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Implications for the Dinosaur Extinction Pattern in Europe

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Implications for the Dinosaur Extinction Pattern in Europe Cretaceous Research (2001) 22, 41–61 doi:10.1006/cres.2000.0236, available online at http://www.idealibrary.com on New dinosaur sites correlated with Upper Maastrichtian pelagic deposits in the Spanish Pyrenees: implications for the dinosaur extinction pattern in Europe *Nieves Lo´pez-Martı´nez, †Jose´ Ignacio Canudo, ‡Lluı´s Arde`vol, §,¶Xabier Pereda Suberbiola, §Xabier Orue-Etxebarria, †Gloria Cuenca-Besco´s, †Jose´ Ignacio Ruiz-Omen˜ aca, §Xabier Murelaga and 0Monique Feist *Depto. Paleontologı´a, Facultad de C.Geolo´gicas-IGE, Universidad Complutense-CSIC, 28040 Madrid, Spain †Dept. Geologı´a, Facultad de Ciencias, Universidad de Zaragoza, 50009 Zaragoza, Spain ‡GeoPlay, PO Box 12, 25620 Tremp, Lleida, Spain §Dept. Estratigrafı´a y Paleontologı´a, Facultad de Ciencias, Universidad del Paı´s Vasco-Euskal Herriko Unibertsitatea, Apdo. 644, 48080 Bilbao, Spain ¶Muse´um National d’Histoire Naturelle, Lab. Pale´ontologie, 8 rue Buffon, 75005 Paris, France 0Institut des Sciences de l’Euvolution, Universite´ Montpellier II, place E. Bataillon, 34095 Montpellier, France Revised manuscript accepted 24 September 2000 Six new dinosaurs sites have been found close to the Cretaceous/Tertiary boundary in Are´n (south-central Pyrenees, Huesca, Spain) in coastal and non-marine deposits of the Are´n and Tremp Formations. The sites contain articulated remains (skull elements, vertebrae, hind-limb bones) and isolated teeth and bones of hadrosaurids, three types of theropod teeth, one sauropod, at least seven types of eggshells (six ornithoid types and one Megaloolithidae probably from a sauropod), remains of other vertebrates, and four charophyte species. The fossil-bearing rocks have been correlated with marine sediments containing planktonic foraminifera from the uppermost Maastrichtian Abathomphalus mayaroensis Biozone. These rich and diversified dinosaur assemblages enable more accurate dating of the faunal changes that took place during the Maastrichtian in Europe and support the hypothesis of a sudden dinosaur extinction at the Cretaceous/Tertiary boundary. 2001 Academic Press K W: Cretaceous/Tertiary boundary; Maastrichtian; marine-continental correlations; dinosaurs; charophytes; Pyrenees. 1. Introduction other vertebrates (bony fishes, amphibians, lepido- saurs, turtles and crocodiles); charophytes occur only Dinosaur remains are mostly found in non-marine at Blasi 2. The fossil-bearing rocks are located on top rocks; therefore, their accurate age determination is of the Are´n Formation and in the lower part of the often problematic. This paper presents one of the rare Tremp Formation. Downdip these dinosaur localities cases where dinosaur sites can be accurately dated, can be correlated with deep marine sediments con- and the first to be directly correlated with the upper- taining planktonic foraminifera from the topmost most Cretaceous planktonic foraminiferal biozone. Maastrichtian Abathomphalus mayaroensis Biozone, Six new dinosaur-rich sites were discovered by L. near the Cretaceous/Tertiary boundary (Figure 2). Arde`vol (GeoPlay) and F. Lo´pez Olmedo (INYPSA) Other European dinosaur localities in non-marine in uppermost Cretaceous rocks on the northern limb rocks previously considered to be Maastrichtian by of the Tremp syncline near Are´n (Huesca, south- authors (e.g. Grigorescu, 1983; Buffetaut & Le central Pyrenees, Spain; Figure 1; Lo´pez-Olmedo & Loeuff, 1991; Buffetaut et al., 1997; Gheerbrant et al., Arde`vol, in press). The sites, named Blasi 1, 1b and 1997; Laurent et al., 1999; Garcia et al., 1999), need 2–5, contain articulated and scattered dinosaur bones, to be recalibrated in the light of the new chrono- teeth and eggshell fragments, together with remains of stratigraphic scale of Gradstein et al. (1995) and 0195–6671/01/010041+21 $35.00/0 2001 Academic Press 42 N. Lo´pez-Mart´ınez et al. refinements in marine-continental biostratigraphic content in Table 1. Blasi 1 is situated on top of a correlation (Riveline et al., 1996). The changes intro- 60-m-thick sandstone unit of the marine Are´n duced to the duration of the stages has led to rocks Formation, which dips 30 to the south. The sand- previously assigned to the Early Maastrichtian (Figure stone is a mixed arenite with white, coarse, well- 3) to be dated as Late Campanian, particularly those rounded quartz grains. Iron nodules and remnants of containing Septorella charophyte assemblages. iron crusts are locally present. Large dinosaur bones The dinosaur-rich Are´n sites, among the youngest and crocodilian teeth are scattered over an area of in the world, are crucial for establishing the pattern of several hundred square metres. Most of the bones are the Late Cretaceous dinosaur succession and extinc- fragmentary limb remains lacking proximal or distal tion in Europe. In particular, they contradict the ends, but there are several associated bones belonging hypothesis that dinosaur diversity decreased before to a hadrosaurid skull (Figures 5–7). their extinction, and that the extinction was