A New Record of the Pliosaur Brachauchenius Lucasi Williston, 1903 (Reptilia: Sauropterygia) of Turonian (Late Cretaceous) Age, Morocco

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A New Record of the Pliosaur Brachauchenius Lucasi Williston, 1903 (Reptilia: Sauropterygia) of Turonian (Late Cretaceous) Age, Morocco Geol. Mag.: page 1 of 11 c Cambridge University Press 2015 1 doi:10.1017/S0016756815000321 A new record of the pliosaur Brachauchenius lucasi Williston, 1903 (Reptilia: Sauropterygia) of Turonian (Late Cretaceous) age, Morocco ∗ D. ANGST ‡† & N. BARDET‡ ∗ LSCE–CEA–CNRS UMR 8212 CE Saclay l’Orme des Merisiers, 91191 Gif-sur-Yvette, France ‡Sorbonne Universités CR2P CNRS-MNHN-UMPC Paris 6, Département Histoire de la Terre, Muséum National d’Histoire Naturelle, CP 38, 8 rue Buffon, F-75005 Paris (Received 12 August 2014; accepted 1 May 2015) Abstract – The site of Goulmima (south Morocco) is well known for its rich marine fauna of Tur- onian age (Late Cretaceous). It has yielded a large variety of invertebrates but also of vertebrate taxa, represented by actinopterygians and marine reptiles including Plesiosauria (Sauropterygia) and Mosasauroidea (Squamata). The Plesiosauria are known so far by two major clades of Plesiosauroidea: the Elasmosauridae (Libonectes atlasense Buchy, 2005) and the Polycotylidae (Thililua longicollis, Bardet, Suberbiola & Jalil, 2003a; Manemergus angirostris Buchy, Metayer & Frey, 2005). Here we describe a new specimen of plesiosaur found in the same outcrop, differing from those previously cited and belonging to the other large plesiosaur clade, the Pliosauroidea. Comparison of this specimen with other Plesiosauria shows that it belongs to Brachauchenius lucasi Williston (1903), a species previously known only from the Cenomanian–Turonian stages of the Western Interior Seaway of North America and in the upper Barremian succession of northern South America (Colombia). The description of this species on a contemporaneous site of North Africa significantly expands its palaeobiogeographic dis- tribution. This discovery confirms the affinities between marine faunas of the Western Interior Seaway and those of North Africa at this time, and also permits a better understanding of the palaeobiology of the Goulmima outcrop. A discussion about the systematical status of Polyptychodon Owen, 1841 is also provided. Keywords: Sauropterygia, Pliosauridae, Turonian, Morocco, palaeobiogeography. 1. Introduction Schumacher, 2008 (paratype UNSM 50136) from Tur- onian Kansas (see Ketchum & Benson, 2010 for details, The Plesiosauria are a group of large marine pred- and online Supplementary Material Table S3 available ators known from latest Triassic to the end of Late at http://journals.cambridge.org/geo). Cretaceous time, when they become extinct during the Compared to other continents, the fossil record of biological crisis of the Cretaceous–Palaeogene (K–Pg) plesiosaurs is scarce in Africa where only five valid boundary (e.g. Vincent et al. 2011). Soon after their first taxa are known. All described from Cretaceous out- appearance in the fossil record at the end of Triassic crops, these are: the leptocleidid Leptocleidus capensis time, they underwent a spectacular radiation and be- (Andrews, 1911) from Early Cretaceous (Valanginian) came widespread both in time and space, being highly South Africa (Andrews, 1911; Cruickshank, 1997); the systematically and ecologically diversified throughout elasmosaurid Zarafasaura oceanis Vincent et al. 2011 the Jurassic–Cretaceous periods and found in all con- from latest Cretaceous (Maastrichtian) Morocco (Vin- tinents (Ketchum & Benson, 2010). cent et al. 2011); and the polycotylids Thililua longicol- During Late Cretaceous time the Plesiosauria were lis Bardet, Suberbiola & Jalil, 2003a and Manemergus particularly diversified and mainly represented by Ple- anguirostris Buchy, Metayer & Frey, 2005 and the elas- siosauroidea, known by the three families Elasmosaur- mosaurid Libonectes atlasense Buchy, 2005 described idae, Polycotylidae and Leptocleididae, as well as by from the lower Upper Cretaceous (Turonian) sediments the declining Pliosauroidea, known by the Pliosauridae of the Goulmima outcrop, southern Morocco (Bardet, only (Ketchum & Benson, 2010). Cretaceous pliosaurs Suberbiola & Jalil, 2003a; Buchy, 2005; Buchy, are only represented by: Kronosaurus Longman, 1924 Metayer & Frey, 2005). from Aptian–Albian Australia and Colombia; Bra- The Goulmima outcrop today represents the richest chauchenius Williston, 1903 from the Turonian West- plesiosaur site from Africa, with the two major clades ern Interior Seaway and Barremian Colombia (Hampe, of Cretaceous plesiosaurs represented as well as the 2005); and Megacephalosaurus eulerti Schumacher, elasmosaurids and the polycotylids. Here we describe Carpenter & Everhart, 2013 (FHSM VP321) and a new plesiosaur specimen from this outcrop belong- ing to the Pliosauridae, a clade previously undescribed †Author for correspondence: [email protected] from this area. http://journals.cambridge.org Downloaded: 17 Aug 2015 IP address: 37.163.202.226 2 D. ANGST & N. BARDET Figure 1. (a) Palaeogeographical location of the Goulmima area in southern Morocco (modified from Bardet, Suberbiola & Jalil, 2003a). (b) Probable stratigraphical range (Mammites horizon) of Brachauchenius lucasi specimen MNHN GOU 11 (modified from Cavin et al. 2010). Abbreviations: FHSM – The Sternberg Museum where the specimen could potentially come from (per- of Natural History, Hays, Kansas (USA); MNHN – sonal observation by NB). Muséum National d’Histoire Naturelle, Paris (France); In the Goulmima area, the Cenomanian–Turonian UNSM – University of Nebraska State Museum, Lin- limestone bar is easily identified. It is famous for its coln, Nebraska, USA. Turonian nodulous fossiliferous levels, including am- monites as well as marine vertebrates such as bony fishes (Cavin, 2001; Cavin et al. 2010) and large marine 2. Geographical and geological context reptiles, including the basal mosasauroid Tethysaurus In North Africa, the ‘middle’ part of the Cretaceous Bardet, Suberbiola & Jalil, 2003b and the three ple- (Aptian–Turonian) deposits is represented by a clas- siosaur taxa mentioned in the previous section (see sical trilogy forming in the landscape the recognizable Bardet et al. 2008 for details). Specimen MNHN cliff ‘Hamada’ that crops out extensively SW–NE from GOU 11 was probably preserved in such one large the Gulf of Agadir (Morocco) to the Gulf of Gabès nodule, although it remains impossible to confirm (Tunisia) (Choubert & Faure-Muret, 1962). as it was purchased already prepared (NB, personal The series classically includes, from the base to observation). the top: (1) red sandstones with crossed stratific- The fossils are generally very well preserved in small ations (Ifezouane Formation); (2) lagunar gypsous to very large concretion nodules that mainly occur marly sandstones and green marls (Aoufous Forma- near the top of a Cenomanian–Turonian limestone bar tion) (these two units are grouped into the classical (Fig. 1b). The main fossiliferous levels are located in ‘Continental Intercalaire’ of the French authors or Kem the ‘Unit 4’ (lower Turonian) of Ferrandini et al. (1985), Kem beds, Aptian–Albian in age); and (3) a marine cal- now reappraised by Ettachfini & Andreu (2004) as the careous massive bar (Akrabou Formation), represent- ‘Unit T2a’ of the Akrabou Formation (middle Turo- ing the beginning of the Cenomanian–Turonian trans- nian). Ammonites are rather frequent in these levels, gression in the area. especially belonging to the genus Mammites (see Cavin The specimen MNHN GOU 11 comes from the et al. 2010 for details). Goulmima region, located in the southern slope of the The Goulmima area was the centre of a basin which High Atlas of Morocco (Fig. 1a). Although its exact underwent large subsidence during the Cenomanian– location remains unknown because this specimen was Turonian transgression. It corresponds to an open mar- purchased, fieldwork has allowed us to locate several ine carbonated platform (Fig.1) related to the maximum fossiliferous localities near the villages of Tadighourst of the Cenomanian–Turonian transgressive phase and Asfla, north of Goulmima (Er-Rachidia Province), (Ferrandini et al. 1985), with dominant Tethyan but also http://journals.cambridge.org Downloaded: 17 Aug 2015 IP address: 37.163.202.226 A new record of a Cretaceous pliosaur, Morocco 3 Central Atlantic influences (Ettachfini & Andreu, 2004; Table 1. Measurements of the different elements of Cavin et al. 2010). Brachauchenius lucasi specimen MNHN GOU 11. NA – not applicable. Maximum Maximum 3. Systematic palaeontology Element length (mm) width (mm) Plesiosauria de Blainville, 1835 Mandible Pliosauroidea (Seeley, 1874) Welles, 1943 Dentary (dorsal view) 945 NA Pliosauroidea Seeley, 1874 Dentary (ventral view) 935 NA Symphysis (dorsal view) 250 NA Brachauchenius Williston, 1903 Symphysis (ventral view) 145 NA Brachauchenius lucasi Williston, 1903 Alveoli 1 right 32.9 31.2 Holotype: USNM 4989, skull, mandible, vertebrae and 2 right 34.7 33.4 ribs; Ottawa County, Kansas (USA); Greenhorn Lime- 3 right 35.1 34.2 4 right 35.1 33.4 stone, Turonian (Schumacher & Everhart, 2005). 5 right 35.7 31.5 6 right 28.9 33.1 Referred specimen: MNHN GOU 11, an incomplete 7 right 31.2 33.2 mandible; Goulmima region, Er-Rachidia province 8 right 39.0 36.7 (Morocco); probably Akrabou Formation (NB, per- 9 right 33.7 39.5 10 right 36.4 36.8 sonal observation), middle Turonian (Ettachfini & 11 right 30.6 34.8 Andreu, 2004). 12 right 34.4 32.6 13 right 30.8 33.3 14 right 31.5 32.1 15 right 31.7 36.5 4. Description 16 right 31.4 32.4 17 right 30.1 30.1 4.a. Mandible 18 right 30.1 30.0 19 right 29.2 27.4 The mandible preserved only both dentaries, broken 20 right 31.0 31.2 into 18 parts but remaining in connection.
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