Vertebrate Assemblages from the Early Late Cretaceous of Southeastern Morocco: an Overview

Home , Acanthomorpha, Aoufous, Araripesuchus, Asteracanthus, Bahariya Formation, Calamopleurus

Journal of African Earth Sciences 57 (2010) 391–412

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Journal of African Earth Sciences

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Geological Society of Africa Presidential Review No. 16 Vertebrate assemblages from the early Late Cretaceous of southeastern Morocco: An overview

L. Cavin a,*, H. Tong b, L. Boudad c, C. Meister a, A. Piuz a, J. Tabouelle d, M. Aarab c, R. Amiot e, E. Buffetaut b, G. Dyke f, S. Hua g, J. Le Loeuff f a Dpt. de Géologie et Paléontologie, Muséum de Genève, CP 6434, 1211 Genève 6, Switzerland b CNRS, UMR 8538, Laboratoire de Géologie de l’Ecole Normale Supérieure, 24 rue Lhomond, 75231 Paris Cedex 05, France c Faculté des Sciences et Techniques, BP, 509, Boutalamine, Errachidia, Morocco d Musée Municipal, 76500 Elbeuf-sur-Seine, France e IVPP, Chinese Academy of Sciences, 142 XiZhiMenWai DaJie, Beijing 100044, China f School of Biology and Environmental Science, University College Dublin, Belﬁeld, Dublin 4, Ireland g Musée des Dinosaures, 11260 Espéraza, France article info abstract

Article history: Fossils of vertebrates have been found in great abundance in the continental and marine early Late Cre- Received 16 June 2009 taceous sediments of Southeastern Morocco for more than 50 years. About 80 vertebrate taxa have so far Received in revised form 9 December 2009 been recorded from this region, many of which were recognised and diagnosed for the first time based on Accepted 11 December 2009 specimens recovered from these sediments. In this paper, we use published data together with new field Available online 23 December 2009 data to present an updated overview of Moroccan early Late Cretaceous vertebrate assemblages. The Cre- taceous series we have studied encompasses three Formations, the Ifezouane and Aoufous Formations, Keywords: which are continental and deltaic in origin and are often grouped under the name ‘‘Kem Kem beds”, Africa and the Akrabou Formation which is marine in origin. New field observations allow us to place four Mesozoic Vertebrata recognised vertebrate clusters, corresponding to one compound assemblage and three assemblages, Palaeoecology within a general temporal framework. In particular, two ammonite bioevents characterise the lower part Palaeogeography of the Upper Cenomanian (Calycoceras guerangeri Zone) at the base of the Akrabou Formation and the upper part of the Lower Turonian (Mammites nodosoides Zone), that may extend into the Middle Turonian within the Akrabou Formation, and allow for more accurate dating of the marine sequence in the study area. We are not yet able to distinguish a specific assemblage that characterises the Ifezouane Formation when compared to the similar Aoufous Formation, and as a result we regard the oldest of the four vertebrate ‘‘assemblages” in this region to be the compound assemblage of the ‘‘Kem Kem beds”. This well-known vertebrate assemblage comprises a mixture of terrestrial (and aerial), freshwater and brackish vertebrates. The archosaur component of this fauna appears to show an intriguingly high proportion of large-bodied carnivorous taxa, which may indicate a peculiar trophic chain, although collecting biases alter this palaeontological signal. A small and restricted assemblage, the OT1 assemblage, possibly corresponds to a specific, localised ecosystem within the Kem Kem beds compound assemblage. Microfossils and facies from the Aoufous Formation, corresponding to the top of the compound assemblage, provide evidence of extremely abiotic conditions (hypersalinity), and thus of great environmental instability. At the base of the Akrabou Formation the first ammonite bioevent, Neolobites, corresponds to the onset of the marine transgression in the early Late Cenomanian while the Agoult assemblage (Late Cenomanian?) contains a variety of small fish species that have Central Tethyan affinities. Finally, the youngest Mam- mites bioevent in the late Early Turonian corresponds to a deepening of the marine environment: this sequence is isochronous with the Goulmima assemblage, a diverse collection of fish and other marine taxa, and shows affinities with taxa from the South Atlantic, the Central Tethys and the Western Interior seaway of North America, and further highlights the biogeographical importance of these North African Late Cretaceous assemblages. Ó 2009 Elsevier Ltd. All rights reserved.

* Corresponding author. Fax: +41 22 4186301. E-mail address: [email protected] (L. Cavin).

Contents

1. Introduction ...... 392 2. Geological setting...... 393 3. Vertebrate assemblages ...... 394 3.1. The compound assemblage of the Kem Kem beds ...... 394 3.1.1. Faunal list ...... 395 3.1.2. Age, palaeoenvironment and palaeogeography ...... 399 3.2. The OT1 assemblage ...... 402 3.2.1. Faunal list ...... 402 3.2.2. Age, palaeoenvironment and palaeogeography ...... 402 3.3. The marine transgression or Neolobites bioevent ...... 402 3.4. The Agoult assemblage ...... 402 3.4.1. Faunal list ...... 405 3.4.2. Age, palaeoenvironment and palaeogeography ...... 405 3.5. The Goulmima assemblage ...... 405 3.5.1. Faunal list ...... 405 3.5.2. Age, palaeoenvironment and palaeogeography ...... 405 4. Discussion and conclusions ...... 408 Acknowledgments ...... 409 References ...... 409

1. Introduction in 1952. In the late 1940s and early 1950s, Lavocat (1948, 1949, 1951, 1954a,b) published a series of notes on his palaeontological The Cretaceous deposits surrounding the northern, eastern and discoveries in this area of Morocco, with special emphasis on his southern borders of the Palaeozoic Tafilalt and Mader basins in dinosaur finds. In 1954, he published a geological synthesis of Morocco have yielded numerous vertebrate remains for more than the Kem Kem area (Lavocat, 1954b) which is still one of the best 50 years. The first discoveries of fossil vertebrates in the ‘Grès accounts of the local geology. The first record of marine fish frag- Infracénomaniens’ of the Kem Kem region were made by French ments in the Turonian of the High-Atlas and Midelt area had been geologists, including Choubert who found tooth plates of lungfish made slightly earlier by Dubar (1949) and over the following dec- and ganoid scales south of Taouz in 1938 (Choubert, 1948; Chou- ades only a few vertebrate remains from the ‘mid’ Cretaceous of bert et al., 1952). Indeed, the first illustrations of vertebrate fossils this area of Morocco were described, including isolated fish frag- from the Kem Kem beds were published by Choubert et al. (Fig. 1A) ments described by Tabaste (1963). In 1971, a German team from

Fig. 1. Plates from Choubert et al. (1952): (A) the first illustrations of vertebrate fossils from the Kem Kem beds. Fig. 2 shows a theropod tooth referable to Carcharodontosaurus (not Megalosaurus) saharicus. Fig. 3, labelled as Crocodilus sp., is probably a Spinosaurus tooth; (B) invertebrates from the Cenomanian including Neolobites vibrayeanus (Fig. 3). L. Cavin et al. / Journal of African Earth Sciences 57 (2010) 391–412 393 the University of Göttingen discovered vertebrate fossils in the 2009). In the Early Cretaceous, North Africa was extensively cov- Taouz area, including fish (Wenz, 1980, 1981) and dinosaurs ered with deltaic and brackish deposits, which were first referred (Buffetaut, 1989); over the last 20 years, local people in this part to as the ‘Continental Intercalaire’ by Kilian (1931). As a package, of Morocco have engaged in very active excavation work to collect this series of sediments has proved difficult to date because of Cretaceous fossils for commercial purposes (while Palaeozoic the absence of marine incursions, but recent attempts to do so invertebrates from the Tafilalt and Mader basins are also being ex- based on vertebrate assemblages indicate that Nigeran outcrops tracted and sold). Because of such commercial activity, beautifully of the ‘Continental Intercalaire’ [which does not comprise a Palae- preserved fossil specimens have been discovered in both the Kem ozoic component in this country, the so-called ‘Continental post- Kem beds and in the Goulmima area, but often with little associ- tassilian’ of Furon and Lombard (1964)], might be as old as late ated information about sedimentology, stratigraphy and geograph- Middle Jurassic (Rauhut and López-Arbarello, 2009). In Morocco, ical location. Moreover, because of the rudimentary techniques the ‘Continental Intercalaire’ occurs in the southeastern part of used by local collectors to remove fossils from the field, specimens the country, and apparently contains the uppermost part of the are often incomplete, and numerous specimens or fragmentary series only, which is Cenomanian in age (see below). The succes- fossils with no commercial value are often neglected and not col- sion is topped by coastal and then open marine sediments that cor- lected (McGowan and Dyke, 2009). Although important for the lo- respond to the great Cenomanian–Turonian worldwide cal economy, this collecting activity is harmful for the preservation transgression (Gale, 2000). of Moroccan palaeontological heritage and for scientific studies, In Morocco the ‘Continental Intercalaire’ – locally called the even though natural history museums worldwide have acquired ‘Kem Kem beds’ – belongs to the so-called ‘hamadas Cretaceous– Moroccan specimens from fossil dealers, allowing some heritage Tertiary palaeogeographic domain’ in contrast to other Creta- preservation and research to proceed. In any case, we know that ceous–Tertiary Plateaus in the region, including the Phosphate Pla- specimens corresponding to about 80 early Late Cretaceous verte- teau north of the Atlas Mountains and the coastal Laayoune-Bou brate taxa from this region housed in various public collections Craa Plateaus in the southwest. The hamada domain extends to have been described so far. Most of these fossils, however, were the south of the High-Atlas, mainly over the Sahara craton (Zouhri found by local people and thus have little associated field data. et al., 2008) and belongs to the Plateforme Préafricaine (Ferrandini In this paper we present the results of fieldwork sporadically et al., 1985; Ettachfini and Andreu, 2004). It comprises the Imini conducted by us over the course of 15 years, specifically the first and Goulmima narrow plateaus wedged between the Anti-Atlas stratigraphic framework for understanding the succession of and the High-Atlas and is limited to the north by the major Moroccan Cretaceous vertebrate assemblages. Although we found South-Atlasic fault, the Guir Hamada in the East, the Kem Kem Pla- few complete specimens of vertebrates during our field trips, a teau in the South, and the Draa Hamada in the southwest (Zouhri large amount of fragmentary material was collected, together with et al., 2008). In the Moroccan Hamada domain, the time interval stratigraphic and geographical observations. These data are very spanned by the known vertebrate occurrences, the Cenomanian– important because they allow us to present, for the first time, a re- Turonian, is characterised by important changes in lithofacies that gional synthesis of the succession of vertebrate assemblages in the reflect a marked shift in depositional environments. early Late Cretaceous of southeastern Morocco. It has also proved difficult to precisely define many of the Cre- taceous Formations in Southeast Morocco. In the Anti-Atlas area, Choubert (1948) recognised the ‘trilogie mésocrétacée’ with, from 2. Geological setting bottom to top, the ‘Grès Infracénomaniens’, the ‘Marnes à gypses cénomaniennes’, and the ‘Calcaires cénomano-turoniens’. Although At the beginning of the Mesozoic, the mountains of the Varisc the lowest unit had not yielded fossils at the time, Choubert (1948) orogeny located in the region corresponding to the modern Wes- regarded it as probably exclusively Albian in age and suggested tern Sahara were peneplained. At the same time rifting associated that it corresponds to the ‘Continental Intercalaire’ defined by Kil- with the opening of the Central Atlantic, at the origin of the Central ian (1931). In the middle unit, Choubert (1948) mentioned the Atlantic Magmatic Province (CAMP) 200 Ma ago, caused an uplift of occurrence of Cératodes[sic] africanus and ganoid scales (identified the Moroccan side of the rift that prevented continuous deposition by Camille Arambourg), and in the upper part the occurrence of the of Mesozoic sediments in western Morocco, ultimately leading to ammonite Neolobites vibrayeanus. Later, Choubert et al. (1952) the extensive exposures of Palaeozoic and Precambrians rocks in mentioned the recent discoveries of fish and reptiles in the ‘Grès this region (Fabre, 2005; Michard et al., 2008; Frizon de Lamotte Infracénomaniens’ by Lavocat, and noted again that this fauna et al., 2009). Although the movement of Africa relative to a fixed looked similar to the assemblage from the ‘Continental Interca- Europe was an eastward lateral displacement from about 175 Ma laire’. Interestingly, Choubert et al. (1952) observed that this caused by the opening of The Central Atlantic, continental (terres- assemblage is also similar to the Cenomanian assemblage from trial and freshwater) biogeographic affinities with South America Egypt discovered by Stromer at the beginning of the 20th Century. remained strong in this region during the whole of the Early Creta- Although they all noticed resemblances with Cenomanian assem- ceous. Because of the opening of the South Atlantic, the movement blages, Choubert and other authors, such as Lavocat, still consid- of Africa relative to Europe changed progressively into a N–S con- ered this unit to be infracenomanian in age. More recently, the vergence between ca. 92 and 46 Ma and eventually led to the sur- lowest two parts of the trilogy were gathered into the informally rection of the Atlas system, which was contemporaneously uplifted named Kem Kem beds (Sereno et al., 1996), with a lower unit (cor- from Morocco to Algeria and Tunisia from the Middle Eocene to Re- responding to the ‘Grès Infracénomaniens’) and an upper marly cent (Frizon de Lamotte et al., 2009). unit (corresponding to the ‘marnes à gypses cénomaniennes’). During the Middle and Late Jurassic, most parts of Morocco These names have been widely used ever since. were emerged as shown by series of continental and brackish In the ‘Sillon Préafricain’, along the southern slope of the High- deposits in the central and eastern High-Atlas, as well as in the Atlas, Dubar (1948) also defined three formations in the Cretaceous Middle-Atlas regions (Charrière et al., 1994, 2005). Following series which are, from bottom to top: the sandstone Ifezouane For- emergence, the first marine ingression is Early Barremian in age mation, referred to the Albian without palaeontological evidence, (Charrière et al., 2005). During the Barremian–Aptian two narrow the Aoufous Formation, mainly clayey sandstones and green marls elongated marine gulfs extended northwards along the Middle-At- with gypsum, referred to the Albian–Cenomanian, and the Akrabou las and from the western Essaouira basin (Frizon de Lamotte et al., Formation which corresponds to the Cenomanian–Turonian 394 L. Cavin et al. / Journal of African Earth Sciences 57 (2010) 391–412 limestone hamada. Sereno et al. (1996, note 21) also remarked that ented depressions running across the Tafilalt basin (Choubert et al., the underlying beds west of Goulmima consist of a marginal mar- 1952). Lavocat (1948, 1954b) then showed that the ‘Infracénoma- ine-evaporite facies characterised by alternating marl, limestone nien’ is present below the Tertiary deposits of the Guir Hamada, and gypsum that do not form two distinct, subequal subunits. Sec- thus demonstrating a connection between the basin of the ‘Sillon tions published by Ettachfini (2008) and our own observations in Préafricain’ in the north and the Taouz Basin in the south. the southeastern vicinity of the town of Goulmima indicate, how- Consequently, there is no sedimentological or palaeogeograph- ever, that the Ifezouane Formation (the lower sandstone unit) oc- ical evidence in favour of separating the early Late Cretaceous ser- curs in most of the eastern and southern outcrops of the ies located in the ‘Sillon Préafricain’ from the series located in the Cretaceous series of the ‘Sillon Préafricain’ (as already mentioned ‘Taouz Basin’ or the Kem Kem area. Following Ettachfini and And- by Choubert, 1948), although it is generally less well exposed than reu (2004), we thus propose use of the names Ifezouane, Aoufous the ‘Grès Infracénomaniens’ (or ‘lower unit’) in the Kem Kem area. and Akrabou formations to classify the Cretaceous sediments in From a palaeogeographical point of view, Choubert (1948) con- the area. We also suggest retention of the term ‘Kem Kem beds’ sidered that Moroccan Cretaceous sediments were deposited in to characterise the vertebrate assemblages of both the Ifezouane two different basins: the Atlasic basin (including the ‘Sillon Préa- and Aoufous formations as so far we have not been able to distin- fricain’), covering the present Atlas range and reaching the Atlantic guish two different sets of taxa in this package of sediments. Thus in the west, and the Taouz Basin, formed by the Taouz hamada and the expression ‘compound assemblage’ for the Kem Kem beds is the Kem Kem. He suspected that sporadic communications be- used in this paper to characterise the vertebrate taxa found in tween both basins were probable, possibly along north–south ori- these two formations. Because this succession of vertebrate assemblages was deposited in a single sedimentary basin, and because no large hiatuses are present in the deposits, we consider that they mirror the biotic and abiotic evolution that occurred in this region of northwest Africa between the Early Cenomanian and the Middle Turonian, over a timescale of around 8 million years. Localities visited by the authors during the last decade are shown in Fig. 2;inMay 2008 eight stratigraphic sections were measured (Fig. 3).

