53-2.56
                         El Codo-2               2.40-2.57
                         El Codo-3               2.57-2.66
                         El Codo-4               2.58-2.62
                         El Codo-5               2.22-2.31
                         El Codo-6               2.73-2.84
                         El Codo-7               2.82-2.88
                         El Codo-8               2.62-2.92
                         El Codo-9               2.72-2.84
                         El Codo-10              3.08-3.26
                         El Codo-11              2.38-2.52
                         El Codo-12              2.38-2.59
                         El Codo-13              2.29-2.49
                         El Codo-14              2.24-2.48
                         El Codo-15              3.07-3.13
                         El Codo-16              2.84-2.99
                         El Codo-17              2.64-2.72
                         El Codo-18              2.53-2.67
                         El Codo-19              2.77-2.86
                         El Codo-20              2.51-2.63
                         El Codo-21              3.18-3.33
                         El Codo-22              2.70-2.75
                         El Codo-23              2.16-2.35
                         El Codo-24              2.28-2.42
                         El Codo-25              2.84-2.87
                         El Codo-26              2.20-2.32
                         El Codo-27              2.86-2.92
                         El Codo-28              2.77-2.97
                         El Codo-29              2.76-2.99
                         El Codo-30              2.31-2.45
                         El Codo-31              2.71-2.79
                         El Codo-32              3.08-3.20
                         El Codo-33              2.54-2.76
                         El Codo-34              2.44-2.57
                         El Codo-35              2.47-2.56
                         El Codo-36              3.15-3.19
                         El Codo-37              2.29-2.33
                         El Codo-38              2.78-2.83
                         El Codo-39              2.28-2.34
                         El Codo-40              2.17-2.25
                         El Codo-41              1.98-2.07

