Geol. Mag.: page 1 of 11 c Cambridge University Press 2015 1 doi:10.1017/S0016756815000321 A new record of the pliosaur lucasi Williston, 1903 (Reptilia: ) of (Late ) 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 (). It has yielded a large variety of invertebrates but also of vertebrate taxa, represented by actinopterygians and marine including (Sauropterygia) and Mosasauroidea (Squamata). The Plesiosauria are known so far by two major clades of : the ( atlasense Buchy, 2005) and the (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 . Comparison of this specimen with other Plesiosauria shows that it belongs to Brachauchenius lucasi Williston (1903), a species previously known only from the –Turonian stages of the Western Interior Seaway of North America and in the upper 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 Owen, 1841 is also provided. Keywords: Sauropterygia, , 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 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 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 –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: 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.

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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 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

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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. The right 21 right 29.6 28.9 ramus is more complete, but the left is only partially 22 right 29.9 31.5 23 right 27.3 25.8 preserved. No suture is visible and it is impossible to 1 left 32.7 30.2 say if the splenial was preserved. The anterior part of 2 left 31.3 32.8 the mandible is large and robust, measuring 945 mm in 3 left 34.6 33.3 4 left 37.8 34.7 length (Table 1). This indicates that the whole mandible 5 left 36.6 33.3 measured roughly 1500 mm in total length. 6 left 34.5 32.0 The symphysis is of medium size, being 250 mm 7 left 34.7 37.5 8 left 35.2 31.9 length dorsally and 145 mm ventrally (Figs 2, 3; on- a 32.5 34.1 line Supplementary Material Video S1 available at b 30.6 33.3 http://journals.cambridge.org/geo), which corresponds c 30.5 33.7 d 32.6 34.3 to 26% of the preserved dentary. The maximum ramus e 31.2 34.9 height is 120 mm on the posterior part of the ramus f 27.6 32.1 and decreases slowly forward, being 68 mm high just g 33.6 29.5 h 32.6 29.3 before the symphysis. No diastema is visible between i 31.7 33.3 the last symphysal alveolus and the next. The right dentary bears 24 alveoli, 6 of which are located within the symphysis. All alveoli have roughly (5 mm length) and still located in the replacement alve- the same diameter, averaging about 32 mm × 35 mm olus, while the 7th alveolus is largest, of length 25 mm. (Table 1). The alveoli are lingually bordered by a rim It has a rounded cross-section and the enamel is orna- that extends anteriorly up to the symphysis and that mented with heavily striae all around and some branch- bears the alveoli for the replacement teeth. As in all ing striae toward the base of the crown. The teeth are other plesiosaurs, the replacement teeth are located large, posteriorly recurved rounded cones. Thanks to postero-lingually to the primary alveoli. the various longitudinal dentary fractures, we can ob- serve that the alveoli are relatively deep (50 mm). The 4.b. Dentition symphysal teeth are more closely spaced than in the remainder of the ramus, with an interval of 2–3 mm No tooth is complete, but several are partially pre- versus 5 mm. served. In the alveolus 9 of the right dentary ramus, the root of a mature tooth is eroded and broken at the base of the crown; it is 25 mm × 35 mm in diameter, 5. Discussion which permits the maximum width of mature tooth 5.a. Systematical attribution roots of this specimen to be estimated (Fig. 2). Two replacement teeth are preserved on alveoli 2 The specimen MNHN GOU 11 has been com- and 7 of the right ramus (Fig. 2). The first is very small pared with a selection of Plesiosauria for which the

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Figure 2. (Colour online) (a) Photograph, with the localization and detail of the three teeth (1–3); (b) 3D reconstruction; and (c) interpretative drawing of Brachauchenius lucasi specimen MNHN GOU 11 in dorsal view, with the numeration of the different elements measured.