3. Vertebrate assemblages

3.1. The compound assemblage of the Kem Kem beds

The Ifezouane Formation is composed mainly of detritic sandstones with cross-stratified structures (Fig. 4B) and is the richest in terms of numbers of disarticulated vertebrate fossils, especially at localities situated in the southern part of the outcrop area (Fig. 4A). The Ifezouane Formation rests unconformably on Palaeo- zoic rocks and its base – composed of conglomerates and breccia – fossilised the peneplain. Its thickness is very variable, from 0 to 250 m, and normally decreases from the south to the north (Chou- bert, 1948). In contrast the Aoufous Formation is characterised by marls that contain intercalations of detritic sandstone and micro- conglomerates (Fig. 4D). In localities in the southern Kem Kem area, this formation has yielded only a few isolated vertebrate remains, mainly Onchopristis teeth, while localities to the north appear to be richer (Fig. 2): at the Douira locality, for example, abundant vertebrate remains are found in the upper unit, where they are systematically excavated by local collectors (Fig. 4D). The thickness of the Aoufous Formation is between 100 and 200 m; its facies and microfossil content are described below (Sec- tion 3.1.2). The vertebrate occurrences that are known from both the Ifezouane and Aoufous formations are discussed together in this paper as a single compound assemblage, that of the Kem Kem beds. Vertebrate fossils that comprise the Kem Kem compound assemblage have been collected along a series of outcrops that span some 250 km along a westwardly inclined ‘U’ around the western end of the Anti-Atlas belt (Fig. 2). The more northerly of the recognised fossiliferous outcrops have been recorded northwest of Erfoud (at the localities of Tarda and Jorf, not shown in Fig. 2. Satellite photograph (A) and simplified geological map (B) to show the Fig. 2), while to the east localities have been recorded in the vicin- localities discussed in the text (stars). 1, Tadirhoust-Asfla; 2, Tadirhoust; 3, spring of ity of the village of Douira (Takemout, Zaouia, Douira) and then ex- Goulmima; 4, road of Goulmima; 5, Ziz; 6, Takemout; 7, Zaouia; 8, Douira; 9, Jebel tend to the east (Hasi Beraber) and south, especially to the east of Al Qabla; 10, Slilim; 11, Khetitila Srhira; 12, Bou Laâlou; 13, Chaaft; 14, El Begâa; 15, the town of Taouz (El Begâa, El Begâa – Taouz). Fossiliferous out- El Begâa – Taouz; 16, Tizi Tazguart; 17, Tizi Momrad; 18, Gara Sbâa – Agoult; 19, Tazoughard – Belkasem; 20, Tazougard SW. Black stars indicate localities corre- crops have also been traced southwest (Ouzina) towards the Kem sponding to the stratigraphic column shown in Fig. 3. Kem area (Tizi Tazguart, Tizi Momrad, Gara Sbâa) and occur farther L. Cavin et al. / Journal of African Earth Sciences 57 (2010) 391–412 395

Fig. 3. Stratigraphic column measured through the eight localities in Fig. 2. south (Lavocat, 1954b); we have not yet visited this region and no complete skull has recently been reported (Dutheil and Brito, published data are available at present. 2009). In addition to O. numidus, Sereno et al. (1996) and Dutheil As mentioned by Lavocat (1954b), fossils occur in low density in (1999a) recorded seven other, less abundant, elasmobranch taxa these deposits which makes systematic excavation difficult; how- from the Kem Kem beds: Asteracanthus aegyptiacus, Distobatus nu- ever, some areas may yield rich collections of bones and bone frag- tiae, Tribodus sp., ’Lissodus’ sp., Serratolamna amonensis, Cretoxyrhi- ments, as noted initially by Lavocat (1954b, Fig. 5A) and confirmed nidae indet., and Marckgrafia lybica (Dutheil, 1999a). Fin spines of by two of us (H.T., J.T.) during a three-week series of systematic hybodonts are also rather common: Dutheil (1999a) recognised excavations conducted at several localities in the Kem Kem beds two forms that he referred to Asteracanthus and to Tribodus/‘Lisso- (May 2008). According to Lavocat (1954b), fossils are sometimes dus’. In addition, the dental plates of lungfish are abundant in the concentrated at the surface by erosion but most of these concen- Kem Kem beds (recorded from the localities of Douira, El Begâa, trations have now vanished as a result of continuous surface col- Tizi Tazguart and Tizi Momrad, among others); indeed, in his pa- lecting. At these more ephemeral localities fossils are associated pers Lavocat variously referred to these deposits by the names with the kerkoub facies (Fig. 4C), characterised at the surface by ‘grès à Onchopristis’ or ‘grès à Ceratodus’. Lungfish remains were concentrations of sandstone spheres that range from around 2 to first referred to Ceratodus humei by Tabaste (1963) – a taxon in- 10 cm in diameter (Lavocat, 1954b). cluded in the genus Protopterus by Martin (1984a) but later referred back to Ceratodus by Churcher and De Iuliis (2001) and 3.1.1. Faunal list Churcher et al. (2006) – while others have been referred to the neo- The most abundant vertebrate remains found in the Kem Kem ceratodontid ‘Neoceratodus’ africanus (Martin, 1982, 1984b). Coela- beds are large isolated rostral teeth of the sclerorhynchid shark canth skull fragments are also commonly encountered at these Onchopristis numidus (Fig. 4C). For the most part these teeth are sites (i.e., Douira, Tizi Momrad); interestingly, the one currently found isolated, but fragments of rostral cartilage, sometimes asso- recognised Kem Kem coelacanth was referred to Mawsonia lavocati ciated with rostral teeth, are also quite common and an almost by Tabaste (1963) and Wenz (1980, 1981), and represents a genus 396 L. Cavin et al. / Journal of African Earth Sciences 57 (2010) 391–412

Fig. 4. Cretaceous outcrops and facies of SE Morocco. (A) outcrops in the Ifezouane Formation, with galleries that have been dug for collecting fossils (arrows) at El Begâa; (B) cross-bedding stratiﬁcation in the Ifezouane Formation at El Begâa; (C) kerkoub facies with Onchopristis and theropod teeth (arrows), top of the Ifezouane Formation at Douira; (D) outcrops in the Aoufous Formation at Douira; (E) Akrabou Formation, showing the bank with Neolobites vibrayeanus, sea-urchins and mollusks, excavated by local researchers (arrows). The top of the Aoufous Formation lies to the bottom left under the mass of falling rocks. East of Taouz; (F) Excavation for fossils from the Agoult assemblage; (G) Close up showing siliceous beds (arrows); (H) View of the Cretaceous deposits between the cities of Errachidia and Goulmima. The reddish lowest part and the middle part covered with fallen rocks of the cliff correspond to the Aoufous Formation. The carbonate beds that top the cliff correspond to the Akrabou Formation. Note the Jurassic southern slope of the High-Atlas in the background; (I) Cliff located 10 km to the north of the town of Goulmima (spring of Goulmima), showing beds excavated by local researchers (arrows); (J) close up of outcrop in the Akrabou Formation showing the succession of limestone and marly limestone beds with nodules (arrows). These sediments sometimes contain ammonites and ﬁshes from the Goulmima assemblage.

that is otherwise only known from South America, characterising a Several taxa of Cladistia (polypterids) have been recorded from west-Gondwana Province. In agreement with this ‘South American the Aoufous Formation; two are represented by isolated pinnulae connection’, some more recently described skull remains also show from detritic beds (Dutheil, 1999a) while three others are known features characteristic of the Brazilian Cretaceous genus Axelrod- on the basis of articulated specimens from the clayey locality ichthys (Cavin and Forey, 2004). OT1 (see below). In addition, at least two semionotiforms are L. Cavin et al. / Journal of African Earth Sciences 57 (2010) 391–412 397

Fig. 5. Vertebrate macrofossils from the Kem Kem compound assemblage. (A) excavation conducted by René Lavocat (sitting on the ground) at Gara Sbâa in 1950 (Courtesy: late René Lavocat); (B and C) Stromerichthys aethiopicus, jaw fragment (B) and scales(C); (D) Concavotectum moroccensis, posterior part of a braincase in left lateral view; (E and F) Galianemys sp: first left costal plate (E); nuchal (F); (G and H) caudal vertebra of an indeterminate lithostrotian MDE-D108 in lateral (G) and dorsal (H) views. Scales bars equal 2 cm. known to occur in the Kem Kem assemblage: one is a Lepidotes-like lated scales referable to this species are common in many parts species known from undescribed skulls housed in the Natural His- of the Kem Kem area (e.g., the unknown type locality, fragments tory Museum, London (Peter Forey, personal communication, at El Begâa, Gara Sbâa, etc.). 2007), and on the basis of numerous isolated scales found in situ The halecomorph species Calamopleurus africanus was described at most of the localities visited. Some of these scales show the based on a complete articulated skull of uncertain origin collected ‘double-peg pattern’ regarded as a possible synapomorphy for ‘close to Taouz’ (Forey and Grande, 1998); during our 2008 field the family Semionotidae (Cavin et al., 2009). Indeed, the sheer size season several additional isolated fish fragments referable to of the preserved skulls and some loose scales indicates the pres- Stromerichthys aethiopicus were also recovered. This record de- ence of very large individuals, probably up to several metres in serves comment because this genus, described by Weiler (1935) length. The only other currently known semionotiform fish is the on the basis of material from Bahariya, Egypt, and now housed in gar Oniichthys falipoui (Cavin and Brito, 2001): fragments and iso- the Bayerische Staatssammlung für Paläontologie und Geologie in 398 L. Cavin et al. / Journal of African Earth Sciences 57 (2010) 391–412

München, Germany (part of the material was destroyed by the a species of the long-snouted eusuchian Thoracosaurus, T. cherifien- bombing of the Museum during World War II, but some material sis, from the Kem Kem, his fossil material was subsequently shown is still kept in the collection, in particular a fragmentary maxilla by Broin (2002) to belong to an advanced, mesosuchian-grade and one scale) (O. Rauhut and A. López-Arbarello, personal com- crocodilian described as Elosuchus cherifiensis. Buffetaut (1976) munication, 2009), is of uncertain systematic relationships. Most also reported the presence of Libycosuchus in the Kem Kem beds material of Stromerichthys from the Kem Kem beds consists of iso- on the basis of the posterior part of a skull, which, according to lated teeth and scales which have a characteristic shape (i.e., teeth Larsson and Sues (2007), belongs to another genus which they call and jaw fragments from El Begâa, Tizi Tazguart and Tizi Momrad; Hamadasuchus. This is worthy of further discussion because Hama- scales from Boudarite, Takemout, El Begâa, Tizi Tazguart and Tizi dasuchus rebouli was described by Buffetaut (1994) on the basis of Momrad). These teeth are often rather large and have a crown that a dentary, interpreted as belonging to a small, ziphodont tremato- reaches 13 mm in length in the largest specimen, shaped as a champsid while Larsson and Sues (2007) subsequently referred slightly curved cone. This curvature affects either the whole crown additional specimens to Hamadasuchus, including a well-preserved of the tooth or the tip only, while the surface bears very fine stria- skull. It appears likely that Larsson and Sues’s (2007) attribution of tions and the apex has a well-marked acrodine tip. However, one additional specimens to Hamadasuchus is erroneous and that the fragment of maxilla (Fig. 5B) containing two complete teeth and material in question should be placed in a distinct genus, appar- the base of two others is very reminiscent of the specimen de- ently belonging to the Trematochampsidae: Larsson and Sues scribed by Weiler (1935: pl. II, Figs. 17 and 23). Scales, already re- (2007) refer the skull they describe to the Peirosauridae, but this ported from this deposit by Tabaste (1963) and Dutheil (1999a), taxon is in all likelihood a junior synonym of Trematochampsidae typically have an ornamentation consisting of strong longitudinal (Buffetaut, 1988). An additional well-preserved skull (Hua et al., in ridges with a larger central one (Fig. 5C). Fragments with strong prep.) belongs to yet another taxon of trematochampsid and sup- ossifications and coarse ornamentation, slightly reminiscent of ports the idea of a diverse trematochampsid radiation in the Ceno- bones from mawsoniid coelacanths, may also be referrable to manian of Africa. The recent descriptions by Sereno and Larsson Stromerichthys based on comparisons with material described (2009) of Araripesuchus rattoides and Laganosuchus maghrebensis and figured by Weiler (1935: Fig. 2). This taxon, referred to the further increases the known diversity of crocodilians from the family Gigantodontidae by Weiler (1935), certainly deserves fur- Kem Kem beds. Indeed, Araripesuchus rattoides provides an addi- ther study. tional example of clear similarities with South American faunas; Teleosteans from the Kem Kem beds are represented by the ich- Araripesuchus was originally described from Brazil, and later from thyodectiform Cladocyclus pankowskii (Forey and Cavin, 2007; from Niger (Buffetaut and Taquet, 1979), whereas the stomatosuchid unknown type and other localities;), by the weird osteo- Laganosuchus provides another faunal element indicating similari- glossomorph Palaeonotopterus greenwoodi (Forey, 1997; Taverne ties with the Bahariya fauna of Egypt, which has yielded Stoma- and Maisey, 1999; Taverne, 2000, 2004; Cavin and Forey, 2001; tosuchus inermis (Stromer, 1925). type locality unknown but fragments recorded at least at Hasi Ber- Above all, however, the Kem Kem beds are now well known for aber), by a new taxon of notopterid (Brito et al., 2009), by the pos- their dinosaur assemblage. Lavocat (1954a) described the rebba- sibly basal ostariophysan Erfoudichthys rosae (Pittet et al., 2009; chisaurid sauropod Rebbachisaurus garasbae on the basis of a par- unknown type locality), and by the tselfatiiform Concavotectum tial skeleton which has still been only partially described. For moroccensis (Cavin and Forey, 2008; unknown type locality, frag- example, a scapula and a dorsal vertebra were figured, but Lavocat ments collected at Douira and Tizi Momrad including several (1954a, 1955b) mentioned the discovery of 11 vertebrae (including braincases from field work in 2008) (Fig. 5D). The lack of well-doc- six articulated caudal vertebrae), a sacrum, 10 ribs, a humerus and umented locality information for most teleostean fish also repre- pelvic bones at the Gara Sbâa locality between 1949 and 1952. A sents an area for future work. very large dorsal vertebra from the Kem Kem beds – from an un- Rage and Dutheil (2008) recently described a number of known locality – kept at the Musée des Dinosaures, Espéraza amphibians and squamates from the Kem Kem beds, including (MDE), is also referrable to R. garasbae, together with a large num- amphibians – the sirenid Kababisha sp., the pipid frog Oumtkoutia bers of isolated teeth. anae and non-pipids frogs – an indeterminate lizard, snakes that The presence of additional sauropod groups in the Kem Kem appear similar to Simoliophis libycus, indeterminate madtsoiids assemblage was first indicated by Russell (1996) who suggested and nigerophiids. The known turtle fauna of the Kem Kem com- the presence of andesaurine sauropods on the basis of amphypl- pound assemblage is also diverse, with four families of pleurodi- atian middle and posterior caudals, although others have sug- rans represented. Of these, euraxemydids comprise Dirqadim gested that these elements may belong instead to a schaefferi, the podocnemidids Hamadachelys escuilliei, the bothre- rebbachisaurid (Salgado et al., 2004). In addition, a single isolated mydids Galianemys whitei and G. emringeri, and the araripemydids procoelous middle caudal vertebra (MDE-D108) can be referred Araripemys sp. (Gmira, 1995; Tong and Buffetaut, 1996; Gaffney to an indeterminate lithostrotian; very similar lithostrotian cau- et al., 2002, 2006). So far, because all the described chelonian fos- dals have been reported from the Cenomanian of Sudan (Werner, sils were collected by local people and purchased by museums, the 1994; Rauhut, 1999). By way of comparison, the Bahariya locality exact locations and stratigraphic horizons of these taxa are un- in Egypt has yielded Aegyptosaurus aegyptiacus (Stromer, 1932), known. However, two large turtle shells figured by Gaffney et al. while Stromer also suggested the presence at Bahariya of a second (2006) and confidently assigned to Galianemys sp., were collected large sauropod belonging to the family Dicraeosauridae, and most at Tizi Tazguart (Fig. 2). According to the nomad who discovered recently Smith et al. (2001) described Paralititan stromeri, from the this locality more than 30 large turtle shells have been unearthed same locality. Although the currently known fossil material from at this site; during our field work, isolated plates of Galianemys the Kem Kem is too incomplete to allow identification of the Egyp- sp. were collected in Tizi Tazguart and Tizi Momrad (Fig. 5E and tian taxa, it nevertheless does demonstrate the coexistence of at F) and a few fragmentary plates of Araripemys sp. were encoun- least two sauropod families in the area and further reinforces sim- tered at El Begâa and Gara Sbâa. ilarities between the Egyptian and Moroccan assemblages. Finally, Crocodilian remains, notably isolated teeth (Larsson and Sidor, O’Leary et al. (2004) described elements from an armoured salta- 1999), are very abundant in the Kem Kem beds and indicate a high saurid from the ‘Continental Intercalaire’ in Mali, although these diversity of taxa with various dietary adaptations as well as several deposits are poorly dated and might be older than those of the forms that remain undescribed. Although Lavocat (1955a) reported Kem Kem beds (Cenomanian, see below). L. Cavin et al. / Journal of African Earth Sciences 57 (2010) 391–412 399