Locality            Egg-bearing level (site)          Oospecies

Barranc de La       Barranc de La Fonguera-1   Megaloolithus siruguei
  Fonguera          Barranc de La Fonguera-2   Megaloolithus siruguei
Barranc de                                     Megaloolithus baghensis
  la Boiga
Orcau-1                                        Megaloolithus baghensis
Els Nerets                                     Megaloolithus mamillare
Suterranya-1                                   Megaloolithus baghensis
Abella                                         Megaloolithus mamillare
Tossal de Sant                                 Megaloolithus mamillare
  Roma d'Abella
Costa de la Coma                               Megaloolithus mamillare
Sant Bartomeu                                  Megaloolithus baghensis
Moli del Baro-1                                Megaloolithus baghensis
Serrat del                                     Megaloolithus baghensis
Cami del Soldat                                Megaloolithus baghensis
Els Terrers                 J-BAR01            Megaloolithus baghensis
Torrent de                  E-MUN06            Megaloolithus baghensis
La Nou-1                                       Megaloolithus mamillare
L'Espinau                                      Megaloolithus baghensis
Perauba-Figuerola                             Megaloolithus aureliensis
Blancafort                                    Megaloolithus aureliensis
Sallent                    Sallent-1           Megaloolithus siruguei
                           Sallent-2           Megaloolithus siruguei
                           Sallent-3           Megaloolithus siruguei
                           Sallent-4           Megaloolithus siruguei
                           Sallent-5           Megaloolithus siruguei
                           Sallent-6           Megaloolithus siruguei
                           Sallent-7           Megaloolithus siruguei
Cal Fontanet             Cal Fontanet-1        Megaloolithus siruguei
                         Cal Fontanet-2        Megaloolithus siruguei
                         Cal Fontanet-3        Megaloolithus siruguei
                         Cal Fontanet-4        Megaloolithus siruguei
Pinyes Est                Pinyes Est-1         Megaloolithus siruguei
                          Pinyes Est-2         Megaloolithus siruguei
                          Pinyes Est-3         Megaloolithus siruguei
                          Pinyes Est-4         Megaloolithus siruguei
                          Pinyes Est-5         Megaloolithus mamillare
Santa Eulalia           Santa Eulalia-1        Megaloolithus siruguei
                        Santa Eulalia-2        Megaloolithus siruguei
                        Santa Eulalia-3        Megaloolithus mamillare
                        Santa Eulalia-4        Megaloolithus siruguei
La Teuleria              La Teuleria-1         Megaloolithus siruguei
                         La Teuleria-2         Megaloolithus siruguei
                         La Teuleria-3         Megaloolithus siruguei
                         La Teuleria-4         Megaloolithus siruguei
                         La Teuleria-5         Megaloolithus siruguei
                         La Teuleria-6         Megaloolithus siruguei
                         La Teuleria-7         Megaloolithus siruguei
                         La Teuleria-8         Megaloolithus siruguei
                         La Teuleria-9         Megaloolithus siruguei
                         La Teuleria-10        Megaloolithus siruguei
Els Enserris             Els Enserris-1        Megaloolithus siruguei
                         Els Enserris-2        Megaloolithus siruguei
                         Els Enserris-3        Megaloolithus siruguei
                         Els Enserris-4        Megaloolithus siruguei
                         Els Enserris-5        Megaloolithus siruguei
                         Els Enserris-5        Megaloolithus siruguei
                         Els Enserris-6        Megaloolithus siruguei
                         Els Enserris-7        Megaloolithus siruguei
                         Els Enserris-8        Megaloolithus siruguei
                         Els Enserris-9        Megaloolithus siruguei
                        Els Enserris-10        Megaloolithus siruguei
                        Els Enserris-11        Megaloolithus siruguei
El Codo                  El Codo-1             Megaloolithus siruguei
                         El Codo-2             Megaloolithus siruguei
                         El Codo-3             Megaloolithus siruguei
                         El Codo-4             Megaloolithus siruguei
                         El Codo-5             Megaloolithus siruguei
                         El Codo-6             Megaloolithus siruguei
                         El Codo-7             Megaloolithus siruguei
                         El Codo-8             Megaloolithus siruguei
                         El Codo-9             Megaloolithus siruguei
                         El Codo-10            Megaloolithus siruguei
                         El Codo-11            Megaloolithus siruguei
                         El Codo-12            Megaloolithus siruguei
                         El Codo-13            Megaloolithus siruguei
                         El Codo-14            Megaloolithus siruguei
                         El Codo-15            Megaloolithus siruguei
                         El Codo-16            Megaloolithus siruguei
                         El Codo-17            Megaloolithus siruguei
                         El Codo-18            Megaloolithus siruguei
                         El Codo-19            Megaloolithus siruguei
                         El Codo-20            Megaloolithus siruguei
                         El Codo-21            Megaloolithus siruguei
                         El Codo-22            Megaloolithus siruguei
                         El Codo-23            Megaloolithus siruguei
                         El Codo-24            Megaloolithus siruguei
                         El Codo-25            Megaloolithus siruguei
                         El Codo-26            Megaloolithus siruguei
                         El Codo-27            Megaloolithus siruguei
                         El Codo-28            Megaloolithus siruguei
                         El Codo-29            Megaloolithus siruguei
                         El Codo-30            Megaloolithus siruguei
                         El Codo-31            Megaloolithus siruguei
                         El Codo-32            Megaloolithus siruguei
                         El Codo-33            Megaloolithus siruguei
                         El Codo-34            Megaloolithus siruguei
                         El Codo-35            Megaloolithus siruguei
                         El Codo-36            Megaloolithus siruguei
                         El Codo-37            Megaloolithus siruguei
                         El Codo-38            Megaloolithus siruguei
                         El Codo-39            Megaloolithus siruguei
                         El Codo-40            Megaloolithus siruguei
                         El Codo-41            Megaloolithus siruguei

Appendix 2.- Systematic palaeontology

Basic organizational group: Dinosauroid

Oofamily: Megaloolithidae Zhao, 1979

Oogenus: Megaloolithus Vianey-Liaud, Mallan, Buscail, and Montgelard, 1994

Megaloolithus aureliensis Vianey-Liaud, Mallan, Buscail, and Montgelard, 1994

Diagnosis according to Garcia and Vianey-Liaud (2001): Megaloolithus with short fan-shaped units; arched growth lines can be horizontal in the small area with flat outer surface; thickness ranges from 0.75 to 1.52 mm; average node diameter about 0.5 mm; pore diameters between 10 and 50 pm; spherical eggs (until 20-22 cm).

Localities, stratigraphic range, and age: This Megaloolithus ooespecies is represented by scattered eggshells at the Blancafort and Perauba-Figuerola localities. The former locality falls on top of the La Malana unit, the regional equivalent of the "grey unit" of the Tremp Formation, late Campanian (Villalba-Breva and MartinClosas, 2012). The Perauba-Figuerola locality locates at the lower part of the "Figuerola de Meia" unit, the regional equivalent of the "lower red unit" of the Tremp Formation, early Maastrichtian (Villalba-Breva and Martin-Closas, 2012).

