Upper Cenomanian–Turonian (Upper Cretaceous) ammonoids from the western Wadi Araba, Eastern...

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Cretaceous Research 31 (2010) 473e499

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Cretaceous Research

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Upper CenomanianeTuronian (Upper Cretaceous) ammonoids from the westernWadi Araba, Eastern Desert, Egypt

Emad Nagma, Markus Wilmsen b,*, Mohamed F. Aly c, Abdel-Galil Hewaidy a

aGeology Department, Faculty of Science, Al-Azhar University, Egyptb Senckenberg Naturhistorische Sammlungen Dresden, Museum für Mineralogie und Geologie, Sektion Paläozoologie, Königsbrücker Landstr. 159, D-01109 Dresden, GermanycGeology Department, Faculty of Science, Cairo University, Giza, Egypt

a r t i c l e i n f o

Article history:Received 16 December 2009Accepted in revised form 19 May 2010Available online 26 May 2010

Keywords:Lower Upper CretaceousAmmonoidsSystematic palaeontologyPalaeobiogeographyEgypt

* Corresponding author.E-mail addresses: [email protected] (E.

senckenberg.de (M. Wilmsen).

0195-6671/$ e see front matter � 2010 Elsevier Ltd.doi:10.1016/j.cretres.2010.05.008

a b s t r a c t

The CenomanianeTuronian (CeT) successions (Galala and Maghra el Hadida formations) exposed on thefootwalls of the slopes of the Galala plateaus in the Wadi Araba area in the northern part of the EasternDesert, Egypt yielded a fairly rich and moderately to well preserved ammonoid fauna. In total, 24 taxahave been identified, 17 of which are systematically described herein. Four genera (Thomelites, Euom-phaloceras, Wrightoceras, Eubostrychoceras) and four species (Euomphaloceras septemseriatum, Vascoceraspioti, Fagesia cf. peroni, Wrightoceras munieri) are recorded for the first time from Egypt. In addition,the taxonomy of ammonoids previously reported from these successions has been re-evaluated. Theammonoid fauna came from Upper Cenomanian as well as Lower and Upper Turonian (pars) strata; theMiddle Turonian strata are barren of ammonoids. The presence of lower Upper Cenomanian ammonoidsin the Galala Formation and mid-/upper Upper Cenomanian ammonoids in the lower part of the Maghrael Hadida Formation shows that the formational boundary between both units does not coincide withthe CeT boundary as often reported in the literature. The palaeobiogeographic affinities of the taxarecorded classify the ammonoid assemblage as part of the (southern) Tethyan Vascoceratid Provinceduring the early Late Cretaceous.

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1. Introduction

Upper Cretaceous strata in northeast Egypt arewell exposed andfossiliferous, yielding rich macrofaunas, especially ammonoids,from many localities in the Eastern Desert and Sinai. In the lasttwenty years, numerous studies have been published on thetaxonomy of CenomanianeTuronian ammonoids of these areas(Luger and Gröschke, 1989; Kassab, 1994, 1996; Aly and Abdel-Gawad, 2001; El-Hedeny, 2002; Hewaidy et al., 2003; Abdel-Gawad et al., 2004, 2007; El Qot, 2006). However, only a few andoften fairly poorly to moderately preserved ammonoids have beendescribed in most of these publications. Therefore, this paper aimsto enhance the knowledge on the distribution and taxonomy of theCenomanianeTuronian ammonoid faunas in the Eastern Desert bymeans of a systematic description of a large collection of moder-ately to well preserved ammonoids collected bed-by-bed from theCenomanianeTuronian Galala and Maghra el Hadida formations. Intotal, 81 ammonoids have been collected during three field

Nagm), markus.wilmsen@

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campaigns (2006e2008) from four sections in the western partof the Wadi Araba area, Eastern Desert, Egypt. In addition tothesystematic description, the ammonoid fauna is placed in apalaeobiogeographic framework.