earlier in Blasi 2 occurs in a 6.5-m-thick interval of grey marls Europe than in North America (Colombo, 1996; that overlie the sandstones of Blasi 1 (Figure 2). Galbrun, 1997), already challenged by Lo´pez- Nearly 5000 kg of sediment have been washed, result- Mart´ınez et al. (1998) and Casanovas et al. (1999a). ing in the extraction of vertebrate microfossils (fish, amphibians, squamates and turtles), some dinosaur 2. Geological setting and depositional teeth (theropods, hadrosaurids, a sauropod) and egg- framework shells (Tables 1, 2). Plant debris, charophytes and gastropods are common, but mammal remains have The dinosaur-bearing strata are exposed along the not been found. Blasi 1b is situated 150 m to the east northern limb of an east–west-trending syncline in an equivalent stratigraphic position. It has only (Tremp) in the south-central Pyrenean thrust unit yielded large, weathered and fragmentary dinosaur (Figure 1). This belt exposes rocks that reflect sediment bones: three dorsal vertebrae and a few rib remains deposition along the axis of an elongate foredeep basin, of a hadrosaurid, and limb bone fragments of an which deepens westward to the Atlantic Ocean. The indeterminate dinosaur (Table 1). foredeep was filled with basinal turbidites and prodelta Blasi 3 is situated in a 1-m-thick channelized shales, followed by deltaic sandstones and fluvial red sandstone body that overlies the marls of Blasi 2 beds (Are´n Sandstone and Tremp Formation, respect- (Figure 2). The fine-grained, matrix-supported feld- ively; Figure 2). These deposits are 3400 m thick and spathic sandstone contains Ophiomorpha burrows. range in age from Santonian to Maastrichtian. The site has yielded about 70 dinosaurian bones, most The sands of the Are´n Sandstone are composed of of which probably belong to a single hadrosaurid quartz with scarce feldspar, chert, quartzite, mica, and individual. They were scattered over an area of about fossil debris, and show medium- and large-scale cross 70 m2 (Tables 1, 2). All together, 19 caudal vertebrae bedding (Nagtegaal et al., 1983). These rocks are were present in an area 4 m2: eight caudal vertebrae transitionally overlain by lagoonal/marsh marls and are preserved in anatomical connection, while red beds of the lower part of the Tremp Formation additional caudal vertebrae and chevron bones are (Liebau, 1973; D´ıaz-Molina, 1987; Krauss, 1990). slightly disarticulated (Figure 4). Other hadrosaurid The Blasi 1 site is located on top of the Are´n material consists of a maxilla fragment, poorly pre- Sandstone whereas Blasi 2–5 are located within the served isolated teeth, two cervical vertebrae, several lower part of the Tremp Formation (Figures 1, 2). distal caudal centra and chevrons, several rib remains, The depositional assemblages of the Upper one ilium and two femora. Moreover, isolated teeth Cretaceous foredeep have been divided into four from a large theropod dinosaur, crocodilian bones and depositional sequences (Are´n1–4) bounded by turtle plates have also been found (Tables 1, 2). sequence boundaries H1–5(Figure 2; Arde`vol et al., Blasi 4 and Blasi 5 occur 3 km to the west, 70 m 2000). The Are´n dinosaur sites occur in delta-front, and 100 m respectively above Blasi 3 (Figure 2). Blasi lagoonal, and fluvial deposits of the Are´n 4 sequence, 4 is on top of a 0.5-m-thick, channelized, micro- dated as latest Maastrichtian (Figure 2). The chrono- conglomerate bed. It contains a few fragments of stratigraphy of the sequences and dinosaur sites is hadrosaurid bones, mainly vertebrae, as well as discussed below (Section 6). The sites are described in crocodilian and turtle remains and internal casts of ascending stratigraphic order. bivalves. Blasi 5 is in a grey mudstone intercalation within shaly red beds; it has yielded turtle plates, 3. Dinosaur sites crocodilian osteoderms and several fragments of The stratigraphic location of the fossil sites is indi- vertebrae and long bones apparently belonging to cated in the cross-section (Figure 2), and their fossil hadrosaurs (Table 1). New dinosaur sites correlated with Upper Maastrichtian pelagic deposits in the Spanish Pyrenees 43 Figure 1. Simplified geologic map of the Tremp and Ager synclines (south-central Pyrenees, Spain). Numbers refer to the stratigraphic sections in Figure 2. The vertebrate fossils from Are´n are mostly frag- sauropod (one tooth and one Megaloolithidae mentary with a high proportion of splinters, with the eggshell-type), three types of theropod teeth and six notable exceptions of the associated remains from ‘ornithoid’ eggshell-types from theropod dinosaurs Blasi 1 and 3 and the articulated remains from Blasi 3, and birds. The fossil material from the Are´n sites which indicate an absence of reworking processes and is housed in the Museo Paleontolo´gico of the rapid burial probably after necrokinesis. Conversely, Universidad de Zaragoza, Spain (MPZ). the microconglomeratic matrix of the Blasi 4 site covers the fracture surfaces of the bones, suggesting resedimentation and transport prior to final burial. 4.1. Hadrosaurid ornithopods Blasi 1 hadrosaurid. Hadrosaurid skull bones and a lower jaw with teeth have been recovered from 4.
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