Material: see Appendix 1.

Description: The outer surface of the eggshell is covered with scattered rounded nodes (0.5 mm in diameter), sometimes coalescent, with flat areas between them (Fig. 1A). Circular-shaped pore apertures (80-120 pm in diameter) are located at the base of nodes, and less frequently in the middle of flatted areas. Eggshell ranges from 0.8 mm to 1.2 mm in thickness. In radial view, short fanshaped shell units (H/W ratio of 1.8) are interlocked with adjacent ones (Fig. 1B). Growth lines are mainly arched at the base of those shell units, whereas they are undulating when crossing fused units. Some tubocanaliculate pore channels have been also observed in radial section, which are 95-120 pm in width.

Comparisons and Discussion: Coalescent-nodular ornamentation like that of eggshells from the Blancafort and Perauba-Figuerola localities has been described in four megaloolithid oospecies: M. baghensis, Pseudomegaloolithus atlasis, Patagoolithus salitraensis and M. aureliensis (Vianey-Liaud et al., 2003; Vianey-Liaud and Garcia, 2003). All these oospecies exhibit similar structural features, such as shell-thickness, shape of shell units, and node and pore dimension; being the high/width ratio (H/W ratio) of the shell units the most notable differences between them. This value is about 2.33 in M. baghensis (Khosla and Sahni, 1995), less than 3 in P. atlasis (Vianey-Liaud and Garcia, 2003), 2.28 in P. salitralensis (Simon, 2006) and 1.75 inM. aureliensis (Selles et al., 2013). Given that H/W ratio established for the studied material is 1.8, this value is more similar to that of M. aureliensis than any other oospecies. Although eggshell thickness can be highly variable within a certain oospecies, and that eggshell-thickness-range may overlap between several oospecies, it is worthy comment that the thickness of the oological material from the studied localities is more similar to M. aureliensis (0.9 mm and 0.98mm, respectively) than any other of the previously referred oospecies (0.7 mm-thick in P. atlasis, 1.5 mm-thick in M. baghensis, and 1.4 mm-thick in P. salitralensis). Although the size of pore apertures of the studied material is somewhat larger than that reported by Garcia and Vianey-Liaud (2001), Selles et al. (2013) described some specimens of M. aureliensis from Coll de Nargo syncline with similar values of pore apertures size than those of the present work. Therefore, eggshells from the Blancafort and Perauba-Figuerola localities are attributed to the oospecies M. aureliensis.

Megaloolithus siruguei Vianey-Liaud, Mallan, Buscail, and Montgelard, 1994

Diagnosis: According to Vianey-Liaud et al (1994) and after Elez and Lopez-Martinez (2000), and Vianey-Liaud and Zelenitsky (2003), Megaloolithus with shell units taller than in M. mammilare; thickness range 2.65 to 2.70 mm; pore diameter range 50 to 80 pm; reticulate pore system.

Localities, stratigraphic range, and age: This Megaloolithus ooespecies is represented by eggs and/or scattered eggshells at the sites of Barranc de la Fonguera-1, and -2, Sallent-1 to -7, Cal Fontanet-1 to -4, Pinyes Est-1 to -4, La Teuleria-1 to -10, Santa Eulalia-1, -2, and -4, El Codo-1 to -40, and Els Enserris-1 to -11. They all fall in the "lower red unit" of the Tremp Formation. Early Maastrichtian - ?early late Maastrichtian.

Material: see appendix 1.

Description: Relatively thick eggshells (1.75 mm to 3.6 mm), fully covered by well-delimited rounded nodes (0.51 - 1.23 mm in diameter, Fig. 1C). Circular to sub-circular pore apertures (130 [micro]m-width), which are located near the base of the nodes. In radial section, the edge of the elongate fan-shaped shell units (H/W ratio = 4) can be easily traceable throughout whole eggshell. Nevertheless, some of them are partially interlocked with their neighbouring ones (Fig. 1D). Growth lines are clearly arched from the base to the top of the shell units. The respiratory channel system is composed of vertical funnel-shaped channels interconnected by transversal channels, altogether forming a reticulate network of channels.

Comparisons and Discussion: Only two oospecies of Megaloolithus exhibit a clear reticulate channel system: M. siruguei and M. microtuberculata. The surface of M. microtuberculata is covered of pronounced small nodes, sometimes with irregular shape (Garcia and Vianey-Liaud, 2001), while M. siruguei is covered of large rounded nodes, as in eggshells described above. In fact, the ornamental pattern is the only difference observed between these oospecies. Given that all the oological remains examined here exhibit large rounded nodes in their outer surface and reticulate pore system, they are classified as Megaloolithus siruguei.