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differ considerably from the large robust teeth of MNHN GOU 11. Mandibular symphysis of Poly- cotylidae usually encompasses at least 12 (Pahasa- pasaurus) and up to 20 alveoli (), much longer and more slender than the symphysis of MNHN GOU 11. Only 9 symphyseal tooth po- sitions are known in Manemergus, but again it is much more slender and elongate and never exhibits the slight spoon-shaped inflation of the MNHN GOU 11 symphysis. The Pliosauroidea have large head and mandible, massive toothed jaws and elongated mandibular sym- physis (O’Keefe, 2001), all characters that fit rather well with the mandible from Morocco. On this basis, MNHN GOU 11 is therefore referred to the Pliosauroid clade. Pliosauroids include two families, the Pliosauridae (Seeley, 1874) and the Rhomaleosauridae (Nopsca, 1928; Ketchum & Benson, 2011a). Rhomaleosauridae were unearthed from Up- per Triassic – Middle Jurassic rocks of West- ern Europe, Argentina and Canada (online Supple- mentary Material Tables S2 and S3, available at http://journals.cambridge.org/geo). This family is char- acterized by a short, reinforced and keeled spatulate symphysis (Smith & Vincent, 2010; Fig. 4), like that of the pliosaurid Andrews, 1909 from the Jur- Figure 3. (Colour online) (a) 3D reconstruction and (b) inter- assic rocks of Europe and India (see Godefroit, 1994). pretative drawing of Brachauchenius lucasi specimen MNHN This special mandible morphology tip is therefore quite GOU 11 in (1) lateral and (2) ventral views. different from MNHN GOU 11. Pliosauridae comprise at least 11 genera (see Ketchum & Benson, 2011a) of which the mandible mandible is preserved. The most recent works on the morphology is known (or can be partially deduced by subject, mainly O’Keefe (2001), Smith & Vincent the skull morphology) in Storrs & (2010), Ketchum & Benson (2010, 2011a, b), Ben- Taylor, 1996 from the lower Lower Jurassic deposits son et al. (2013) and Schumacher, Carpenter & Ever- of England; Hauffiosaurus O’Keefe, 2001 from the hart (2013) have been used for this comparative pur- upper Lower Jurassic rocks of Germany; pose (online Supplementary Material Tables S1 and S2, Owen, 1841, Sauvage, 1873, available at http://journals.cambridge.org/geo.) Lydekker, 1889, Simolestes Andrews, 1909 and Mar- Because of its long mandible, MNHN GOU 11 monectes Ketchum & Benson, 2011a from the Middle– could belong either to Pliosauroidea or to poly- Upper Jurassic rocks of mainly western Europe; Polyp- cotylid Plesiosauroidea and can reasonably be excluded tychodon Owen, 1841 from the lower Upper Cretaceous from other Plesiosauroidea (Plesiosauridae, Elasmo- deposits of England and ; Megacephalosaurus sauridae, Leptocleididae) which all possess a small and Schumacher, Carpenter & Everhart, 2013 from the gracile head as well as short mandibular symphysis. As middle Turionan rocks of Kansas; Kronosaurus Long- polycotylids have already been described from Turo- man, 1924 from the upper Lower Cretaceous deposits nian deposits of Goulmima (Bardet, Suberbiola & Jalil, of Australia and Colombia and finally Brachauchenius 2003a; Buchy, Metayer & Frey, 2005), the possibility Williston, 1903, from the lower Upper Cretaceous unit that MNHN GOU 11 could belong to this clade cannot of the Western Interior Way. be excluded and the specimen has been compared to Simolestes has a short and spatulate symphysis very taxa of both clades. different from that of MNHN GOU 11. The same oc- The Polycotylidae have a long and tapered mand- curs with the very short and plesiosauroid-like mand- ible like some Pliosauridae (e.g. Marmornectes), but ible of the basal Thalassiodracon. On the contrary, their symphysis is generally more elongated and more Hauffiosaurus, Peloneustes and Marmonectes have a slender than that of pliosauroids in general and of very long and slender mandibular symphysis includ- MNHN GOU 11 in particular (Figs 2–4). Moreover, ing around 12 pairs of teeth (Ketchum & Benson, polycotylids bear more slender teeth (Bardet, Suber- 2011b), therefore very different from MNHN GOU biola & Jalil, 2003a), even on taxa with robust teeth 11. Liopleurodon has a robust and moderately long such as Eopolycotulus (Albright III, Gillette & Titus, mandibular symphysis like that of the specimen from 2007b)orDolichorhynchops (O’Keefe, 2008), that Morocco; however, it exhibits distinct heterodonty