The remains of theropod dinosaurs are very abundant indeed in dromeus agilis was described by Sereno et al. (1996) on the basis of the Kem Kem beds, especially isolated teeth. As early as 1948, a partial skeleton. Deltadromeus, a lightly-built theropod, has vari- Lavocat reported the presence of ‘megalosaurid’ teeth in these ously been considered Coelurosauria incertae sedis by Holtz et al. deposits. Indeed, as the teeth of large carnivorous theropods are (2004), a possible noasaurid by Sereno et al. (2004), and a basal especially sought after by fossil collectors, specimens are preferen- ceratosaur by Carrano and Sampson (2008). Finally, the presence tially retained by local dealers and has lead to the observation that of the family Dromaeosauridae in the Kem Kem assemblage was theropods are over-represented in commercially obtained collec- established by Amiot et al. (2004) on the basis of isolated teeth tions of Kem Kem beds fossils (McGowan and Dyke, 2009). In spite while ornithischian dinosaurs have only been reported to date on of this collectorship effect, however, theropod diversity is still ex- the basis of their footprints (Sereno et al., 1996). tremely high in the Kem Kem beds; a number of taxa have been re- Pterosaurs are relatively common in the Kem Kem beds (Well- ported, and were likely present, but are so far known only from nhofer and Buffetaut, 1999; Kellner, 2009), although their fossils incomplete, fragmentary material. One of the first specimens to tend to be fragmentary, which is not unexpected in such detritic, be described in detail was a tibia, which according to Lavocat high-energy deposits. However, the presence of isolated teeth – (1954a), closely resembles Elaphrosaurus bambergii, from the Late abundant at some localities – and disarticulated material demon- Jurassic of Tendaguru (Janensch, 1925), a taxon now usually con- strates a large range of taxa. Wellnhofer and Buffetaut (1999) de- sidered to be a ceratosaur (Tykoski and Rowe, 2004). This tibia scribed specimens referred to at least four taxa, including needs to be re-examined in light of the confirmed presence of representatives of ?Pteranodontidae, ?Azhdarchidae, Tapejaridae abelisaurid theropods in the Kem Kem assemblage (see below). and Ornithocheiridae, while Mader and Kellner (1999) erected Most famously, large-bodied spinosaurs (Spinosauridae) are the taxon Siroccopteryx moroccanus on the basis of a jaw fragment well-represented in the Kem Kem beds, particularly again on the referred to an anhanguerid, which may in fact belong to the genus basis of isolated teeth. However, it seems that early palaeontolo- Coloborhynchus (Barrett et al., 2008). Newly discovered jaw mate- gists who worked in these sediments did not recognise these teeth rial confirms the occurrence of azhdarchids in the Kem Kem beds. as belonging to theropods because they closely resemble those of The single incomplete vertebra considered as avian by Riff et al. large crocodiles; indeed in the first published illustrations of verte- (2004) does not preserve any avian synapomorphies: although re- brate fossils from the Kem Kem beds (Choubert et al., 1952, pl. VI; cords of these taxa remain to be confirmed in the Kem Kem beds Fig. 1A), a tooth designated as ‘Crocodilus sp.’ seems rather to be compound assemblage, birds were present alongside pterosaurs from a spinosaur (Fig. 1). Taquet (1984) was the first to mention (GD, personal observation 2008). the occurrence of spinosaurids in the Kem Kem, while later, Buffe- taut (1989) described an upper jaw fragment from the Taouz area 3.1.2. Age, palaeoenvironment and palaeogeography as Spinosaurus cf. aegyptiacus, Milner (2003) referred more com- For many years the vertebrate compound assemblage of the plete Kem Kem beds jaw remains to S. cf. aegyptiacus, and Dal Sasso Kem Kem beds was regarded as Early Cretaceous in age (i.e., et al. (2005) described the upper jaw of a large individual as Spino- ‘Infracénomanien’ for French geologists), because the youngest saurus cf. aegyptiacus. In fact it seems as though two species of age of these sediments is constrained by the occurrence of the Spinosaurus may well have been present in the Kem Kem beds: Cenomanian ammonite Neolobites vibrayeanus in the overlying Russell (1996) named Spinosaurus maroccanus on the basis of an limestone (see below). However, on the basis of the known rep- isolated cervical vertebra and referred a few other isolated re- tiles, the Kem Kem compound assemblage has also been consid- mains, including a jaw fragment, to this taxon. Its status is ques- ered younger than the Tiouraren Formation of Niger (Taquet, tioned, however, as Sereno et al. (1998) and Buffetaut and Ouaja 1976; Moody and Sutcliffe, 1991; Russell, 1996), recently dated (2002) have considered Spinosaurus maroccanus, which is based to the late Middle or early Late Jurassic (Rauhut and López-Arba- on inadequate material, to be a junior synonym of S. aegyptiacus rello, 2009) and close in age to the Bahariya assemblage of Egypt Stromer, 1915. As a result the number of spinosaurid taxa present (Choubert et al., 1952; Buffetaut, 1989, 2001; Tong and Buffetaut, in the Kem Kem beds remains uncertain, the situation being fur- 1996; Russell, 1996; Sereno et al., 1996; Wellnhofer and Buffetaut, ther complicated by the presence of several tooth morphotypes 1999), particularly because of the occurrence in both assemblages (some teeth, for example, have a smooth surface whereas others of Spinosaurus and Carcharodontosaurus. This age seems to be borne are fluted); it is unclear whether this simply reflects individual var- out by more recent palaeontological data: based on the shark iation or confirms the presence of several species. assemblage of the Kem Kem beds, Sereno et al. (1996) noted strong Another large Kem Kem theropod, Sigilmassasaurus brevicollis, affinities with the Bahariya assemblage. A close relationship be- was described on the basis of a cervical vertebra by Russell tween the tselfatiiform fish Paranogmius doederleini from Bahariya, (1996), who also referred various other isolated vertebrae to this described by Weiler (1935) and re-interpreted by Taverne (2003), taxon. However the validity of Sigilmassasaurus is highly doubtful; and Concavotectum moroccensis from the Kem Kem beds (Cavin Sereno et al. (1998) are probably correct in referring Russell’s and Forey, 2008) provides additional supporting evidence for the (1996) material to Carcharodontosaurus. However, Novas et al. similar age of both assemblages. The Bahariya Formation is well- (2005) noted that the vertebrae are very different from the cervi- dated to the Early Cenomanian (Catuneanu et al., 2006) and as such cals of South American carcharodontosaurids and it is intruiging would indicate an Early Cenomanian age for the Kem Kem beds. In- that vertebrae resembling those of Sigilmassasaurus have also been deed, other similar North African vertebrate assemblages have also found alongside Carcharodontosaurus teeth at Albian localities in been considered Cenomanian in age, including the Wadi Milk For- southern Tunisia (Benton et al., 2000). mation, Sudan (Buffetaut et al., 1990; Werner, 1994), and the Ech- The large carcharodontosaurid theropod Carcharodontosaurus kar Formation, Niger Republic. Of these, the Wadi Milk assemblage saharicus, however, is well-represented in the Kem Kem beds on is thought to be Cenomanian because of the occurrence of teeth the basis of abundant isolated teeth and a well-preserved skull from the shark Asteracanthus aegyptiacus (Werner, 1994) and other (Sereno et al., 1996). This skull was recently proposed as the neo- faunal elements. In particular the dinosaurs of the Wadi Milk are type for Carcharodontosaurus by Brusatte and Sereno (2007).In similar to those known from the Bahariya and the Kem Kem beds, addition, fossil remains of abelisaurids were first reported from also reinforcing a similar age interpretation (Werner, 1994; Rau- the Kem Kem by Russell (1996) who described a dentary fragment hut, 1999). In Niger, the abelisaurids Rugops primus (Sereno et al., as cf. Majungasaurus. The presence of these theropods was also 2004) and the carcharodontosaurid Carcharodontosaurus iguidensis confirmed by Mahler (2005) on the basis of a maxilla while Delta- (Brusatte and Sereno, 2007) found in the Cenomanian Echkar 400 L. Cavin et al. / Journal of African Earth Sciences 57 (2010) 391–412

Formation, indicate taxonomic differentiation when compared ded stratification, intercalations of pinkish sand and conglomerates with the dinosaurs from the Kem Kem beds, although both thero- with quartz pebbles, and some calcareous layers containing lamel- pod families are common in both assemblages. libranchs and gastropods (Ettachfini and Andreu, 2004; Ettachfini, Microfossils found in the Aoufous Formation (see below) have 2008). Many excavations for vertebrate fossils conducted by local so far not proved helpful for dating these sediments. There is no people are located in the upper part of the Ifezouane Formation evidence at the base of the Cretaceous series in the study area (Fig. 4A). for pre-Cenomanian fossils, although this possibility cannot be Our field data indicate that the Aoufous formation is composed definitively ruled out. If we are correct, the sediments preserved of three main facies: (1) variegated (sometimes sandy) claystones in this region correspond only with the uppermost part of the Sah- (+-hcl-and marlstones?); (2) fibrous, or saccharoid, gypsum layers; aran ‘Continental Intercalaire’. Indeed, several of the formations in- and (3) dolomitic limestone. cluded in the ‘Continental Intercalaire’ in other regions of North The micro-bio-components of the Aoufous claystones were Africa have yielded older vertebrate assemblages: Late Jurassic, picked from washed and sieved (71, 125, 200 and 400 lm) sedi- or Hauterivian–Barremian, taxa from the Tiouraren Formation, Ni- ment. Most samples are sterile, but rarely they yield quite abun- ger (Sereno et al., 1994; Rauhut and López-Arbarello, 2009); ?Ap- dant organic components. Our preliminary study indicates tian taxa from the Elrhaz Formation, Niger (Taquet, 1970, 1976, mostly fish remains (scales, vertebrae, teeth) associated with other 1980; Sereno et al., 1994) (but see discussion of the age of this for- indeterminate vertebrate fragments (Fig. 6A, sample ap299, Douira mation in Benton et al., 2000); and Albian taxa from Algeria (Depé- section) within these samples. Only apatitic microfossils are pres- ret and Savornin, 1927), Tunisia (Bouaziz et al., 1988; Benton et al., ent in the claystone samples and no calcite shells are observed. 2000) and other Saharan localities (Lapparent, 1960). This absence may be due to either palaeoenvironmental or preser- As a result, the Kem Kem compound assemblage, as defined vational factors. Only a few samples with carbonaceous residues here, encompasses fossils from two formations: the lower Ifezou- (i.e., sample ap302, Douira section) contain micro rhizocretions ane Formation and the upper Aoufous Formation (Ettachfini and (i.e., preferential calcareous cementation around plant rootlets); Andreu, 2004). Although most of the vertebrate remains described quartz is frequent throughout and marcassite is sometimes pres- to date are from the lower Ifezouane Formation, the Aoufous For- ent, indicative of a reductive environment. mation also contains a number of vertebrate records; however The facies of the carbonaceous Aoufous Formation also contain many occurrences from across the Kem Kem, because they lack a depauparate fauna. Strong dolomitisation (Fig. 6B) is likely to field data, cannot be assigned to either of these formations with have eliminated any organic remains in most samples, although confidence. at Douira some do contain lumachelles consisting of monospecific We now describe the sedimentology and microfossil contents of gastropods (Fig. 6C and D). Thin sections also show abundant gas- both formations separately, and present new data for the Aoufous tropods, thick and thin-shelled ostracods, as well as rare silicified Formation. Finally we discuss the palaeoenvironmental implica- fish scales, bones (Fig. 6E), and very rare agglutinated foraminifers tions of the Kem Kem compound vertebrate assemblage in light (Fig. 6F). Ettachfini and Andreu (2004) noted the presence of undu- of our new findings. lated cryptalgal lamination within this facies, while quartz and No new information is presented for the Ifezouane Formation. dolomite are also abundant. Quartz fragments belong to two size The main facies of this unit consists of sandstones with cross-bed- classes (Fig. 6G); large (250–600 lm) sub-rounded fragments,