Megaloolithus mamillare Vianey-Liaud, Mallan, Buscail, and Montgelard, 1994

Diagnosis according to Vianey-Liaud et al. (1994): Megaloolithus with short fan-shaped units; thickness in the main part of the eggshells from 1.2 to 2.1 mm; average node diameter about 1 mm; pore diameters range from 75 to 120 pm.

Synonymy: Megaloolithus cf. mamillare from the Compuertas locality of Moratalla (1993)

Localities, stratigraphic range, and age: This Megaloolithus ooespecies is represented by eggs and scattered eggshells at the Basturs-1, Basturs-2, Abella and Sant Bartomeu localities, on top of the Aren Sandstone Formation; eggshell fragments in the Els Nerets and Compuertas localities, in the "grey unit" of the Tremp Formation; and eggs and/or eggshell fragments from the Santa Eulalia-3, El Codo-41, Pinyes Est-5, Tossal de Sant Roma d'Abella, Costa de la Coma, and La Nou-1 localities, in the upper part of the "lower red unit" of the Tremp Formation. Late Maastrichtian.

Material: see Appendix 1.

Description: Some nearly complete sub-spherical eggs (19-20 cm in diameter) have been discovered in Tossal de Sant Roma d'Abella locality. Eggshell ranges from 1.39 mm to 2.3 mm in thickness (average of 1.9 mm) with the outer surface covered of small rounded nodes (0.73 mm in diameter). Sometimes, nodes are coalescenced between two or five nodes, forming short and irregular chains (Fig. 1E). Circular-shaped pore apertures (90-110 [micro]m in diameter) are located between ornamental nodes. In radial thin section, fan-shaped shell units (H/W ratio of 2) are clearly traceable; showing well curved growth lines from the base to the top of the crystalline units (Fig. 1F). The respiratory system is tubocanaliculate, with narrow and slender canals, ranging from 50 to 80 [micro]m in wide.

Comparisons and Discussion: Structural characters exhibited by eggshells described above are consistent with three oospecies of Megaloolithus: M. jabalpurensis, M. dhoridungriensis and M. mamillare. The Indian oospecies M. dhoridungriensis differs from our specimens by showing cylindrical-shape shell units, whit a H/W ratio of 2.7 (Mohabey, 1998). M. jabalpurensis and M. mamillare share several structural characters (Vianey-Liaud et al., 1994, 2003), only differing in the size of the egg (15 cm and 20 cm in diameter, respectively), the eggshell thickness (2.3 mm in M. jabalpurensis and 2.1 mm in M. mamillare) and the H/W ratio (2.45 in M. jabalpurensis and 2.1 to 2.2 in M. mamillare). Given that the studied oological material exhibits sub-spherical eggs of 20 cm in diameter, 1.9 mm-thick eggshell, and H/W ratio of 2, it is attributed to the oospecies M. mamillare.

Moratalla (1993, 1998) described very scarce remains attributed to M. cf. mamillare in the Compuertas locality. The author stated that the recovered eggshells were badly preserved, obscuring several features. However, he noted that the thin eggshells collected in the locality exhibited compact and nodular ornamentation and non-fused shell units. Although we had not the chance to analyze this egg material directly on first-hand examination, all evidence point that the abovementioned characters described in the eggshells from the Compuertas locality fit with those of the oospecies M. mamillare. Accordingly, we tentatively assign the oological remains of this locality to M. mamillare.

Megaloolithus baghensis Khosla and Sahni 1995

Diagnosis according to Fernandez and Khosla (in press): Spherical eggs 140-200 mm in diameter; nodes ornamentation, eggshell 1.0-1.70 mm thick; average node diameter about 0.60 mm; fanshaped spheroliths distinct or even partially fused; height/width ratio 2.32:1; pore subcircular to elliptical; swollen-ended, variably spaced basal caps (0.2-0.3 mm in diameter).