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Figure 4. (Colour online) Comparison between (a) Goulmima specimen MNHN GOU 11 and a selection of (b) Rhomaleosauridae, (c) Polycotylidae and (d) Pliosauridae for which the mandible is well known. (a) Drawing of Brachauchenius lucasi specimen MNHN GOU 11 in (upper) dorsal view and (lower) ventral view. (b) Drawing of cramptoni in (upper, from Cruickshank,

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(Tarlo, 1960; Noè, Smith & Walton, 2004) unlike that 5.b. Synonymy and the taxonomic problem of observed in MNHN GOU 11, which exhibits alveoli Polyptychodon Owen, 1841 roughly similar in size. As far as Pliosaurus is con- This problem has already been noted by other authors cerned, P.macromerus (Phillips, 1871), P.brachyspon- (e.g. Albright III, Gillette & Titus, 2007a; Schumacher, dylus (Owen, 1841) and P.brachydeirus (Owen, 1841) 2008) and has a special importance in our work because bear teeth with trihedral or sub-trihedral cross-sections Polyptychodon is described for outcrops of the same (Tarlo, 1960; Ketchum & Benson, 2011a; Knusten, age as our specimen. It is therefore very important to 2012), unlike the circular section of the teeth of the Mo- understand better this taxon, in particular to compare it rocco specimen. All the other species of Pliosaurus – P. correctly with other Plesiosauria. portentificus Noè, Smith & Walton, 2004; P. andrewsi Polyptychodon was described for the first time by Tarlo, 1960; P.kevani Benson et al. 2013; P.carpenteri Owen in 1841 in his Odontography from specimens Benson et al. 2013; and P.westburyensis Benson et al. from the Greensands of England. At that time, he pro- 2013 (Tarlo, 1960; Noè, Smith & Walton, 2004; Ben- posed two new species P.interruptus and P.continuus. son et al. 2013) – present a number of symphyseal teeth P.interruptus is based on a unique isolated tooth and an- (8–15 teeth) larger than that observed on the Goulmina other one included in a mandibular fragment, to which specimen (6 teeth). Kronosaurus shows only 3–4 sym- a skull fragment was later referred (Owen, 1851, pl. 4, physal teeth (Longman, 1924; Druckenmiller, 2006), fig. 1). P. continuus is based on few isolated teeth and compared to the 6 found in MNHN GOU 11. The new isolated and fragmentary postcranial material. Among genus Megacephalosaurus described by Schumacher, all this material, one tooth was chosen as the type; the Carpenter & Everhart (2013) is very similar to the spe- reason for this choice remains unclear and is not ex- cimen from Morocco, but its size is significantly larger plained by Owen. More than one century later Welles (50%). & Slaughter (1963) described a new species of Polyp- Finally, MNHN GOU 11 presents similarities to both tychodon, P.hudsoni, from Turonian deposits of Texas Polyptychodon Owen, 1841 and Brachauchenius Wil- (USA) on the basis of an incomplete skull and mand- liston, 1903. The systematical situation of Polyptycho- ible and isolated teeth, as well as (possibly) a caudal don remains problematic (see following section) and vertebra found later at the same place. These three direct comparisons cannot be made with this taxon. species are therefore known from Cretaceous (Aptian– Brachauchenius includes only the species B. lucasi Santonian) Europe for P. continuus and P. interuptus Williston, 1903, described from Turonian rocks of (Owen, 1861; Persson, 1963; Welles& Slaughter, 1963; the Western Interior Seaway of North America and Papazzoni, 2003) and Turonian North America for P. upper Barremian deposits of northern South Amer- hudsoni. It should be noted that P.interruptus has also ica (Colombia) (Hampe, 2005). It is characterized by been found in Asia (Sepkoski, 2002). Teeth bearing 21 (Carpenter, 1996) to 25 (Albright III, Gillette & a“Polyptychodon morphology” are common in the Titus, 2007a) dentary teeth. Teeth are regular in size Albian–Cenomanian outcrops of Europe (personal ob- along the series and are robust and heavily striated servation by NB). all around, with some branching of the striation to- Different problems arise from this genus. Firstly, as ward the base of the crown (Albright III, Gillette & noted by Welles & Slaughter (1963) and Schumacher Titus, 2007a). This combination of characters is com- (2008), it is based on very poor material, mainly isol- parable with that observed in MNHN GOU 11. The ated teeth. Moreover, these teeth are non-diagnostic as only minor difference observed is that the alveoli of compared to some other pliosaurs, being large cones MNHN GOU 11 occupy almost the total dentary width, with circular cross-section and striated enamel which a character not reported in any other taxon. However, are all characteristics applying to several pliosaur taxa. as the external surface of MNHN GOU 11 bones is Welles & Slaughter (1963) proposed the designation somewhat eroded, this could partly be a preservational of the skull fragment referred to P. interruptus by bias. Owen (1851) as the new type of this species. As for Based on the characters described above, the speci- most nineteenth-century palaeontological descriptions, men MNHN GOU 11 is referred to the pliosaurid genus Owen’s publications (1841, 1851) were very ambigu- Brachauchenius and, this genus being represented by ous as far as the designation of the holotype and re- only one species, de facto to Brachauchenius lucasi ferred specimens are concerned. To the isolated teeth Williston, 1903.