Fig. 6. Microfossils and facies from the Aoufous Formation. (A) microremains, mainly fish, from the Douira locality (sample ap299); (B) dolomitised carbonaceous facies (ap300). Scale bar equals 100 lm; (C and D) gastropod lumachelle (D: ap297); (E) Silicified fish scale (ap300). Scale bar equals 200 lm; (F) Agglutinated foraminifera (ap297). Scale bar equals 50 lm; (G) facies showing two size classes of quartz fragments (ap296). Scale bar equals 200 lm. For sample localities, see Fig. 3. L. Cavin et al. / Journal of African Earth Sciences 57 (2010) 391–412 401 probably of detritic origin, and small (30–50 lm), subangulous levels: fish and small aquatic reptiles at the base and theropods homometric fragments, probably of eolian origin. at the top. This hypothesis, however, requires further quantitative When present, microfaunal assemblages are of very low diver- study utilising theoretical ecological approaches because the sity but contain rather abundant skeletal grains. This pattern apparent scarcity of herbivorous taxa may instead indicate biased may have been caused by: (1) the destruction of calcareous shells collecting in the Kem Kem area (McGowan and Dyke, 2009). Be- by dissolution in claystones and subsequent dolomitisation of most cause we also know that at least two sauropod taxa were present of the carbonaceous layers; (2) extreme abiotic conditions and in the Kem Kem assemblage, any trophic network was likely more great instability of the palaeoenvironment, with the rare fossil re- complex. mains being the result of short transgressive episodes; or (3) a As shown above, the Kem Kem vertebrate compound assem- hypersaline palaeoenvironment, with rare occurrences of gastro- blage shows clear similarities with the Cenomanian Bahariya pods, ostracods and arenaceous foraminifers, known for their eury- assemblage in Egypt, the Wadi Milk assemblage in Sudan, and also haline tolerance (Fluegel, 2004). The common occurrence of with the Libyan assemblage of Draa Ubari (Nessov et al., 1998; gypsum layers also provides evidence for hypersaline conditions Rage and Cappetta, 2002), suggesting that in the early Late Creta- in the Aoufous Formation. ceous the continental faunas from western North Africa were not Our new sedimentological and micropalaeontological observa- separated by an epicontinental sea from those of eastern North tions agree with the model that the Aoufous Formation represents Africa. Dinosaurs from the Cenomanian part of the Echkar Forma- a coastal lagoon, or paralic sebkha, deposited in a medio- to supr- tion in Niger, however, are not the same as those from Sudan alittoral palaeoenvironment, as proposed by Ettachfini and Andreu and from northern Africa (Morocco, Egypt). This might be caused (2004). Indeed, the known vertebrate assemblage is also in general either by: (1) the small number of sampled taxa so far recorded agreement with reconstructed continental palaeoenvironments for from the Cenomanian of Niger; (2) variations related to differences both the Ifezouane and Aoufous formations, as evidenced by a par- in the ecosystems; (3) a slightly different age for the Nigeran local- ticularly abundant terrestrial component (i.e., a lizard, a madtsoiid ity; or (4) latitudinal differences. The latter point, however, is un- snake, dinosaurs and pterosaurs). On the other hand, the aquatic likely since the Nigeran and the Egyptian localities were almost component of these sequences – the sharks, bony fish and some at the same latitude in the early Late Cretaceous. In addition, and of the crocodiles – is indicative of freshwater or brackish environ- as we have already noted, many of the vertebrate taxa from the ments. Amongst turtles, extant podocnemidids live in lacustrine Kem Kem beds show palaeogeographical affinities with South and riverine environments while some taxa, in particular those America, including the fish (Maisey, 2000; Cavin et al., 2001; Cavin, with South American affinities (i.e., Tribodus, Calamopleurus, Clado- 2008), pipid frogs (Rage and Dutheil, 2008), turtles (Tong, 2008) cyclus, etc.) and Western Tethyan or Central Atlantic affinities (i.e., and dinosaurs. Among the Elasmobranchii, for instance, Tribodus Stromerichthys, Simoliophis), may indicate marine influences, was first described from the Santana Formation, Brazil, by Brito although no clear open marine indicators have yet been definitely and Ferreira (1989), then the genus was tentatively reported from identified. Simoliophis is a marine ‘hind-limbed’ snake restricted to Congo and Egypt (Maisey, 2000), and from Tunisia (Cuny et al., the Mediterranean Tethys, more widespread on its southern mar- 2001). This clear vicariant pattern, however, was altered by the dis- gin, but with rare occurrences to the north (Bardet et al., 2008). covery of Tribodus in the Cenomanian of Charente, France (Vullo Fossil indicators of a strictly freshwater environment for the et al., 2005; Vullo and Néraudeau, 2008a). This French occurrence, detritic unit of the Kem Kem beds include sirenid urodels and together with the occurrence of scales of Stromerichthys (Vullo and frogs. Russell (1996) proposed that the environment of this part Néraudeau, 2008b), may reflect either a vicariant event between of the sequence was likely a fluvial plain with westward drainage, Europe and Western Gondwana, or dispersal across Tethys. Clear while Dutheil (1999a) presented rose diagrams which show cross- vicariant patterns between taxa from South America and from stratified beds oriented north–south with a northward drainage. the Kem Kem beds compound assemblage are observed at the spe- Russell (1996) also suggested that the remains of coelacanths cies level: Mawsonia (cf. M. gigas and M. lavocati respectively), and other fish more than 3 m in length indicates the ‘existence of Calamopleurus (C. cylindricus and C. africanus respectively), Cladocy- substantial freshwater bodies upstream, either within the plain clus (C. gardneri and C. pankowskii respectively), Araripesuchus (A. or possibly beyond it to the west’. The presence of large bodies of rattoides and A. gomesii); and at the genus level: euraxemydids water is confirmed by our observations because of the occurrence (Euraxemys and Dirqadim respectively), podocnemidids (Brasilemys of other large fish, including Onchopristis and lungfish, the ‘Lepi- and Hamadachelys respectively), bothremydids (Cearachelys and dotes-like’ semionotiform and Concavotectum moroccensis. Of these, Galianemys respectively) (Gaffney et al., 2006; Tong, 2008), and the latter had a large buccal cavity with proportionally very long diplodocids (Limaysaurus and Rebbachisaurus respectively) (Salgado gill rakers (borne by the branchial arches) and was thus a filter fee- et al., 2004). Other vicariant patterns between both blocks of der (Cavin and Forey, 2008) reminiscent of the recent paddlefish Western Gondwana are observed, although the sister-pairs of taxa Polyodon spathula (Grande and Bemis, 1991). Indeed, the large size on both sides of the split are not well identified. These include the of the freshwater fish and crocodilians from this deposit was also Dipnoi, which shows a clear vicariant event for the Recent lepido- used as evidence for the presence of highly productive waters in sirenids, but the situation is unclear for the neoceratodontids (see mid-Cretaceous Saharan ecosystems (Russell and Paesler, 2003). Apesteguía et al., 2007; Cavin et al., 2007a; Cione et al., 2007 for Interestingly, large-bodied aquatic and terrestrial tetrapods are discussions), as well as for the Cladistia and possibly the Gingly- also common in the Kem Kem compound assemblage and thus a modi (these two lineages were present in South America until direct ecological link between large fish and large carnivorous tet- the end of the Cretaceous; Gayet et al., 2002). Nevertheless, the rapods can be proposed for ichthyophagous predators, such as sauropod and theropod assemblage of the Kem Kem beds resem- crocodiles and spinosaurid theropods. The large body sizes at- bles that known from Bahariya (Egypt) while recent work suggests tained by dinosaurs like Spinosaurus may be explained as an adap- that the theropod assemblage is similar to more or less coeval tation to catch large fish. Indeed, and as already suggested by assemblages in Brazil (Medeiros, 2006), where spinosaurids and Russell (1996), other large theropods like Carcharodontosaurus carcharodontosaurids are also abundant. The pterosaur assem- may have been attracted to the margins of the streams by the rich blage from the Kem Kem beds appears to show a fairly high diver- fish community. Russell (1996) even went so far as to suggest that sity, as do other pterosaur assemblages of similar or slightly older the major food chain linking components of the Kem Kem beds geological age (the Aptian Crato and ?Albian Santana Formations of vertebrate compound assemblage was short, with just two main Brazil and the Albian Cambridge Greensand of the UK), but the 402 L. Cavin et al. / Journal of African Earth Sciences 57 (2010) 391–412 fragmentary nature of the material makes a precise appraisal of described by most authors (e.g. the syntheses of Caron, 1985; Pre- that diversity difficult. moli Silva and Sliter, 1995; Premoli Silva and Verga, 2004; Roba- szynski and Caron, 1979) tend measure usually between 250 and 3.2. The OT1 assemblage 600 lm. Eleven different types of section are distinguished and contain small and simple (mostly rounded chambered) specimens Dutheil (1999a) recorded several fish taxa from a locality called that are difficult to determine in thin section. Nevertheless, consid- OT1, 10 km south of the village of Tafraout, preserved as articu- ering the geometric evidence (i.e. allowing for some intraspecific lated specimens in an elongate clay lens alongside plants, molluscs, test variability), the regularity of the size of the sections and the insects and crustaceans. Although situated in the upper part of unit constancy of the wall type and diagenesis, we assume that all sec- II of the Kem Kem beds, the Aoufous Formation as used here (i.e. in tions illustrated in Fig. 7A–U are the result of sections taken the same sedimentary package as the Kem Kem beds compound through Hedbergella (Asterohedbergella) asterospinosa Hamaoui, assemblage), the faunal composition of this lens is peculiar and 1964. Sections of various primitive type adults, known to be oppor- warrants further discussion. tunistic species that rapidly colonise shallow, newly-opened areas during transgressive periods (Hart and Bailey, 1979; Caron, 1983; 3.2.1. Faunal list Caron and Homewood, 1983; Hart, 1999), can however be homeo- Dutheil (1999a) described an indeterminate batoid from this morphous. For example, Hedbergella (Asterohedbergella) asterospin- locality, the cladistian Serenoichthys kemkemensis (Dutheil, osa is rarely illustrated (e.g. Abdallah and Meister, 1997; Ettachfini, 1999b) and two other, as yet unnamed, cladistians (Dutheil, 2008; Ettachfini and Andreu, 2004; Ettachfini et al., 2005; Hama- 2009), an indeterminate actinopterygian, Diplospondichthys mor- oui, 1964) and does not appear in the syntheses on planktonic eaui (Filleul and Dutheil, 2004), and Spinocaudichthys oumtkouten- foraminiferas mentioned above. Often only the most characteristic sis which is the oldest known freshwater acanthomorph (Filleul sections (with a spine) are shown; various sections through Hed- and Dutheil, 2001). Diplospondichthys in particular exhibits a bergella (Asterohedbergella) asterospinosa are figured in Fig. 7A–U. strange combination of characters, especially diplospondylous ver- In Morocco (i.e. the western High-Atlas, central High-Atlas, tebrae, which led Filleul and Dutheil (2004) to compare this species Middle-Atlas, and High Moulouya), Hedbergella (Asterohedbergella) with various taxa among halecomorphs, basal neopterygians and asterospinosa is only known from the Upper Cenomanian; its last Anguilliformes. Note also that Diplomystus sp., as well as an inde- occurrence marks the base of the ‘imprecision interval’ of the terminate clupeomorph were figured in an early publication Cenomanian/Turonian boundary (Ettachfini, 2008). (Dutheil, 1999a) but are still pending description. Rage and Dutheil The assemblage encountered here is different from the typical (2008) also reported the presence of amphibians and squamates at Rotalipora cushmani zone assemblage generally assigned by the OT1, but because they were found in the clastic sediments situated Neolobites vibrayeanus ammonite found in the same layers (see be- just below and above the clay lens containing the fishes (Dutheil, low). The low diversity of plankton and the scarcity, or absence, of personnal communication, 2009), they belong instead to the com- associated benthic foraminiferas suggests a peculiar environment pound assemblage of the Kem Kem beds as defined here and not with probable unfavourable bottom waters; oxygen and organic the OT1 assemblage sensu stricto. flux are of great ecological importance (Van der Zwaan et al., 1999). Surprisingly, macrobenthos like gastropods, bivalves and 3.2.2. Age, palaeoenvironment and palaeogeography sea-urchins are rather abundant. The OT1 locality is located within the Aoufous Formation, re- In the studied area, the first ammonite bioevent is characterised garded here as being early Cenomanian in age. The environment by Neolobites vibrayeanus (d’Orbigny) associated with the nautilids of this site has been interpreted as a quiet lake, or pool, with fast Eutrephoceras and Angulithes (see Basse and Choubert, 1959; Mei- fossilisation occurring, evidenced by the preservation of muscles ster and Rhalmi, 2002)(Fig. 7). In the Southern part of our field in most of the fish (Dutheil, 1999a). Because the placement of Dip- area, these taxa are often associated with a rich invertebrate mac- lospondichthys within actinopterygians is problematic, Spinocaud- rofauna, including bivalves, gastropods and echinoids (Fig. 4E). ichthys has been hypothesised to occupy an indeterminate This bioevent is short in time and corresponds precisely to the low- position among basal acanthomorphs, indeed the unresolved posi- er part of the Upper Cenomanian (more precisely the lower part of tions of both taxa are really due to an unexpected combination of the Calycoceras guerangeri Zone) and is recorded at Gara Sbâa, Tizi characters, it is possible that the fish assemblage of OT1 is highly Momrad, Taouz and Belkassem. This event clearly represents an endemic. early transgressive trend in the area. Indeed, this Cenomanian Neo- lobites bioevent is closely related to the outset of the Upper Ceno- 3.3. The marine transgression or Neolobites bioevent manian marine transgression which spreads on the North African craton. To the east, the sea reached the South of the Saharan re- The contact between the Aoufous Formation and the overlying gions [i.e. South Algeria (Amédro et al., 1996), South Tunisia (Mei- carbonate Akrabou Formation is generally hidden by masses of fall- ster and Abdallah, 1996, 2005) as far as Damergou in Niger en rocks. Where the contact is visible, for example to the east of the (Meister et al., 1992, 1994)]. To the west, in the ‘Sillon Préafricain’, village of Douira, the Akrabou Formation lies conformably on top of this bioevent appears not to have crossed a NE–SW line that con- the Aoufous Formation. The Akrabou Formation was studied in de- nects Tazougart (Belkassem) to Agoult (Gara Sbâa) and indicates tail by Ettachfini and Andreu (2004) and Ettachfini (2008), who dis- the possible western border of the sea at this time. In North Africa cussed its lithostratigraphy, paleontology, paleogeography, and and in the Middle East, Neolobites is closely related to shallow mar- paleoenvironments, and found evidence of four transgressive– ine environments and generally marks the beginning of the influ- regressive sequences on this open marine platform. ence of the Southern Tethys ocean. The Upper Cenomanian transgression is characterised in this area by carbonaceous facies, which have been studied using thin 3.4. The Agoult assemblage sections. Our preliminary study shows that during the Neolobites bioevent, any benthos was very scarce, calcispheres are common, Cavin and Dutheil (1999) provided a preliminary study of a and planktonic foraminifers are abundant. These taxa are generally small fish assemblage, preserved as negatives on fine-grained simple, little evolved (no keel) and of very small morphotypes. Sec- sandstone slabs, referred as ‘Daoura’. All their described fossils tions measure around 70–100 (max 150) lm, whereas the species were found about 100 km south of Erfoud, probably on the escarp- L. Cavin et al. / Journal of African Earth Sciences 57 (2010) 391–412 403

Fig. 7. Macro- and microfossils from the Neolobites bioevent (base of the Akrabou Formation). (A–U) Various sections of Hedbergella (Asterohedbergella) asterospinosa Hamaoui, 1964 from samples ap280, ap281, ap285 and ap305. Scale bars equal 25 lm (A–L) and 50 lm (M–U). For sample localities, see Fig. 3. (V and W) Neolobites vibrayeanus (d’Orbigny); (X) Angulithes sp. Scale bar equals 100 mm (V–X). ment of the Kem Kem plateau near the Oued (wadi) Daoura, dinosaur remains in the 1950s. Indeed, dinosaur remains, as well although precise data on stratigraphical and geographical prove- as other fossils typical of the compound assemblage from the the nance are lacking. Murray et al. (2007) have reported on a resem- Kem Kem beds (see above), occur in the Aoufous Formation at bling fish assemblage from southeastern Morocco, but indicate the bottom of this cliff, while the Agoult fossil assemblage is pre- differences because the Daoura assemblage is characterised by served in a series of finely laminated carbonate beds that form sandstone preservation, while specimens from the new locality the top of the hill. These fossiliferous beds are intensively exploited are preserved in carbonates. Murray and Wilson (2009) gave more by the local people; several quarries are worked along the border information about this second locality, near the village of Agoult, of the gara (Fig. 4F). Within the carbonate sequence, intercalate southeast of the town of Alnif. beds with a more siliceous matrix form nodule-like structures In May 2008 some of us (A.P., C.M., L.B., L.C.) visited the locality (Fig. 3G) and although a more precise sedimentological study is mentioned by Murray and Wilson (2009) and found that it is lo- necessary, we suspect that the fish specimens described by Cavin cated at the top of the Gara Sbâa hill, where Lavocat excavated and Dutheil (1999) come from these siliceous beds, while the 404 L. Cavin et al. / Journal of African Earth Sciences 57 (2010) 391–412

Fig. 8. Fish from the Agoult assemblage. (A) pycnodontiform indet., (B) cf. Lusitanichthys africanus; (C and D) indeterminate teleosts.