2. Geological setting and stratigraphic framework

Wadi Araba is located in the northern part of the EgyptianEastern Desert (Fig. 1). This area is bounded to the north and southby the Galala plateaus and to the east by the Gulf of Suez. WadiAraba is about 30 km wide and extends westward to the centralEocene limestone plateau of the Eastern Desert. It has a NEeSWtrend, following the direction of the regional Syrian Arc anticlinestructure, one of the best-known structural features in northernEgypt (e.g., Said, 1990). The Syrian Arc can be traced from Syria tothe central Western Desert of Egypt, via Sinai and the northern partof the Eastern Desert. The Galala plateaus represent a major branchof the Syrian Arc in the Eastern Desert, characterized by LateCretaceous uplift in the north and subsidence farther to the south.Folding and/or uplift of the Syrian Arc began in post-Cenomaniantimes (Aal and Lelek, 1994) and reached its acme during the LateCretaceous (Kuss et al., 2000).

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Fig. 1. Geographic position of the study area in Egypt (lower right) with indication of localities of the four measured sections.

E. Nagm et al. / Cretaceous Research 31 (2010) 473e499474

Palaeogeographically, Egypt was situated during the Cretaceousat the southern margin of the Neotethys Ocean (Fig. 2). The Cen-omanianeTuronian interval was probably the most widespreadCretaceous transgression in northeastern Egypt, with maximumflooding during Early Turonian times (maximum flooding surfaceK140 of Sharland et al., 2001). Consequently, lower Upper Creta-ceous marine strata are very widespread.

The CenomanianeTuronian successions of the two Galalaplateaus consist mainly of inner- to mid-shelf mixed siliciclasticand carbonate sequences represented by the Galala and Maghra elHadida formations. The four measured sections (Fig. 3) start withunfossiliferous, brownish sandstones of the non-marine MalhaFormation (Lower Cretaceous, eventually ranging up into theCenomanian). Bioturbation in the upper part of the Malha Forma-tion indicates initial marine influence and a gradual interfingeringof marine and non-marine deposits. Lagoonal, but fully marineenvironmental conditions are documented by the muddy, fossilif-erous sediments of the Cenomanian Galala Formation, consisting ofsiltstone, sandy marly silt, and limestones in the lower part, fol-lowed by intercalations of marl and limestones in the upper part. Itcontains abundant oysters as well as rudistid and chondrodontidbivalves, large gastropods, and a diverse assemblage of echinoids(Nagm, 2009). The Galala Formation ranges in thickness from 55 min the Saint Anthony section to 95 m in the East Wadi Ghonimasection. Based on the occurrence of Neolobites vibrayeanus (seebelow) and the characteristic associated macrofauna, an early LateCenomanian age is inferred for the Galala Formation in the studyarea (Nagm, 2009).

The lower boundary of the Maghra el Hadida Formation ischaracterized by a major unconformity which separates it fromthe underlying Galala Formation. The formation starts witha brown, fine- to medium-grained calcareous sandstone unitcharacterized by strong lateral thickness shifts from a few metresto >20 m. The overlying succession of the Maghra el HadidaFormation is characterized by an increase in carbonate content,

represented by yellow, soft marls intercalated with fine-grainedwacke- to packstones containing a rich and diverse ammonoidassemblage of the middle and upper Upper Cenomanian as well asthe Lower Turonian. The middle part of the Maghra el HadidaFormation is characterized by marlstone, becoming more siltyupward, with occasional hummocky cross-bedded sandstoneintercalations, suggesting deposition in a storm-influenced shelfsetting. The topmost part of the Maghra el Hadida Formationconsists of brownish, medium-grained sandstones topped byfossiliferous marly limestones yielding Upper Turonian ammo-noids. The measured thickness of the Maghra el Hadida Formationranges from 59 m at Saint Anthony to 118 m at East Wadi Gho-nima. The presence of lower Upper Cenomanian ammonoids inthe Galala Formation and mid- to upper Upper Cenomanianammonoids in the lower part of the Magra el Hadida Formation(see below) shows that the boundary between both formationsdoes not coincide with the CeT boundary as often reported in theliterature (e.g., Hewaidy et al., 2003). Three ammonite zones havebeen recognized in the Upper Cenomanian, from base to top, theN. vibrayeanus, Metoicoceras geslinianum, and Vascoceras cauvinitotal range zones (Nagm, 2009; Nagm et al., 2010). The threezones are equivalent to the Upper Cenomanian standard zones ofCalycoceras naviculare, M. geslinianum and Neocardioceras juddii(cf. Gradstein et al., 2004). The Lower Turonian likewise containsthree zones, characterized by the taxon ranges of Vascocerasproprium, Choffaticeras spp., and Wrightoceras munieri. In thestandard zonation, the first two zones correspond to the lowerLower Turonian Watinoceras coloradoense Zone whereas theW. munieri Zone is equivalent to the upper Lower TuronianMammites nodosoides Zone (see Nagm et al., 2010 for furtherdetails). The Middle Turonian is barren of ammonoids, while thelower Upper Turonian is represented by the Coilopoceras requie-nianum Zone, corresponding to the (lower part of the) Sub-prionocyclus neptuni (standard) Zone. The ammonoid biozonationof the lower Upper Cretaceous of the Wadi Araba area is the topic