1996) dorsal view and (lower, from Smith & Dyke, 2008) ventral view. (c) Drawing of Dolichorhynchops herschelensis in (upper) ventral view and (lower) dorsal view (from Sato, 2005). (d) Gallardosaurus itturraldei drawing in ventral view (from Gasparini, 2009). Polyptychodon hudsoni photo in dorsal view (from Welles & Slaughter, 1963). Brachauchenius lucasi photo in dorsal view (from Albright III, Gillette & Titus, 2007a). Photographs of Simolestes vorax in (upper, from Andrews, 1913) ventral view and (lower, from Tarlo, 1960) dorsal view. Photograph of Liopleurodon ferox in (upper) dorsal view and (lower) ventral view (Andrews, 1913). Pliosaurus brachydeirus drawing in dorsal view (from Noè, Smith & Walton, 2004). Peloneustes philarchus photograph (upper, from Tarlo, 1960) in dorsal view and outline (lower, from Lydekker, 1889) in dorsal view. Scale bars: 10 cm.

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Figure 5. (Colour online) Palaeobiogeographical map of the Turonian distribution of the Plesiosauria.

originally described (also not necessarily from the is interesting to note that in their recent phylogenetic same individual), he associated several bone elements work, Benson et al. (2013) refer at least some spe- without any homologous elements of comparison; his cimens originally described as Polyptychodon to Bra- only justification was that they had the same strati- chauchenius. graphical origin. At this time, when only Polyptychodon 2) Polyptychodon is a valid genus. In this case, as had been described from Upper Cretaceous deposits of noted by Schumacher (2008), a paratype bearing dia- England, this reasoning seemed logical (another com- gnostic characters, for example the skull fragment men- mon situation in nineteenth-century palaeontological tioned by Owen (1851), must be defined. It should be descriptions). adequate to prove that isolated specimens actually be- Another problem arises from the proposition of the long to the same species, and to permit the erection of new species P. hudsoni by Welles & Slaughter (1963). a new and more rigorous diagnosis. Pending the dis- As noted by Albright III, Gillette & Titus (2007a), covery of new and more complete specimens, this is Welles & Slaughter described the new specimen from difficult in the current state of knowledge of this taxon. Texas without comparing it to pliosaurid taxa already known from Cretaceous rocks of North American, especially Brachauchenius lucasi. Further, Williston (1907) referred to this species a specimen (USNM 5.c. Palaeobiogeography and palaeoecological 2361, see Schumacher, Carpenter & Everhart, 2013) interpretations recovered from the same stratigraphical unit as their Brachauchenius lucasi was previously known only P. hudsoni specimen. It is noteworthy that P. hudsoni from several Cenomanian–Turonian outcrops from the resembles B. lucasi in having teeth with branching stri- Western Interior Seaway of North America (Williston, ations, but differs from the holotype of Polyptychodon, 1907; Carpenter, 1996; Hampe, 2005; Liggett et al. P.interruptus, which lacks branching striations. 2005; Albright III, Gillette & Titus, 2007a) and from Two possibilities can therefore be envisaged for upper Barremian deposits of northern South America Polyptychodon as follows. (Colombia) (Hampe, 2005). The description of a new 1) Polyptychodon is a nomen dubium because is it specimen of Brachauchenius lucasi in the Turonian not based on any diagnostical material. In this case, the outcrops of Morocco therefore allows the greatly en- systematical position of P. hudsoni should be revised larged palaeobiogeographical distribution of this taxon and this species, if valid, should be referred to another to North Africa (Fig. 5). Moreover, some affinities genus (possibly Brachauchenius, or to a new genus). It between North American and North African faunas