Fig. 9. Microfossils from the base of the Akrabou Formation situated between the Neolobites and the Mammites bioevents. (A) Alveolinids indet (ap307); (B) Ovalveolina? (ap307); (C) Prealveolina sp. (ap307); (D) Biconcava bentori (ap282); (E) Praebulimina sp. (ap310); (F) Dicyclina sp. (ap307); (G) Nezzazata sp. (ap282); (H) Nezzazata isabellae (ap281); (I and J) Nummofallotia apula (ap305); (K) Pseudolituonella reicheli (ap282); (L) Pseudorhipidionina casertana (ap305); (M and N) Trochospira avnimelechi (ap282); (O and P) Actinostomaria stellata (ap306); (Q and R) Likanella? sp. A, nom. prov. (ap286); (S) Genotella pfenderae (ap286); (T) Heteroporella lepina (ap305); (U) Boueina? sp. (ap305); (V) Girvanella sp. (ap305); (W) Neomeris? cf. N. circularis (ap310); (X) microfacies with vadose pisoids (ap264). Scale bars equal 50 lm (G–J and W), 100 lm (D, L–N and Q), 200 lm (A–C, F, K, O and P, T–V, and X) and 250 lm (R and S). For localization of the samples, see Fig. 3.

specimens described by Murray et al. (2007) and Murray and here the name ‘Agoult’ to refer to this assemblage, geographically Wilson (2009) come from the carbonate beds. If this hypothesis more justiﬁed than ‘Daoura’, and to avoid confusion with fossils re- is correct then Daoura, Agoult and Gara Sbâa all correspond to corded from the Gara Sbâa locality, such as Rebbachisaurus garasbae, the same locality. However, pending additional ﬁeldwork we retain which belong to the compound assemblage of the Kem Kem beds. L. Cavin et al. / Journal of African Earth Sciences 57 (2010) 391–412 405

3.4.1. Faunal list blage (Cavin and Dutheil, 1999; Murray and Wilson, 2009), but also Cavin and Dutheil (1999) reported the occurrence of a with Cenomanian assemblages from Slovenia (Cavin et al., 2000) Pycnodontidae indet., an otocephalian referrable to Clupavus afric- and Lebanon (Forey et al., 2003). Nevertheless it is also very differ- anus (Cavin, 1999a), a clupemorph referrable to aff. Spratticeps, an- ent from the Early Turonian fish assemblage from Goulmima, other referrable to a paraclupeid indet., and Rhynchodercetis at the which shows mainly South Atlantic affinities (see below). Daoura site. In addition, Murray et al. (2007) and Wilson et al. (2009) also mentioned two paraclupeids (one possibly correspond- 3.5. The Goulmima assemblage ing to the aff. Spratticeps, although this taxon is regarded as slightly more derived than the ellimmichthyiformes by Taverne, 1997), a In his explanations of the geological map of the High-Atlas, Du- Rhynchodercetis, an euteleostean and an acanthomorph, as well bar (1949) reported the occurrence of calcareous nodules with fish as a macrosemiid later assigned to Agoultichthys chattertoni by fragments associated with an ammonite assemblage in the Turo- Murray and Wilson (2009). Visiting this locality in May 2008, we nian of the Midelt area. Farther west, in the Goulmima area, the discovered further specimens of macrosemiids, pycnodontiforms, equivalent fossiliferous beds are exploited for commercial pur- clupavids, clupeomorphs, dercetids and possible tselfatiiforms poses, mostly near the villages of Tadirhoust and Asfla, north of (Fig. 8), together with plant remains and isopod crustaceans. the city of Goulmima. Interesting fossiliferous exposures also lie about 10 km east and north of Goulmima (Fig. 3I). Vertebrate re- 3.4.2. Age, palaeoenvironment and palaeogeography mains are usually preserved in calcareous nodules, either com- During our 2008 fieldtrip, a specimen of the ammonite Neolo- pletely embedded in the nodules, or with part of the body bites vibrayeanus was found at the base of the limestone sequence enclosed. Large specimens, especially marine reptiles, are some- that forms the top of the gara, about 25 m below the fossiliferous times preserved within several distinct nodules. Although fossils outcrop. This indicates an age younger than early Late Cenomanian from this assemblage can be chemically prepared by dissolving for the Agoult assemblage. On the basis of faunal comparisons be- the matrix with formic acid, in many cases the nodule core is com- tween several ‘mid’-Cretaceous assemblages, in particular the Jbel posed of siliceous material that prevents complete preparation of Tselfat assemblage first described by Arambourg (1943, 1954), Ca- specimens. Ammonites are preserved either within nodules, or lie vin and Dutheil (1999) suggested a Late Cenomanian age for both free in the marly beds (Fig. 4J). the Agoult (‘Daoura’) and the Jbel Tselfat assemblages. Rock samples collected during the 2008 fieldtrip at the base of the Akrabou 3.5.1. Faunal list Formation, in a position located under the Mammites bioevent or Chondrichthyans are represented by at least two batoids and a Goulmima assemblage (see below) but above the Neolobites bio- sclerorhynchid (Underwood et al., 2009). The other fishes include event (i.e. in the sequence containing the Agoult assemblage), pro- indeterminate pycnodontiforms preserved either as isolated small vide new information about the palaeoenvironment of this time remains found loose in the matrix of nodules and as gut contents interval. Above the Neolobites bioevent and its specific planktonic (Cavin, 1997a), or as large articulated individuals (under study by microfauna, various benthic (and rare planktic) foraminiferas as L.C.). Individual fish recorded thus far include the ichthyodectid well as stromatoporoids, solenoporaceae, corals, algae, bryozoans, Ichthyodectes bardacki, the osmeroidid Osmeroides rheris, the ara- bivalves, ostracods, serpulids, calcispheres, gastropods and echino- ripichthyid Araripichthys corytophorus (Cavin, 1997b), a small ench- derms can be seen to gradually colonise the shelf. The preliminary odontid, Enchodus sp., sometimes preserved as gut contents (Cavin, study of our thin sections shows, amongst the sections of benthic 1999b), and the pachyrhizodontid Goulmimichthys arambourgi (Ca- foraminiferas: alveolinids (Ovalveolina?, Prealveolina sp., Fig. 9A– vin, 1995, 2001; Fig. 10C), the most common vertebrate from this C), Biconcava bentori (Fig. 9D), Ammobaculites sp., Biplanata pene- assemblage. Marine reptiles are represented by two polycotylid ropliformis, bolivinids, buliminids (Fig. 9E), Chrysalidina gradata, plesiosaurs, Thililua longicollis (Bardet et al., 2003a) and Manemer- Cuneolina sp., Dicyclina schlumbergeri (Fig. 9F), discorbids, epistom- gus anguirostris (Buchy et al., 2005), as well as the mosasauroid inids, Haplophragmoides sp., Ichnusella? sp., various miliolids, Tethysaurus nopcsai (Bardet et al., 2003b), and indeterminate prot- Montcharmontia appeninica, Nezzazata spp. (Fig. 9G and H), Nezzaz- ostegid turtles (under study by H.T.) (Fig. 10A and B). atinella picardi, Nummofallotia apula (Fig. 9I and J), Peneroplis sp., Pseudolituonella reicheli (Fig. 9K), Pseudorhapydionina dubia, Pseu- 3.5.2. Age, palaeoenvironment and palaeogeography dorhipidionina casertana (Fig. 9L), Pseudotextulariella sp., Rotorbinel- Within the Akrabou Formation a second ammonite bioevent has la mesogeensis, Spiroloculina? sp., Spiroplectammina sp., Textularia been defined which corresponds to the layer containing vertebrate spp., Trochammina sp., and Trochospira avnimelechi (Fig. 9M and remains, or the Goulmima assemblage (Cavin, 1999b). This bio- N). The stromatoporoid is attributed to Actinostomaria stellata event is longer in time than the first Neolobites bioevent and is cor- (Fig. 9O and P) and the solenoporaceae to Parachaetetes asvapatii, related with the late Early Turonian, mainly with the Mammites while six species of algae (determination by M. Conrad) have been nodosoides Zone. The ammonite fauna is also quite diversified with attributed to Likanella? sp. A (Fig. 9Q and R, very similar to Likanella Mammites (Fig. 10D), Fagesia (Fig. 10E), Neoptychites, Choffaticeras, hammudai, pl. 4, Fig. 11 in Kuss, 1994), Genotella pfenderae (Konishi Nannovascoceras, and Hoplitoides (see Basse and Choubert, 1959; and Epis, 1962) Granier et al., 1991, emend. Granier and Berthou, Kennedy et al., 2008) all present. The occurrence of Romaniceras 2002 (Fig. 9S), Heteroporella lepina Praturlon 1966 (Fig. 9T), Bouein- (Yubariceras) in the upper part of the section also suggests a Middle a? sp. (Fig. 9U), Girvanella sp. (Fig. 9V), and Neomeris? cf. N. circu- Turonian age. Ammonites have been recorded from the outcrops laris Badve and Nayak, 1983 (Fig. 9W). The foraminiferal located in the northwestern part of the studied area only (Goulm- association indicates a shallow environment (complex foramini- ima and Asfla) but they have been previously reported in areas lo- fers, alveolinids, etc.) occurring within a restricted area (scarcity cated to the north (Settat area) and to the east (Rekkame area) or absence of plankton and nekton) for most of the series, and (Basse and Choubert, 1959). Thin sections sampled across the sec- the associated muddy facies (mudstone, wackestone) suggests a ond ammonite bioevent show small and poorly diversified micro- quiet environment. Samples barren of fauna but containing vadose fossil assemblages. Buliminids represent 95% of the foraminiferal pisoids (Fig. 9X), indicative of probable emersive episodes, are fauna, always with a diagenetic micritised wall. The calcispheres numerous in the El Begâa – Taouz section. (calcareous dinoflagellate cysts) collected have at least four differ- The described fish assemblage from Agoult shows affinities ent wall types, possibly corresponding to four species, while pel- with Central Tethyan faunas, in particular the Jbel Tselfat assem- lets, gastropods, small bivalves, and vertebrate fragments are also 406 L. Cavin et al. / Journal of African Earth Sciences 57 (2010) 391–412

Fig. 10. Macrofossils from the Goulmima assemblage (or Mammites bioevent), Akrabou Formation. (A and B), indeterminate protostegid turtle in dorsal (A) and left lateral (B) views; (C), nodule with fragment of the skull of a probable Goulmimichthys arambourgi; Mammites nodosoides (D) and Fagesia sp. (E). Scale bars equal 40 mm (A–C) and 50 mm (D and E). present. This poorly diversified association is representative of an the western Tethys as indicated by recent findings. Araripichthys, environment with conditions not favourable for life, indeed this for instance, was first regarded as an endemic fish from Brazil (Sil- might indicate a likely reason for the the death, and possibly for va Santos, 1985; Maisey and Blum, 1991), but its distribution is the preservation, of the fishes and other vertebrates present be- now spread as far as Venezuela (Maisey and Moody, 2001) and cause of the lack of organisms responsible of the decay of car- Mexico (Blanco and Cavin, 2003) while its stratigraphic distribu- casses. In contrast, the large numbers of small isolated remains tion extends from the Aptian–Albian (Brazil, Venezuela) to the of juvenile Enchodus led Cavin (1999b) to suggest that this environ- Turonian (Morocco, Mexico). Similarly Goulmimichthys, first recog- ment of deposition may have been used as a nursery ground for nised based on specimens from Goulmima and previously regarded that species. as restricted to that locality, now extends as far as Colombia (Para- In contrast to the Cenomanian bioevent, the Turonian Mammites mo Fonseca, personnal communication) and Mexico (Blanco and – Romaniceras (Yubariceras) bioevent is present everywhere in the Cavin, 2003). It is interesting to note that pachyrhizodontids, section except in the south (Kem Kem area) and in the south-east including Goulmimichthys, are a rather frequently encountered along a line that runs from Zelmou to Belkassem and Tarda. This family in the Early and Late Cretaceous, recorded in the Southern suggests, at least in our area, a reduction of the influence of the Atlantic, Australia and in the Boreal domain – including the Euro- open sea toward the south. Indeed, the composition of the fauna pean Chalk and northern part of the Western Interior Seaway is classic for the shallow seas of the southern Tethys; ammonites (Cumbaa and Murray, 2008) – but are not yet known from the very are quite ubiquitous and there is no evidence for Atlantic influence. rich fish assemblages of the southeastern Tethys in Lebanon. Note The Goulmima fish assemblage shows biogeographical affinities as well that the fish taxa from the compound Kem Kem vertebrate with the northern South Atlantic (Cavin et al., 2001) but also with assemblage, as well as some of the tetrapod taxa and fish from the L. Cavin et al. / Journal of African Earth Sciences 57 (2010) 391–412 407

Table 1 Overview of the vertebrate taxa found in the four assemblages.

KK OT1 Agoult Goulmima Euselachii Onchopristis numidus Indet. Batoid 1 Batoid 2 Asteracanthus aegyptiacus Distobatus nutiae Tribodus sp. ’Lissodus’ sp. Serratolamna amonensis Cretoxyrhinidae indet. Marckgraﬁa lybica Dipnoi Ceratodus humei ‘‘Neoceratodus” africanus Actinistia Mawsonia lavocati cf. Axelrodichthys Actinopterygii indet. Stromerichthys aethiopicus Diplospondichthys moreaui Cladistia Species 1 Serenoichthys kemkemensis Species 2 Species 3 Species 4 Semionotiformes cf. Lepidotes Oniichthys falipoui Macrosemiidae Agoultichthys chattertoni Halecomorphi Calamopleurus africanus Pycnodontiformes Indet. Indet. Ichthyodectiformes Cladocyclus pankowskii Ichthyodectes bardacki Osteoglossomorpha Palaeonotopterus greenwoodi Notopteridae indet. Elopomorpha Osmeroides rheris Euteleostei Euteleostei indet. Araripichthyidae Araripichthys corythophorus Pachyrhizodontidae Goulmimichthys arambourgi Otocephala Lusitanichthys africanus Erfoudichthys rosae Clupeomorpha Diplomystus sp. Paraclupeidae indet. Clupeomorph indet. Aff. Spratticeps Tselfatiiformes Concavotectum moroccensis Aulopiformes Rhynchodercetis sp. Enchodus sp. Acanthomorpha indet. Spinocaudichthys oumtkoutensis Indet. Sirenidae Kababisha sp. Pipidae Oumtkoutia anae Anura Indet. Sauria Indet. Mosasauroidea Tethysaurus nopcsai Ophidia Simoliophis libycus Madtsoiidae indet. Nigerophiidae indet. Euraxemydidae Dirqadim schaefferi Podocnemididae Hamadachelys escuilliei Bothremydidae Galianemys whitei G. emringeri Araripemydidae Araripemys sp. Protostegidae Indet. Polycotylidae Thililua longicollis Manemergus anguirostris Mesosuchian indet. Elosuchus cheriﬁensis Notosuchia Libycosuchus sp. Araripesuchus rattoides Stomatosuchidae Laganosuchus maghrebensis Trematochampsidae Hamadasuchus rebouli Indet. Rebbachisauridae Rebbachisaurus garasbae Lithostrotia Indet. Spinosauridae Spinosaurus cf. aegyptiacus Charcarodontosauridae Carcharodontosaurus saharicus Abelisauroidea cf. Majungasaurus Deltadromeus agilis Dromaeosauridae Indet. Pterosauria ?Pteranodontidae Azhdarchidae indet. Tapejaridae indet. Coloborhynchus sp. Aves Indet. 408 L. Cavin et al. / Journal of African Earth Sciences 57 (2010) 391–412

Fig. 11. Synthetic chart showing the succession of the ‘mid’-Cretaceous vertebrate assemblages in SE Morocco within a stratigraphic framework and with palaeobiogeographical afﬁnities (arrows).