Synonymies: Megaloolithuspseudomamillare from Suterranya-1 locality; Vianey-Liaud and Lopez-Martinez (1997)

Megaloolithus trempii from Orcau, Suterranya, Vicari, Llabusta, and Fontllonga L localities; Moratalla (1998)

Megaloolithus cf. mamillare from the Suterranya and Orcau localities; Moratalla (1993, 1998)

Megaloolithus sp.; Vila et al. (2011)

Localities, stratigraphic range, and age: This Megaloolithus ooespecies is represented by eggs at Suterranya-1 and by scattered eggshells at the Orcau-1, Orcau, Llabusta, Vicari, Suterranya, Fontllonga L, Barranc de la Boiga, Moli del Baro-1, Serrat del Rostiar-2, Cami del Soldat and L'Espinau localities. The Suterranya-1 localitiy is found on top of the Aren Sandstone Formation; the Orcau, Llabusta, Vicari, and Suterranya are found in the "grey unit" of the Tremp Formation; the Fontllonga L, Barranc de la Boiga, Moli del Baro-2, Serrat del Rostiar-2, Cami del Soldat and L'Espinau localities are located in the "lower red unit" of the Tremp Formation. Late Maastrichtian.

Material: see Appendix 1.

Description: complete eggs from Suterranya-1 and Suterranya localities are sub-spherical in shape. The smallest egg is 15-18 cm in diameter, while the largest is 21-23 cm, and the eggs of Suterranya are about 21 cm in diameter (Moratalla, 1993). The outer surface of the shell is covered of coalescent rounded nodes ranging from 0.2 to 0.6 mm in diameter. Nevertheless, most part of the shell surface consists in flat areas and isolate nodes (Fig. 1G). Pore opening are 0.1-0.2 mm in diameter frequently located at the flattered surfaces. The eggshell thickness ranges from 1.12 to 1.85 mm, with an average of 1.39 mm. Eggshell units are broad fan-shaped (H/W ratio about 1.9), frequently fused with adjacent ones, and showing irregular morphology in their upper part (Fig. 1G). In radial thin section, the pore channels are narrow and straight (75-90 [micro]m-width), corresponding to tubocanaliculate pore system. Growth lines appear slightly arched from the base to the top of the shell units.

Comparisons and Discussion: The oological material described above resembles M. aureliensis oospecies in the ornamental patterns, and H/W ratio. However, this material is somewhat thicker than M. aureliensis (1.1-1.9 mm and 0.8-1.4 mm, respectively). In addition, they differ in the shape of shell units, being more irregular in the studied material than that of M. aureliensis. M. baghensis also share several features with the studied material. For instance, the range of shell thickness of M. baghensis (1-1.7 mm; Mohabey, 1998) is similar to that that of our specimens (1.1-1.9 mm-thick). Both ootaxa exhibit similar size of node (0.65 mm in diameter in M. baghensis and 0.5 mm in studied material) and pore apertures (0.15 mm in diameter in M. baghensis and 0.1-0.2 mm in studied material). Therefore, the oological remains described in the abovementioned localities are attributed to the oospecies M. baghensis.

Moratalla (1993, 1998) reported a new oospecies named Megaloolithus trempii in the Orcau, Suterranya-1, Vicari, Llabusta, and Fontllonga L localities of the Tremp and Ager basins. A recent revision of this ootaxon reveals identical microstructural characters (e.g. shell thickness, ornamental pattern, shape of shell units) than those of Megaloolithus baghensis, and hence both oospecies should be synonymised. The eggs sizes reported by Moratalla (1993, 1998) are slightly larger than those described in the diagnosis, albeit this can be explained as the result of a tangential-view section of the eggs or even due to regional tectonic deformation of the eggs (see Vila et al. 2010). Thus, the occurrence of M. baghensis in the Moratalla's (1993, 1998) localities is here stated. Moratalla (1993, 1998) also reported the presence of M. cf. mamillare in the Orcau and Suterranya localities. On the base of descriptions given by the author, we note that all characters (such as the ornamentation composed of rounded nodes and flatted surfaces, irregular shell units, and partially fused shell unit) fit with the oospecies M. baghensis. Therefore, we propose to synonymize the material referred to M. cf. mamillare in these localities with M. baghensis. Similarly, Vila et al. (2011) reported an egg and eggshell fragments attributed to Megaloolithus sp. from the J-BAR01 and E-MUN06 localities, respectively, in the Vallcebre Syncline. We re-examined the material concluding that they can also be attributed to M. baghensis because they show a surface covered with coalescent nodes, partially fused short fan-shaped shell units, undulating growth lines, and relatively thin eggshells (0.7-0.9 mm-thick).