http://journals.cambridge.org Downloaded: 17 Aug 2015 IP address: 37.163.202.226 A new record of a Cretaceous pliosaur, Morocco 9 have been already noted for other vertebrate taxa, more ANDREWS, C. W. 1909. On some new Plesiosauria from the specifically teleostean fishes (Cavin et al. 2010). Oxford Clay of Peterborough. Annals and Magazine of The sedimentary unit in which MNHN GOU 11 was Natural History, London 4, 418–29. likely fossilized corresponds to a shallow marine envir- ANDREWS, C. W.1911. Description of a new plesiosaur (Ple- siosaurus capensis, sp. nov.) from the Uitenhage Beds onment related to the maximum of the Cenomanian– of Cape Colony. Annals of the South African Museum Turonian transgressive phase (e.g. Cavin, 1999). The 7, 309–22. faunal relationships between the long distant West- ANDREWS, C. W. 1913. A Descriptive Catalogue of the Mar- ern Interior Seaway and the Goulmima Basin could ine Reptiles of the Oxford Clay. Based on the Leeds Col- have been due to the occurrence of W–E-aligned cur- lection in the British Museum (Natural History), Lon- rents within the Tethys during early Late Cretaceous don. London: British Museum, 205 pp. time. These currents probably facilitated the disper- BARDET,N.,HOUSSAYE,A.,RAGE,J.-C.&PEREDA SUBER- BIOLA, X. 2008. The Cenomanian-Turonian (late Creta- sion of other contemporary marine faunas, such as the ceous) radiation of marine squamates (Reptilia): the role teleostean fish, but also other marine reptiles such as of the Mediterranean Tethys. Bulletin de la Société Géo- squamates (Caldwell & Cooper, 1999; Bardet et al. logique de France 179, 605–22. 2008). BARDET,N.,SUBERBIOLA,X.P.&JALIL, N.-E. 2003a.Anew The Turonian Goulmima outcrop is well known for polycotylid plesiosaur from the Late Cretaceous (Turo- its high diversity of marine vertebrates, including tele- nian) of Morocco. Comptes Rendus Palevol 2, 307–15. ostean fish (Cavin, 1997, 1999, 2001) and marine rep- BARDET,N.,SUBERBIOLA,X.P.&JALIL, N.-E. 2003b.Anew mosasauroid (Squamata) from the Late Cretaceous (Tur- tiles. The latter are represented by two major clades, onian) of Morocco. Comptes Rendus Palevol 2, 607–16. the Plesiosauria and the Squamata (Bardet, Suberbiola BENSON,R.B.J.,EVANS, M., SMITH,A.S.,SASSOON,J., & Jalil, 2003a, b; Buchy, 2005; Buchy, Metayer & Frey, MOORE-FAY E ,S.,KETCHUM,H.F.&FORREST, R. 2013. 2005). The Plesiosauria are the most diverse and up un- A giant pliosaurid skull from the Late Jurassic of Eng- til now were represented by three different Plesiosaur- land. PLoS ONE 8(5). oid taxa, two polycotylids (Thililua and Manemergus) BLAINVILLE DE, H. D. 1835. Description de quelques es- and an elasmosaurid (Libonectes morgani). The de- pèces de reptiles de la Califormie, précédée de l’analyse d’un système général d’Erpetologie et d’Amphibiologie. scription of this specimen belonging to the pliosaurid Nouvelles Annales du Muséum National d’Histoire Brachauchenuis lucasi allows the faunal list of the Tur- Naturelle de Paris 4, 233–96. onian Goulmima outcrops to be improved. Regarding BUCHY, M.