Goulmima assemblage, have been considered to share close rela- the four assemblages we have discussed and Fig. 11 shows the suc- tionships with taxa from the Santana Formation (Maisey, 2000; cession of them in a stratigraphic framework. Of the known assem- Cavin et al., 2001). This Brazilian assemblage is generally considered blages, the compound fauna of the Kem Kem beds is by far the Albian in age, which casts doubt on close taxonomic compositional most diverse, but it is likely that it actually comprises several dis- similarities with northwest African faunas. An age not older than Al- tinct assemblages that we have been unable to distinguish in our bian (Pons et al., 1990), however, and even Cenomanian has been work to date. Richest diversity seems to occur in the lower Ifezou- proposed (Martill, 1993; Tong and Buffetaut, 1996) for the Santana ane Formation, but the diversity of vertebrates from the Aoufous assemblage which may go some way to explaining the similarities Formation is still poorly known. we have discussed. We argue that affinities with the probably Early Our review demonstrates that the vertebrates from the com- Cenomanian Kem Kem assemblage and the well-dated Early Turo- pound Kem Kem assemblage are represented by taxa that encom- nian Goulmima assemblage reinforces the argument for a Cenoma- pass a wide range of sizes, from amphibians and squamates nian age for the Brazilian Santana assemblage. (represented by microfossils) up to very large saurischian dino- Finally, because diversification of basal mosasauroids occurred saurs (represented by isolated fragments, bones and partially artic- on the northern margin of the Mediterranean Tethys and in the ulated skeletons). Apparent completeness of the fauna is on the Western Interior Sea during the Late Cenomanian and Early/Middle one hand very favourable for reconstructing the Kem Kem palaeo- Turonian, the occurrence at Goulmima of Thililua is one of the ecosystem, but it is important to take into account collecting bias, southernmost occurrences known for this period (Bardet et al., for instance the proportional differences observed in collections of 2008). the teeth of herbivorous and carnivorous dinosaurs (McGowan and Dyke, 2009). The Kem Kem compound assemblage also contains a clear mix of terrestrial and aquatic components, the latter mainly 4. Discussion and conclusions freshwater in origin with a few brackish and marine fossils, but no marine ingressions have been recorded within this sequence The succession of vertebrate assemblages known from south- so far. The large size of individuals of several aquatic taxa, together eastern Morocco is one of the most diverse in the world from the with the peculiar diet of some of them (e.g. planktivorous) indicate ‘mid’ Cretaceous. Assemblages contain all the main vertebrate the occurrence of large bodies of water. It is likely that a mosaic of groups from aerial, terrestrial, freshwater, and marine environ- ecosystems was probably present, but we have not been able to ments; Table 1 show an overview of the vertebrate taxa found in distinguish them so far. Indeed the OT1 assemblage may represent L. Cavin et al. / Journal of African Earth Sciences 57 (2010) 391–412 409 an example of such a localised ecosystem in which fish seem to be Acknowledgments taxonomically (even perhaps ecologically) distinct from those of the compound Kem Kem assemblage. The top of the series contain- We thank Pierre-Alain Proz (Museum, Geneva) for making the ing this assemblage, the Aoufous Formation, was certainly depos- thin sections, Prof. Roland Wernli (University of Geneva) and Dr. ited in an environment with abiotic conditions less favourable for Marc A. Conrad (Geneva) for their help in the identification of vertebrate life, and with clues of extreme abiotic conditions microfossils, and the team of volunteers from SESNE Society of El- (hypersalinity) and a great instability of the environment. beuf for assistance with excavations during the 2008 fieldtrip The ammonite Neolobites bioevent marks the beginning of a ma- (Daniel Aubry, Laurent Gauthier, Perrine Luquet, Fabien Bricongne, jor marine transgression in the early Late Cenomanian. In our study Guy Godefroy). Dr. Oliver W.M. Rauhut and Dr. Adriana López- area, however, this bioevent seems to occur in the eastern part Arbarello (München) are thanked for their thorough reviews of this only, relative to the Southern Tethys. It is associated with a shallow paper; the late Dr. René Lavocat kindly provided photographs of his marine environment in a newly opened sea (transgressive phase), fieldwork at Gara Sbâa (Fig. 4A). as indicated by abundant small and ‘primitive’ planktonic foraminiferas. Above the Neolobites bioevent, microfossils indicate a shallow internal environment within a quiet restricted area, with a References number of possibly emersive episodes. The taxonomic composition and the small size of the fishes from the Agoult assemblage also Abdallah, H., Meister, C., 1997. The Cenomanian–Turonian boundary in the Gafsa- correspond with interpretation of this quiet shallow marine pala- Chott area (southern part of central Tunisia): biostratigraphy, eoenvironment while at the same time pointing to strong palaoe- palaeoenvironments. Cretaceous Research 18, 197–236. Amédro, F., Busson, G., Cornée, A., 1996. Révision des Ammonites du Cénomanien obigeographical affinities with the central and the southern supérieur et du Turonien inférieur du Tinrhert (Sahara Algérien), implications margin of Tethys ocean. biostratigraphiques. Bulletin du Muséum d’Histoire naturelle, 4e Série, 18- The Goulmima assemblage, isochronous with the Mammites Section C, 2–3, 179–232. Amiot, R., Buffetaut, E., Tong, H., Boudad, L., Kabiri, L., 2004. Isolated theropod teeth bioevent, is indicative of a deeper marine environment during from the Cenomanian of Morocco and their palaebiogeographical significance. the late Early Turonian. Although ammonites from this Mammites Revue de Paléobiologie 9, 143–149. bioevent show no clear palaeogeographical affinities, the verte- Apesteguía, S., Agnolín, F., Claeson, K., 2007. Review of Cretaceous dipnoans from brates from the Goulmima assemblage show strong affinities with Argentina (Sarcopterygii, Dipnoi) with descriptions of new species. Revista del Museo Argentino de Ciencias Naturales 9 (1), 27–40. the South Atlantic, the Western Tethys and even the Western Inte- Arambourg, C., 1943. Notes préliminaire sur quelques Poissons fossiles nouveaux. rior Seaway in North America, which were already directly Bulletin de la Société Géologique de France, 5e Série XIII, 281–288. connected. Arambourg, C., 1954. Les poissons crétacés du Jebel Tselfat. Notes et Mémoires du Service Géologique du Maroc 118. 188 p. The succession of vertebrate assemblages described in this paper Bardet, N., Pereda Suberbiola, X., Jalil, N.-E., 2003a. A new polycotylid plesiosaur encompasses a time interval of probably less than 10 million years from the Late Cretaceous (Turonian) of Morocco. Comptes Rendus Palevol 2, (from the base of the Cenomanian to the Lower Turonian). It is 307–315. Bardet, N., Pereda Suberbiola, X., Jalil, N.-E., 2003b. A new mosasauroid (Squamata) therefore interesting to note that few vertebrate lineages can be fol- from the Late Cretaceous (Turonian) of Morocco. Comptes Rendus Palevol 2, lowed between assemblages, and is likely due to changing environ- 607–616. mental conditions in North Africa at the time. This may indicate Bardet, N., Houssay, A., Rage, J.-C., Pereda Suberbiola, X., 2008. The Cenomanian– Turonian (late Cretaceous) radiation of marine squamates (Reptilia), the role of that abiotic factors were the main driving force shaping the taxo- the Mediterranean Tethys. Bulletin de la Société Géologique de France 179, nomic composition of the assemblages. Indeed, the palaeobiogeo- 605–622. graphical affinities of Moroccan fauna are strongly constrained by Barrett, P.M., Butler, R.J., Edwards, N.P., Milner, A.R., 2008. Pterosaur distribution in time and space: an atlas. In: Hone, D.W.E., Buffetaut, E. (Eds.), paleoecological conditions, which permitted connections with the Flugsaurier: Pterosaur Papers in Honour of Peter Wellnhofer. Zitteliana west, the north and the east at different times. Deep changes in B28, pp. 61–107. the faunal composition of the vertebrate assemblages can also be Basse, E., Choubert, G., 1959. Les faunes d’ammonites du «Cénomanien–Turonien» related to the peculiar global conditions of this time interval – high de la partie orientale du domaine atlasique marocain et de ses annexes sahariennes. In: 20ème Congrès Géologique International de Mexico, 1956, sea stand, high sea temperature – which may have favoured the Symposium Crétacé 2, pp. 58–82. biological radiations of some lineages, in particular fish (Cavin Benton, M., Bouaziz, S., Buffetaut, E., Martill, D.M., Ouaja, M., Soussi, M., Trueman, C., et al., 2007b). Other groups, such as crocodiles and pterosaurs, also 2000. Dinosaurs and other fossil vertebrates from fluvial deposits in the Lower Cretaceous of southern Tunisia. Palaeogeography, Palaeoclimatology, have high taxonomic diversity in the Kem Kem beds, but further Palaeoecology 157, 227–246. study is necessary to test whether this effect corresponds to a gen- Blanco, A., Cavin, L., 2003. New teleostei from the Agua Nueava Formation uine biological event or to a sampling bias. (Turonian), Vallecillo, NE Mexico. Comptes Rendus Palevol 2, 299–306. Bouaziz, S., Buffetaut, E., Ghanmi, M., Jaeger, J.J., Martin, M., Mazin, J.M., Tong, H., Our overview further emphasises the huge potential of the ‘mid’ 1988. Nouvelles découvertes de vertébrés fossiles dans l’Albien du Sud tunisien. Cretaceous fossil vertebrate record from Southeast Morocco for Bulletin de la Société Géologique de France 4 (2), 335–339. studying past ecosystems and their evolution through time. In par- Brito, P.M., Ferreira, P.L.N., 1989. The first hybodont shark, Tribodus limae n.g, n.s.p, from the Lower Cretaceous of Chapada do Araripe (North-East Brazil). Anais da ticular, we show that assemblages, usually treated separately and Academia Brasileira de Ciências 61, 53–57. even regarded as having been accumulated in distinct basins – Brito, P.M., Dutheil, D., Meunier, F., 2009. Presence of Notopteridae (Teleostei: the ‘Sillon Préafricain’ and the Taouz Basin – actually formed in a Osteoglossomorpha) in the Kem Kem beds, Cretaceous of Morocco. In: Jalil, N.-E. (Ed.), 1st International Congress on North African Vertebrate Palaeontology, single sedimentary basin and therefore should be considered as Program & Abstracts, Marrakech, 25–27 May 2009, p. 66. presenting a continuum in time. In conclusion, our review leads Broin, F.L.de, 2002. Elosuchus, a new genus of crocodile from the Lower Cretaceous to two issues that have to be addressed in future studies: (1) it will of the North of Africa. Comptes Rendus Palevol 1, 275–285. be a challenge to correlate the southeastern and the northwestern Brusatte, S.L., Sereno, P.C., 2007. A new species of Carcharodontosaurus (Dinosauria, Theropoda) from the Cenomanian of Niger and a revision of the genus. Journal parts of the Moroccan study area with either micro- or macrofaun- of Vertebrate Paleontology 27, 902–916. as, keeping in mind that we have no palaeontological and sedimen- Buchy, M.-C., Metayer, F., Frey, E., 2005. Osteology of Manemergus anguirostris n. tological evidence of a complete profile covering the whole Late gen. et sp., a new plesiosaur (Reptilia, Sauropterygia) from the Upper Cretaceous of Morocco. Palaeontographica, Abteilung A 272, 97–120. Cenomanian and Early Turonian and (2) systematic prospecting Buffetaut, E., 1976. Der Land-Krokodilier Libycosuchus Stromer und die Familie across the whole area, combined with systematic excavations in Libycosuchidae (Crocodylia, Mesosuchia) aus der Kreide Afrikas. Mitteilungen particularly interesting spots, must be conducted in order to pro- der Bayerischen Staatssammlung für Paläontologie und Historische Geologie 16, 17–28. vide a better understanding of lateral variations between assem- Buffetaut, E., 1988. The ziphodont mesosuchian crocodile from Messel: a blages and the total composition of single sites. reassessment. Courier Forschungsinstitut Senckenberg 107, 211–221. 410 L. Cavin et al. / Journal of African Earth Sciences 57 (2010) 391–412