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This paper is a contribution to the projects CGL201130069-C02-01,02/BTE and CGL2010-16447, subsidized by the Ministerio de Economia y Competitividad of Spain. B. Vila acknowledges support from the Ministerio de Economia y Competitividad (Subprograma Juan de la Cierva MICINN-JDC, 2011) and the Committee for Research and Exploration of the National Geographic Society. This paper is part of the unpublished PhD dissertation of AGS supported by a FI grant (2008FI-AGAUR). The Generalitat de Catalunya (Departament de Cultura) supported research and fieldwork. The authors thank Miguel Moreno-Azanza (Universidad de Zaragoza) and Geraldine Garcia (University of Poitiers) for their helpful reviews, Xabier Pereda-Suberbiola (Universidad del Pais Vasco/Euskal Herriko Unibertsitatea) for his corrections, Antoni Lacasa (Institut Estudis Ilerdencs), Lluis Ardevol (Geoplay), Carme Llompart (Universitat Autonoma de Barcelona), Josep Marmi (Institut Catala de Paleontologia), Rodrigo Gaete (Museu de la Conca Della), Josep Montane, and Ferran Albert for providing information on some localities, and the volunteers and field crew that assisted in sieving and collecting samples. We thank Patricia Perez (Museo Nacional de Ciencias Naturales) and staff members of Serveis Cientifico-Tecnics (Campus UB). Rupert Glasgow reviewed the English.


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A.G. Selles (1), B. Vila (1,2) *

(1) Institui Caiala de Paleontologia Miquel Crusafont, Carrer de l'Escola Industrial, 23, 08201, Sabadell, Spain.

(2) Grupo Aragosaurus-IUCA, Paleontologia, Facultad de Ciencias, Universidad de Zaragoza, Calle Pedro Cerbuna, 12, 50009 Zaragoza, Spain

e-mail addresses: (A.G.S.); (B.V, * corresponding author)

Received: 19 December 2013 / Accepted: 18 December 2014 / Available online: 25 March 2015

Table 1.-Nomenclature and synonymies of some of the south-Pyrenean
dinosaur localities discussed in the present work.

Locality name                        Previous names and equivalences

Presa de Sant Antoni                 "Presa de San Antonio" or Point 1
                                     of Lapparent and Aguirre (1956, p.
                                     379), "Sant Antoni-2" of Ardevol
                                     et al. (1995), Presa de Tremp
                                     after Vila et al. (2012).

Norets after Pereda-Suberbiola et    Point 2 of Lapparent and Aguirre
al. (2003)                           (1956, p. 380)

Els Nerets after Casanovas-          V-1 (Vilamitjana-1) of Ardevol et
Cladellas et al. (1985)              al. (1995)

Suterranya-Cami de Montesquiu        Point 3 of Lapparent and Aguirre
                                     (1956, p. 380)

Suterranya-Mina de lignit            Point 4 of Lapparent and Aguirre
                                     (1956, p. 380), "Suterranya" of
                                     Escaso et al. (2010)

Suterranya                           "Suterrana" of Moratalla (1993)

Suterranya-1 after Ardevol et al.    "L'Abeller" of Prieto-Marquez et
(1995)                               al. (2000),"Suterrana" of Torices
                                     et al. (in press)

Orcau-1 after Ardevol et al.         "Orcau" of Lapparent and Aguirre
(1995)                               (1956)

Orcau-2 after Ardevol et al.         "Orcau" of Llompart et al. (1984)

Orcau after Moratalla (1993)         --

Moror                                "Moro" of Brinkmann (1984)

Moror-1                              "Moro" of Vianey-Liaud and Lopez-
                                     Martinez (1997), Lopez-Martinez
                                     (2000) and Bravo and Marugan-
                                     Lobon (2013)

Serrat Pedregos after Lopez-         Cellers-2 of Lopez-Martinez (2000)
Martinez and Vicens (2012)

Montrebei after Lopez-Martinez,      misspelled "Montrebey" in Torices
(2000)                               Hernandez (2002)

Fontllonga-6 after Vianey-Liaud      "Fontllonga" of Casanovas-
and Lopez-Martinez (1997)            Cladellas and Santafe-Llopis

Fontllonga L after Moratalla         "Fontllonga" of Moratalla (1993)

Fontllonga of Casanovas et al.       "Fontllonga R" of Lopez-Martinez
(1999)                               (2003)

Santa Eulalia-1 to -4                misspelled "Santa Eularia" in
                                     Lopez-Martinez (2000)
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Title Annotation:texto en ingles
Author:Selles, A.G.; Vila, B.
Publication:Journal of Iberian Geology
Date:Jan 1, 2015
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