-C. 2005. An Elasmosaur (Reptilia: Sauroptery- the small to medium size of all the other taxa, Bra- gia) from Turonian (Upper Cretaceous) of Morocco. chauchenuis lucasi was probably the top predator of Carolinea 63, 5–28. the area. BUCHY, M.-C., METAYER,F.&FREY, E. 2005. Osteology of Manemergus anguirostris n. gen. et sp., a new plesiosaur (Reptilia, Sauropterygia) from the Upper Cretaceous of Acknowledgements. NB thanks François Escuillié (Eldo- Morocco. Palaeontographica 272, 97–120. nia, Gannat, France) for the donation of the specimen and for CALDWELL,M.W.&COOPER, J. A. 1999. Redescription, providing the geographical and stratigraphical occurrence palaeobiogeography and palaeoecolgy of Coniasaurus data which were acquired during fieldwork in the Goulmima crassidens Owen, 1850 (Squamata) from the Lower area in 2001. We also thank Florent Goussard for help with Chalk (Cretaceous; Cenomanian) of SE England. Zo- the 3C treatment of the specimen and Renaud Vacant for ological Journal of the Linnean Society 127, 423–52. assistance with the preparation the specimen. CARPENTER, K. 1996. A review of short-neck plesiosaurs from the Cretaceous of the Western Interior, North America. Neues Jahrbuch fuer Geologie und Palaeon- Declaration of interest tologie. Abhandlungen 201, 259–87. CAV I N, L. 1997. Nouveaux Teleostei du gisement du Tur- None onien inférieur de Goulmima (Maroc). Compte Rendus de l’Académie des Sciences de Paris 325, 19–24. CAV I N, L. 1999. Occurrence of a juvenile teleost, Enchodus Supplementary material sp., in a fish gut content from the Upper Cretaceous of Goulmima, Morocco. Palaeontology 60, 57–72. To view supplementary material for this article, please CAV I N, L. 2001. Osteology and phylogenetic relationships visit http://dx.doi.org/10.1017/S0016756815000321 of the teleost Goulmimichthys arambourgi Cavin, 1995 from the Upper Cretaceous of Goulmima, Morocco. Ec- logae Geologicae Helvetica 133, 25–52. CAV I N,L.,TONG,H.,BOUDAD,L.,MEISTER,C.,PIUZ,A., References TABOUELLE,J.,AARAB, M., AMIOT,R.,BUFFETAUT,E., DYKE,G.,HUA,S.&LE LOEUFF, J. 2010. Vertebrate ALBRIGHT III, L. B., GILLETTE,D.D.&TITUS, A. L. 2007a. Plesiosaurs from the upper Cretaceous (Cenomanian- assemblages from the early Late Cretaceous of south- Turonian) of Souther Utah, Part 1: New eastern Morocco: An overview. Journal of African Earth records of the Pliosaur Brachauchenius lucasi. Journal Sciences 57, 391–412. of Vertebrate Paleontology 27, 31–40. CHOUBERT,G.&FAURE-MURET, A. 1962. Evolution du domaine atlasique marocain depuis les temps paléo- ALBRIGHT III, L. B., GILLETTE,D.D.&TITUS, A. L. 2007b. Plesiosaurs from the upper Cretaceous (Cenomanian- zoïques. Bulletin de la Société Géologique de France, Turonian) tropic shale of Southern Utah, Part 2: Poly- Mémoire Hors-série (Livre à la mémoire du Professeur cotylidae. Journal of Vertebrate Paleontology 27, 41–58. Paul Fallot) 1, 447–527.

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