Buffetaut, E., 1989. New remains of the enigmatic dinosaur Spinosaurus from the Loeuff, J. (Eds.), Late Palaeozoic and Mesozoic Continental Ecosystems of SE Cretaceous of Morocco and the affinities between Spinosaurus and Baryonyx. Asia. Geological Society of London, pp. 123–137 (Special Publication 315). Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 2, 79–87. Charrière, A., Dépêche, F., Feist, M., Grambast-Fessard, N., Jaffrezo, M., Peybernès, B., Buffetaut, E., 1994. A new crocodilian from the Cretaceous of southern Morocco. Ramalho, M., 1994. Microfaunes, microflores et paléoenvironnements successifs Comptes Rendus de l’Académie des Sciences, Paris, Série II 319, 1563–1568. dans la Formation d’El Mers (Bathonien–?Callovien) du synclinal de Skoura Buffetaut, E., 2001. On the age of the mid-Cretaceous vertebrate-bearing Formations (Moyen-Atlas, Maroc). Geobios 27, 157–174. of North Africa. In: Ferre, B., Fouray, M., Tabouelle, J. (Eds.), Colloquium on the Charrière, A., Haddoumi, H., Mojon, P.O., 2005. Découverte de Jurassique supérieur Cenomanian Stage. Bulletin de la Société d’études des Sciences Naturelles du et d’un niveau marin du Barrémien dans les «couches rouges» continentales du Muséum d’Elbeuf, 2001, pp. 20–22. Haut Atlas central marocain: implications paléogéographiques et structurales. Buffetaut, E., Ouaja, M., 2002. A new specimen of Spinosaurus (Dinosauria, Comptes Rendus Palévol 4, 385–394. Theropoda) from the Lower Cretaceous of Tunisia, with remarks on the Choubert, G., 1948. Essai sur la paléogéographie du Mésocrétacé marocain. Volume evolutionary history of the Spinosauridae. Bulletin de la Société Géologique Jubilaire de la Société des Sciences Naturelles du Maroc, 307–329. de France 173 (5), 415–421. Choubert, G., Clariond, L., Hindermeyer, J., 1952. Livret-guide de l’excursion C 36. Buffetaut, E., Taquet, P., 1979. An early Cretaceous terrestrial crocodilian and the Anti-Atlas central et oriental. Congrès Géologique International, XIXe Session opening of the South Atlantic. Nature 280, 486–487. (Alger 1952), Série: Maroc, No. 11. Rabat, 89 p. Buffetaut, E., Bussert, R., Brinkmann, W., 1990. A new nonmarine vertebrate fauna in Churcher, C.S., De Iuliis, G., 2001. A new species of Protopterus and a revision of the Upper Cretaceous of northern Sudan. Berliner geowissenschaftliche Ceratodus humei (Dipnoi: Ceratodontiformes) from the Mut Formation of Abhandlungen (A) 120 (1), 183–202. eastern Dakhleh Oasis, Western Desert of Egypt. Journal of Palaeontology 44, Caron, M., 1983. La spéciation chez les Foraminifères planctiques, une réponse 305–323. adaptée aux contraintes de l’environnement. Zitteliana 10, 671–676. Churcher, C.S., De Iuliis, G., Kleindienst, M.R., 2006. A new genus for the dipnoan Caron, M., 1985. Cretaceous planktic foraminifera. In: Bolli, H.M., Sauders, J.B., species Ceratodus tuberculatus Tabaste, 1963. Geodiversitas 28 (4), 635–647. Perch-Nielsen, K. (Eds.), Plankton Stratigraphy. Cambridge University Press, Cione, A.L., Gouiric, S., Goin, F., Poiré, D., 2007. Atlantoceratodus, a new genus of Cambridge, pp. 17–86. lungfish from the upper Cretaceous of South America and Africa. Revista Museo Caron, M., Homewood, P., 1983. Evolution of early planktic foraminifers. Marine La Plata, Paleontología 10 (62), 1–12. Micropaleontology 7, 453–462. Cumbaa, S.L., Murray, A.M., 2008. New Late Cretaceous pachyrhizodontid and Carrano, M.T., Sampson, S.D., 2008. The phylogeny of Ceratosauria (Dinosauria: enchodontid fishes and associated ichthyofauna from the Northwest Territories, Theropoda). Journal of Systematic Palaeontology 6, 183–236. Canada. In: Arratia, G., Schultze, H.-P., Wilson, M.V.H. (Eds.), Mesozoic Fishes 4 – Catuneanu, O., Khalifa, M.A., Wanas, H.A., 2006. Sequence stratigraphy of the Lower Homology and Phylogeny. Verlag Dr. Friedrich Pfeil, München, Germany, pp. Cenomanian Bahariya Formation, Bahariya Oasis, Western Desert. Egypt 229–256. Sedimentary Geology 190, 121–137. Cuny, G., Suteethorn, V., Buffetaut, E., Ouaja, M., 2001. Freshwater hybodont sharks Cavin, L., 1995. Goulmimichthys arambourgi n. g., n. sp., un Pachyrhizodontidae in the Aptian–Albian of Tunisia and Thailand. In: Ferre, B., Fouray, M., Tabouelle (Actinopterygii, Teleostei) d’une nouvelle localité à nodules fossilifères du J. (Eds.), Colloquium on the Cenomanian Stage. Bulletin de la Société d’étude des Turonien inférieur marocain. Comptes Rendus de l’Académie des Sciences, Paris, Sciences Naturelles d’Elbeuf, pp. 27–29. Série IIa 321, 1049–1054. Dal Sasso, C., Maganuco, S., Buffetaut, E., Mendez, M.A., 2005. New information on Cavin, L., 1997a. Supposed and direct evidence of trophic relationships within the the skull of the enigmatic theropod Spinosaurus, with remarks on its size and fish community of the Lower Turonian of Goulmima, Morocco. Geological affinities. Journal of Vertebrate Paleontology 25 (4), 888–896. Society of Denmark, On Line Series, 1. . Timimoun (Sahara algérien). Bulletin de la Société Géologique de France 27, Cavin, L., 1997b. Nouveaux Teleostei du gisement du Turonien inférieur de 257–265. Goulmima (Maroc). Comptes Rendus de l’Académie des Sciences, Paris, Série Dubar, G., 1949. Carte géologique provisoire du Haut Atlas de Midelt, échelle au 1/ IIa 325 (7), 19–24. 200 000e. Notice explicative, Notes et Mémoires du Service Géologique du Cavin, L., 1999a. A new Clupavidae (Teleostei, Ostariophysi) from the Cenomanian Maroc, 59bis, 60 p. of Daoura (Morocco). Comptes Rendus de l’Académie des Sciences, Paris, Série Dutheil, D.B., 1999a. An overview of the freshwater fish fauna from the Kem Kem IIa 329, 689–695. beds (Late Cretaceous, Cenomanian) of southeastern Morocco. In: Mesozoic Cavin, L., 1999b. Occurrence of a juvenile teleost, Enchodus sp., in a fish gut content Fishes 2 – Systematics and Fossil Record. Friedrich Pfeil, München, pp. 553–563. from the Upper Cretaceous of Goulmima, Morocco. Special Paper in Dutheil, D.B., 1999b. The first articulated fossil cladistian, Serenoichthys Palaeontology 60, 57–72. kemkemensis, gen. et sp. nov., from the Cretaceous of Morocco. Journal of Cavin, L., 2001. Osteology and phylogenetic relationships of the teleost Vertebrate Paleontology 19 (2), 243–246. Goulmimichthys arambourgi Cavin, 1995 from the Upper Cretaceous of Dutheil, D.B., 2009. Two new short-bodies Cladistia (Actinopterii) from the Kem Goulmima, Morocco. Eclogae Geologicae Helvetiae 133, 25–52. Kem beds (Cenomanian of Morocco). In: Jalil, N.-E. (Ed.), 1st International Cavin, L., 2008. Palaeobiogeography of Cretaceous Bony Fishes (Actinistia, Dipnoi Congress on North African Vertebrate Palaeontology, Program & Abstracts, and Actinopterygii). In: Cavin, L., Longbottom, A., Richter, M. (Eds.), Fishes and Marrakech, 25–27 May 2009, p. 31. the breakup of Pangaea. Geological Society of London, pp. 165–183 (Special Dutheil, D.B., Brito, P.M., 2009. Articulated cranium of Onchopristis numidus Publication 295). (Sclerorhynchidae, Elasmobranchii) from the kem Kem beds, Morocco. In: Cavin, L., Brito, P.M., 2001. A new Lepisosteidae (Actinopterygii, Ginglymodi) from Jalil, N.-E. (Ed.), 1st International Congress on North African Vertebrate the Cretaceous of the Kem Kem beds, Southern Morocco. Bulletin de la Société Palaeontology, Program & Abstracts, Marrakech, 25–27 May 2009, p. 66. Géologique de France 172 (5), 141–150. Ettachfini, M., 2008. La transgression au passage Cénomanien–Turonien sur le Cavin, L., Dutheil, D.B., 1999. A new Cenomanian ichthyofauna from southeastern domaine atlasique marocain. Strata Série 2. 225 p. Morocco and its relationships with other early Late Cretaceous Moroccan Ettachfini, M., Andreu, B., 2004. Le Cénomanien et le Turonien de la plate-forme faunas. Geologie en Mijnbouw 78, 261–266. Préafricaine du Maroc. Cretaceous Research 25, 277–302. Cavin, L., Forey, P.L., 2001. Osteology and systematic affinities of Palaeonotopterus Ettachfini, E.M., Souhel, A., Andreu, B., Caron, M., 2005. La limite Cénomanien– greenwoodi Forey, 1997. Zoological Journal of the Linnean Society 133, 25–52. Turonien dans le Haut Atlas central, Maroc. Geobios 38, 57–68. Cavin, L., Forey, P.L., 2004. New mawsoniid coelacanth (Sarcopterygii, Actinistia) Fabre, J., 2005. Géologie du Sahara occidental et central. Tervuren African remains from the Cretaceous of the Kem Kem beds, SE Morocco. In: Tintori, A., Geoscience Collection 108. 572. Arratia, G. (Eds.), Mesozoic Fishes III – Systematics, Paleoenvironments and Ferrandini, M., Philip, J., Babinot, J.-F., Ferrandini, J., Tronchetti, G., 1985. La plate- Biodiversity. Dr Pfeil Verlag, pp. 493–506. forme carbonatée du Cénomano–Turonien de la région d’Erfoud-Errachidia Cavin, L., Forey, P.L., 2008. A new tselfatiiform teleost from the mid-Cretaceous (Sud-Est marocain), stratigraphie et paléoenvironnements. Bulletin de la (Cenomanian) of the Kem Kem beds, Southern Morocco. In: Arratia, G., Schultze, Société Géologique de France I (8), 559–564. H-P., Wilson, M.V.H. (Eds.), Mesozoic Fishes IV – Homology and Phylogeny. Dr Filleul, A., Dutheil, D.B., 2001. Spinocaudichthys oumtkoutensis, a freshwater Pfeil Verlag, pp. 199–216. acanthomorph from the Cenomanian of Morocco. Journal of Vertebrate Cavin, L., Jurkovšek, B., Kolar-Jurkovšek, T., 2000. Succession of fish assemblages in Paleontology 21 (4), 774–780. the Upper Cretaceous of the Trieste-Komen Plateau, Slovenia. Geologija 43 (2), Filleul, A., Dutheil, D.B., 2004. A peculiar diplospondylous actinopterygian fish from 165–195. the Cretaceous of Morocco. Journal of Vertebrate Paleontology 24 (2), 290–298. Cavin, L., Boudad, L., Duffaud, S., Kabiri, L., Le Lœuff, J., Rouget, I., Tong, H., 2001. Fluegel, E., 2004. Microfacies of Carbonate Rocks. Analysis, Interpretation and L’évolution paléoenvironnementale des faunes de poissons du Crétacé Application. Springer Verlag. 976 p. supérieur du bassin de Tafilalt et des régions avoisinantes (Sud-Est du Forey, P.L., 1997. A Cretaceous notopterid (Pisces, Osteoglossomorpha) from Maroc), implications paléobiogéographiques. Comptes Rendus de l’Académie Morocco. South African Journal of Science 93, 564–569. des Sciences, Paris, Série IIa 333, 677–683. Forey, P.L., Cavin, L., 2007. A new species of Cladocyclus (Teleostei, Ichthyo- Cavin, L., Suteethorn, V., Buffetaut, E., Tong, H., 2007a. A new Thai Mesozoic lungfish dectiformes) from the Cenomanian of Morocco. Palaeontologia Electronica 10 (Sarcopterygii, Dipnoi) with an insight into post-Palaeozoic dipnoan evolution. (3), 10 p. . Zoological Journal of the Linnean Society 149, 141–177. Forey, P.L., Grande, L., 1998. An African twin to the Brazilian Calamopleurus Cavin, L., Forey, P.L., Lécuyer, C., 2007b. Correlation between environment and Late (Actinopterygii, Amiidae). Zoological Journal of the Linnean Society 123, 179– Mesozoic ray-finned fish evolution. Palaeogeography, Palaeoclimatology, 195. Palaeoecology 245, 353–367. Forey, P.L., Lu, Y., Patterson, C., Davies, C.E., 2003. Fossil fishes from the Cenomanian Cavin, L., Deesri, U., Suteethorn, V., 2009. The Jurassic and Cretaceous bony fish (Upper Cretaceous) of Namoura, Lebanon. Journal of Systematic Palaeontology 1 record (Actinopterygii, Dipnoi) from Thailand. In: Buffetaut, E., Cuny, G., Le (4), 227–330. L. Cavin et al. / Journal of African Earth Sciences 57 (2010) 391–412 411

Frizon de Lamotte, D., Leturmy, P., Missenard, Y., Sami Khomsi, S., Geoffrey Ruiz, G., Martill, D.M., 1993. Fossils of the Santana and Crato Formations, Brazil. Saddiqi, O., Guillocheau, F., Michard, A., 2009. Mesozoic and Cenozoic vertical Palaeontological Association Field Guides to Fossils 5. 159 p. movements in the Atlas system (Algeria, Morocco, Tunisia): an overview. Martin, M., 1982. Nouvelles données sur la phylogénie et la systématique des Tectonophysics 475, 9–28. Dipneustes postpaléozoïques. Comptes Rendus de l’Académie des Sciences, Furon, R., Lombard, J., 1964. Geological Map of Africa: 1/500 000; Explanatory Note. Paris, Série II 294, 611–614. Natural Resources Research, vol. 3. UNESCO, Paris, 39 p. Martin, M., 1984a. Deux Lepidosirenidae (Dipnoi) crétacés du Sahara, Protopterus Gaffney, E.S., Tong, H., Meylan, P.A., 2002. Galianemys, a new side-necked turtle humei (Priem) et Protopterus protopteroides (Tabaste). Paläontologische (Pelomelusoides, Bothremydidae) from the Late Cretaceous of Morocco. Zeitschrift 58, 265–277. American Museum Novitates 3379. 20 p. Martin, M., 1984b. Révision des Arganodontidés et des Néocératodontidés (Dipnoi, Gaffney, E.S., Tong, H., Meylan, P.A., 2006. Evolution of the side-necked turtles, the Ceratodontiformes) du Crétacé africain. Neues Jahrbuch für Geologie und families Bothremydidae, Euraxemydidae, and Araripemydidae. Bulletin of the Paläontologie, Abhandlungen 169 (2), 225–260. American Museum of Natural History 300. 698 p. McGowan, A.J., Dyke, G.J., 2009. A surfeit of theropods in the Moroccan Late Gale, A.S., 2000. The Cretaceous world. In: Culver, S.J., Rawson, P.F. (Eds.), Biotic Cretaceous? Comparing diversity estimates from field data and fossil shops. Response to Global Change: The Last 145 Million Years. Cambridge University Geology 37, 843–846. Press, Cambridge, pp. 4–19. Medeiros, M.A., 2006. Large theropod teeth from the Eocenomanian of northeastern Gayet, M., Meunier, F., Werner, C., 2002. Diversification in Polypteriformes and Brazil and the occurrence of Spinosauridae. Revista Brasileira de Paleontologia special comparison with the Lepisosteiformes. Palaeontology 45 (2), 361–376. 9, 333–338. Gmira, S., 1995. Étude des chéloniens fossiles du Maroc. Cahiers de Paléontologie, Meister, C., Abdallah, H., 1996. Les ammonites du Cénomanien supérieur et du 140 p. Turonien inférieur de la région de Gafsa-Chotts, Tunisie du centre-sud. Geobios, Grande, L., Bemis, W.E., 1991. Osteology and phylogenetic relationships of fossil and Supplément au Fascicule 29 (5), 3–49. recent paddlefishes (Polyodontidae) with comments on the interrelationships Meister, C., Abdallah, H., 2005. Précision sur les successions d’ammonites du of Acipenseriformes. Society of Vertebrate Paleontology, Supplement to Journal Cénomanien – Turonien dans la région de Gafsa, Tunisie du centre-sud. Revue of Vertebrate Paleontology 11 (1). 121 p. de Paléobiologie 24 (1), 111–199. Hamaoui, M., 1964. On a new subgenus of Hedbergella (Foraminiferida). Israel Meister, C., Rhalmi, M., 2002. Quelques ammonites du Cénomanien–Turonien de la Journal of Earth-Sciences 13, 133–142. région d’Errachidia-Boudnid-Erfoud (partie méridionale du Haut Atlas Central, Hart, M.B., 1999. The evolution and biodiversity of Cretaceous planktonic Maroc). Revue de Paléobiologie 21 (2), 759–779. Foraminiferida. Geobios 32 (2), 247–255. Meister, C., Alzouma, K., Lang, J., Mathey, B., 1992. Les ammonites du Niger (Afrique Hart, M.B., Bailey, H.W., 1979. The distribution of planktonic Foraminiferida in the occidentale) et la transgression transsaharienne au cours du Cénomanien– mid-Cretaceous of N.W. Europe. Aspekte der Kreide Europas, IUGS Series A (6), Turonien. Geobios 25 (1), 55–100. 527–542. Meister, C., Alzouma, K., Lang, J., Mathey, B., Pascal, A., 1994. Nouvelles données sur Holtz, T.R., Chapman, R.E., Lamanna, M.C., 2004. Mesozoic biogeography of les ammonites du Niger oriental (Ténéré, Afrique occidentale) dans le cadre de Dinosauria. In: Weishampel, D.B., Dodson, P., Osmólska, H. (Eds.), The la transgression du Cénomanien–Turonien. Geobios 27, 189–219. Dinosauria, second ed. University of California Press, Berkeley, pp. 627–642. Michard, A., Frizon de Lamotte, D., Saddiqi, O., Chalouan, A., 2008. An outline of the Janensch, W., 1925. Die Coelurosaurier und Theropoden der Tendaguru-Schichten geology of Morocco. In: Michard, A., Saddiqi, O., Chalouan, A., Frizon de Lamotte, Deutsch-Ostafrikas. Palaeontographica 1 (Suppl. 7), 1–99. D. (Eds.), Continental Evolution: The Geology of Morocco. Springer Verlag, Kellner, A.W.A., 2009. Brief review of the pterosaur record in Africa. In: Jalil, N.-E. Berlin, Heidelberg, pp. 1–32. (Ed.), 1st International Congress on North African Vertebrate Palaeontology, Milner, A.C., 2003. Fish-eating theropods, a short review of the systematics, biology Program & Abstracts, Marrakech, 25–27 May 2009, p. 34. and palaeobiogeography of spinosaurs. In: Huerta Hurtado, P., Torcida Kennedy, W.J., Gale, A.S., Ward, D.J., Underwood, C.J., 2008. Lower Turonian Fernandez-Baldor, F. (Eds.), Actas de las II Jornadas Internacionales sobre ammonites from Goulmima, southern Morocco. Bulletin de l’Institut Royal Paleontologıa de Dinosaurios y su Entorno (2001). Salas de Los Infantes, des Sciences de Belgique, Sciences de la Terre 78, 149–177. Colectivo Arqueologico–Paleontologico de Salas, pp. 129–138. Kilian, C., 1931. Des principaux complexes continentaux du Sahara. Comptes Moody, R.T.J., Sutcliffe, P.J.C., 1991. The Cretaceous deposits of the Iullemme- Rendus Sommaires de la Société Géologique de France 9, 109–111. den Basin of Niger, central West Africa. Cretaceous Research 12, 137– Kuss, J., 1994. Cretaceous (Albian–Turonian) Calcareous algae from Egypt and 157. Jordan – systematics, stratigraphy and paleogeography. Abhandlungen der Murray, A.M., Wilson, M.V.H., 2009. A new Late Cretaceous Macrosemiid fish Geologischen Bundes Anstalt 50, 295–317. (Neopterygii, Halecostomi) from Morocco, with temporal and geographical Lapparent, A.F.de, 1960. Les Dinosauriens du ‘Continental Intercalaire’ du Sahara range extensions for the family. Palaeontology 52 (2), 429–440. central. Mémoire de la Société Géologique de France (N.S.) 88A, 1–57. Murray, A., Wilson, M., Chatterton, B., 2007. A Late Cretaceous actinopterygian Larsson, H.C.E., Sidor, C.A., 1999. Unusual crocodyliform teeth from the Late fauna from Morocco. Journal of Vertebrate Paleontology 27 (3), 122A. Cretaceous (Cenomanian) of Southern Morocco. Journal of Vertebrate Nessov, L.A., Zhegallo, V.I., Averianov, A.O., 1998. A new locality of Late Cretaceous Paleontology 19 (2), 398–401. snakes, mammals and other vertebrates in Africa (western Libya). Annales de Larsson, H.C.E., Sues, H.-D., 2007. Cranial osteology and phylogenetic relationships Paléontologie, Paris 84, 265–274. of Hamadasuchus rebouli (Crocodyliformes, Mesoeucrocodylia) from the Novas, F.E., de Valais, S., Vickers-Rich, P., Rich, T., 2005. A large Cretaceous theropod Cretaceous of Morocco. Zoological Journal of the Linnean Society 149 (4), from Patagonia, Argentina, and the evolution of carcharodontosaurids. 533–567. Naturwissenschaften 92, 226–230. Lavocat, R., 1948. Découverte de Crétacé à vertébrés dans le soubassement de O’Leary, M.A., Roberts, E.M., Head, J.J., Sissoko, F., Bouaré, M.M., 2004. Titanosaurian l’Hammada du Guir (Sud marocain). Comptes Rendus de l’Académie des (Dinosauria, Sauropoda) remains from the ‘‘Continental Intercalaire” of Mali. Sciences 226, 1291–1292. Journal of Vertebrate Paleontology 24, 923–930. Lavocat, R., 1949. Les gisements de vertébrés crétacés du Sud marocain. Comptes Pittet, F., Cavin, L., Poyato-Ariza, F.J., 2009. A New Ostariophysan Fish from the early Rendus Sommaires de la Société Géologique de France 19 (5), 125–126. Late Cretaceous (Cenomanian) of SE Morocco, with a Discussion of its Lavocat, R., 1951. Découverte de restes d’un grand Dinosaurien sauropode dans le Phylogenetic Relationships. In: Grande, T. (Ed.), A Comprehensive Review of Crétacé du Sud marocain. Comptes Rendus de l’Académie des Sciences 232, Gonorynchiformes and of Ostariophysan Relationships. Science Publishers, Inc., 169–170. pp. 332–355. Lavocat, R., 1954a. Sur les dinosauriens du Continental Intercalaire des Kem-Kem de Pons, D., Berthou, P.-Y., Campos, D.A., 1990. Quelques observations sur la la Daoura. Comptes Rendus, 19th International Geological Congress 1, 65–68. palynologie de l’Aptien Supérieur et de l’Albien du Bassin d’Araripe (NE du Lavocat, R., 1954b. Reconnaissance géologique dans les Hammadas des confins Brésil). In: Atas 1 Simp. Bacia do Araripe e bacias interiores do Nordeste, Crato, algéro-marocains du Sud. Notes et Mémoires du Service Géologique du Maroc pp. 241–252. 116. 147 p. Premoli Silva, I., Sliter, W.V., 1995. Cretaceous planktonic foraminifera Lavocat, R., 1955a. Titres et travaux scientifiques. Imprimerie Centrale, Romorantin. biostratigraphy and evolutionary trends from the Bottaccione section, Gubbio, 95 p. Italy. Palaeontographia Italica 82. 89 p. Lavocat, R., 1955b. Découverte d’un Crocodilien du genre Thoracosaurus dans le Premoli Silva, I., Verga, D., 2004. Practical Manual of Cretaceous Planktonic Crétacé Supérieur d’Afrique. Bulletin du Muséum National d’Histoire Naturelle, Foraminifera. In: Verga, D., Rettori, R. (Eds.), Internationl School on Planktonic Paris 2 (27), 338–340. Foraminifera, 3° Course, Cretaceous. Universities of Perugia and Milan, Mader, B.J., Kellner, A.W.A., 1999. A new Anhanguerid pterosaur from the Tipografia Pontefelcino, Perugia, Italy. Cretaceous of Morocco. Boletim do Museu Nacional (Nova Serie), Geologia 45, Rage, J.-C., Cappetta, H., 2002. Vertebrates from the Cenomanian, and the 1–11. geological age of the Draa Ubari fauna (Libya). Annales de Paléontologie 88, Mahler, L., 2005. Record of Abelisauridae (Dinosauria, Theropoda) from the 79–84. Cenomanian of Morocco. Journal of Vertebrate Paleontology 25, 236–239. Rage, J.C., Dutheil, D.B., 2008. Amphibians and squamates from the Cretaceous Maisey, J.G., 2000. Continental break up and the distribution of fishes of (Cenomanian) of Morocco. A preliminary study, with description of a new genus Western Gondwana during the Early Cretaceous. Cretaceous Research 21, of pipid frog. Palaeontographica, Abteilung A 285, 1–22. 281–314. Rauhut, O.W.M., 1999. A dinosaur fauna from the Upper Cretaceous (Cenomanian) Maisey, J.G., Blum, S., 1991. Araripichthys. In: Maisey, J.G. (Ed.), Santana Fossils: An of northern Sudan. Palaeontologia Africana 35, 61–84. Illustrated Atlas. TFH Publications, Neptune, NJ, pp. 208–217. Rauhut, O.W.M., López-Arbarello, A., 2009. Considerations on the age of the Maisey, J.G., Moody, J.M., 2001. A review of the problematic teleost fish Tiouaren Formation (Iullemmeden Basin, Niger, Africa), implications for Araripichthys, with a description of a new species from the Lower Cretaceous Gondwanan Mesozoic terrestrial vertebrate faunas. Palaeogeography, of Venezuela. American Museum Novitates 3324, 1–27. Palaeoclimatology, Palaeoecology 271, 259–267. 412 L. Cavin et al. / Journal of African Earth Sciences 57 (2010) 391–412

Riff, D., Mader, B., Kellner, A.W.A., Russell, D., 2004. An avian vertebra from the Taverne, L., 2000. Nouvelles données ostéologiques et phylogénétiques sur continental Cretaceous of Morocco, Africa. Arquivos do Museu Nacional Rio de Palaeonotopterus greenwoodi, notoptéridé (Teleostei, Osteoglossomorpha) du Janeiro 62 (2), 217–223. Cénomanien inférieur continental (Crétacé) du Maroc. Stuttgarter Beiträge zur Robaszynski, F., Caron, M. (Eds.), and European Working Group on Planktonic Naturkunde, Serie B (Geologie und Paläontologie) 293, 1–24. Foraminifera, 1979. Atlas of Mid Cretaceous Planktonic Foraminifera (Boreal Sea Taverne, L., 2003. Redescription critique des genres Thryptodus, Pseudothryptodus et and Tethys). Parts 1–2, Cahiers de Micropaléontologie, 366 p. Paranogmius, poissons marins (Telostei, Tselfatiiformes) du Crétacé supérieur Russell, D.A., 1996. Isolated Dinosaur bones from the Middle Cretaceous of the des Etats-Unis, d’Egypte et de Libye. Belgian Journal of Zoology 133 (2), 163– Tafilat, Morocco. Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e 173. Série, Section C, 18 (2–3), 349–402. Taverne, L., 2004. On a complete hyomandibular of the Cretaceous Moroccan Russell, D.A., Paesler, M.A., 2003. Environments of Mid-Cretaceous Saharan notopterid Palaeonotopterus greenwoodi (Teleostei, Osteoglossomorpha). dinosaurs. Cretaceous Research 24, 569–588. Stuttgarter Beiträge zur Naturkunde Serie B (Geologie und Paläontologie) 348, Salgado, L., Garrido, A., Cocca, S.E., Cocca, J.R., 2004. Lower Cretaceous 1–7. rebbachisaurid sauropods from Cerro Aguada Del León, Neuquén Province, Taverne, L., Maisey, J.G., 1999. A notopterid skull (Teleostei, Osteoglossomorpha) northwestern Patagonia, Argentina. Journal of Vertebrate Paleontology 24 (4), from the continental early Cretaceous of Southern Morocco. American Museum 903–912. Novitates 3260, 1–12. Sereno, P.C., Larsson, H.C.E., 2009. Cretaceous Crocodyliforms from the Sahara. Tong, H., 2008. Les tortues du Cénomanien continental des Kem Kem et leurs ZooKeys 28, 1–143. implications paléobiogéographiques. Deuxième Rencontre sur la Valorisation et Sereno, P.C., Wilson, J.A., Larsson, H.C.E., Sues, H.-D., 1994. Early Cretaceous la Préservation du Patrimoine Paléontologique, Meknès, 9–13 May. Abstracts dinosaurs from the Sahara. Science 266, 267–271. volume, pp. 69–70. Sereno, P.C., Dutheil, D.B., Iarochène, M., Larsson, H.C.E., Lyon, G.H., Magwene, Tong, H., Buffetaut, E., 1996. A new genus and species of pleurodiran turtle from the P.M., Sidor, C.A., Varricchio, D.J., Wilson, J.A., 1996. Predatory Dinosaurs from Cretaceous of southern Morocco. Neues Jahrbuch für Geologie und the Sahara and Late Cretaceous Faunal Differentiation. Science 272, 986– Paläontologie, Abhandlungen 199 (1), 133–150. 991. Tykoski, R.S., Rowe, T., 2004. Ceratosauria. In: Weishampel, D.B., Dodson, P., Sereno, P.C., Beck, A.L., Dutheil, D.B., Gado, B., Larsson, H.C.E., Lyon, G.H., Marcot, J.D., Osmólska, H. (Eds.), The Dinosauria, second ed. University of California Press, Rauhut, O.W.M., Sadleir, R.W., Sidor, C.A., Varrichio, D.D., Wilson, G.P., Wilson, Berkeley, pp. 151–168. J.A., 1998. A long-snouted predatory dinosaur from Africa and the evolution of Underwood, C., Cleason, K.M., Ward, D.J., 2009. Batoids from the Turonian of SE spinosaurids. Science 282, 1298–1302. Morocco. In: Jalil, N.-E. (Ed.), 1st International Congress on North African Sereno, J.A., Wilson, J.A., Conrad, J.L., 2004. New dinosaurs link southern landmasses Vertebrate Palaeontology, Program & Abstracts, Marrakech, 25–27 May 2009, p. in the Mid-Cretaceous. Proceedings of the Royal Society of London B 271 (1546), 43. 1325–1330. Van Der Zwaan, G.J., Duijnstee, I.A.P., Den, D.M., Ernst, S.R., Jannink, N.T., Silva Santos, R. da, 1985. Araripichthys castilhoi novo gênero e especie de teleostei da Kouwenhoven, T.J., 1999. Benthic foraminifers; proxies or problems? A review Formação Santana, Chapada do Araripe, Brasil. In: Campos, D.A., Ferreira, C.S. of paleocological concepts. Earth-Science Review 46 (1–4), 213–236. Brito, I.M., Viana, C.F. (Eds.), Coletânea de Trabalhos Paleontológicos, Série Vullo, R., Néraudeau, D., 2008a. When the ‘‘primitive” shark Tribodus Geologia 27 (2). Rio de Janeiro: Ministério das Minas e Energia—D.N.P.M., pp. (Hybodontiformes) meets the ‘‘modern” ray Pseudohypolophus (Rajiformes), 133–140. the unique co-occurrence of these two durophagous Cretaceous selachians in Smith, J.B., Lamanna, M.C., Lacovara, K.J., Dodson, P., Smith, J.R., Poole, J.C., Charentes (SW France). Acta Geologica Polonica 58 (2), 249–255. Giegengack, R., Attia, Y., 2001. A giant sauropod dinosaur from an Upper Vullo, R., Néraudeau, D., 2008b. Cenomanian vertebrate assemblages from Cretaceous mangrove deposit in Egypt. Science 292, 1705–1706. southwestern France, a new insight into the European mid-Cretaceous Stromer, E., 1915. Ergebnisse der Forschungsreisen Prof. E. Stromer in den Wüsten continental fauna. Cretaceous Research 29 (5–6), 930–935. Agyptens. II. Wirbeltier-Reste der Baharîje-Stufe (unterstes Cenoman). 3. Das Vullo, R., Néraudeau, D., Allain, R., Cappetta, H., 2005. Un nouveau gisement à Original des Theropoden Spinosaurus aegyptiacus nov. gen., nov. spec. microrestes de vertébrés continentaux et littoraux dans le Cénomanien Abhandlungen der Königlich Bayerischen Akademie der Wissenschaften, inférieur de Fouras (Charente-Maritime, Sud-Ouest de la France). Comptes Mathematisch-Physikalische Klasse 28, 1–32. Rendus Palevol 4, 95–107. Stromer, E., 1925. Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wüsten Weiler, W., 1935. Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wüsten Ägyptens. II. Wirbeltier-Reste der Baharije-Stufe (unterstes Cenoman). 7. Ägyptens. II. Wirbeltierreste der Baharîje-Stufe (unterstes Cenoman). Neue Stomatosuchus inermis Stromer, ein schwach bezahnter Krokodilier und 8. Ein Untersuchungen an den Fischresten. Abhandlungen der Bayerischen Akademie Skelettrest des Pristiden Onchopristis numidus Huag sp. Abhandlungen der der Wissenschaften, Mathematische-naturwissenschaftliche Abteilung, Neue Königlich Bayerischen Akademie der Wissenschaften, Mathematisch- Folge 32, 1–57. Physikalische Klasse 30, 1–22. Wellnhofer, P., Buffetaut, E., 1999. Pterosaur remains from the Cretaceous of Stromer, E., 1932. Wirbeltiere-Reste der Baharije-Stufe (unterstes Cenoman). 11. Morocco. Paläontologische Zeitschrift 73 (1/2), 133–142. Sauropoda. Abhandlungen der Bayerischen Akademie der Wissenschaften Wenz, S., 1980. A propos du genre Mawsonia, Coelacanthe géant du Crétacé Mathematisch naturwissenschafiliche Abteilung, Neue Folge 10, 3–21. inférieur d’Afrique et du Brésil. Mémoire de la Société Géologique de France, Tabaste, N., 1963. Étude de restes de poissons du Crétacé saharien. Mémoire, IFAN N.S. 139, 187–190. 68, mélanges Ichthyologiques, 437–485. Wenz, S., 1981. Un coelacanthe géant, Mawsonia lavocati Tabaste, de l’Albien–base Taquet, P., 1970. Sur le gisement de dinosauriens et de crocodiliens de Gadoufaoua du Cénomanien du Sud marocain. Annales de Paléontologie (Vertébrés) 67 (1), (République du Niger). Comptes Rendus de l’Académie des Sciences, Paris, (D) 1–20. 271, 38–40. Werner, C., 1994. Die kontinentale Wirbeltierfauna aus der unteren Oberkreide des Taquet, P., 1976. Géologie et Paléontologie du gisement de Gadoufaoua (Aptien du Sudan (Wadi Milk Formation). Berliner Geowissenschaftliche Abhandlungen (E) Niger). Cahiers de Paléontologie, CNRS Paris, 91 p. 13, 221–249. Taquet, P., 1980. Succession et repartition des gisements de vertébrés du Crétacé du Wilson, M.V.H., Murray, A.M., Chatterton, B.D.E., Gibbs, S., 2009. Late Cretaceous Sahara. Mémoire de la Société Géologique de France, N.S. 139, 185–186. actinopterygian fishes from Morocco. In: Jalil, N.-E. (Ed.), 1st International Taquet, P., 1984. Une curieuse spécialisation du crâne de certains Dinosaures Congress on North African Vertebrate Palaeontology, Program & Abstracts, carnivores du Crétacé, le museau long et étroit des Spinosauridés. Comptes Marrakech, 25–27 May 2009, p. 44. Rendus de l’Académie des Sciences, Paris Série II 299, 217–222. Zouhri, S., Kchikach, A., Saddiqi, O., El Haïmer, F.Z., Baidder, L., Michard, A., 2008. The Taverne, L., 1997. Les clupéomorphes (Pisces, Teleostei) du Cénomanien (Crétacé) Cretaceous–Tertiary Plateaus. In: Michard, A., Saddiqi, O., Chalouan, A., Frizon de Kipala (Kwango), Zaire), ostéologie et phylogénie. Belgian Journal of Zoology de Lamotte, D. (Eds.), Continental Evolution: The Geology of Morocco. Springer 127 (1), 75–97. Verlag, Berlin, Heidelberg, pp. 331–358.