Fig. 2. Late Cenomanian palaeogeographic setting (modified from Philip and Floquet, 2000). The rectangle indicates the position of the study area.

E. Nagm et al. / Cretaceous Research 31 (2010) 473e499 475

of another paper (Nagm et al., 2010) and the reader is referredthereto for further details.

The stratigraphic and facies analyses of the Galala andMaghra elHadida formations (Nagm, 2009) document an overall trans-gressive development during the Late Cenomanian and EarlyTuronian with peak transgression in the early Early Turoniancoinciding with maximum flooding K140 of Sharland et al. (2001).Crustacean microcoprolites and calcareous algae of the successionare the topics of the papers by Senowbari-Daryan et al. (2009) andBucur et al. (2010).

3. Systematic paleontology

The ammonoids studied are arranged systematically accordingto the scheme of Wright et al. (1996). In order to keep thischapter (and the references) as short as possible, the synonymiesonly contain the most recent ones as well as those being ofregional interest or those being important for the discussion.Likewise, authors of higher rank systematic categories are notincluded in the references. The terminology used for thedescription of the taxa follows the glossary in the Treatise onInvertebrate Paleontology, Part L, Mollusca 4 (Wright et al., 1996).All linear dimensions, taken with a Vernier Caliper, are given inmillimetres. The abbreviations that are used in the present workare: maximum diameter (D), whorl breadth (Wb), whorl height(Wh), breadth of umbilicus (U). Figures in parentheses refer todimensions as a percentage of diameter. The material is kept inthe Museum für Mineralogie und Geologie (MMG) of the Senck-enberg Naturhistorische Sammlungen Dresden (SNSD), Germany(repository AfK).

Order: Ammonoidea Zittel, 1884Suborder: Ammontina Hyatt, 1889Superfamily: Hoplitoidea Douvillé, 1890? Family: Engonoceratidae Hyatt, 1900Genus Neolobites Fischer, 1882

Neolobites vibrayeanus (d’Orbigny, 1841)Figs. 4, 5AeF

1841 Ammonites vibrayeanus d’Orbigny, p. 322, pl. 96, figs.1e3.

1981 Neolobites vibrayeanus (d’Orbigny); Kennedy and Juignet,p. 23, figs. 3e4, 6a; text-fig. 5 (with synonymy).

1989 Neolobites vibrayeanus (d’Orbigny); Luger and Gröschke, p.366, pl. 39, fig. 3; text-fig. 5.

1996 Neolobites vibrayeanus (d’Orbigny); Kassab, pl. 1,figs. 1e3.

2001 Neolobites vibrayeanus (d’Orbigny); Aly and Abdel-Gawad,p. 29, pl. 1, figs. 2e4; text-fig. 3a,b.

2002 Neolobites vibrayeanus (d’Orbigny); El-Hedeny, p. 401,fig. 3aeb, fig. 7d, fig. 8e.

2003 Neolobites vibrayeanus (d’Orbigny); Hewaidy et al., p. 340,pl. 1, figs. 7e8.

2004 Neolobites vibrayeanus (d’Orbigny); Abdel-Gawad et al., pl.1, fig. 2.

2005 Neolobites vibrayeanus (d’Orbigny); Wiese and Schulze,p. 933, figs. 4e9 (with synonymy).

2006 Neolobites vibrayeanus (d’Orbigny); El Qot, p. 116, pl. 24,figs. 4e5.

2008 Neolobites vibrayeanus (d’Orbigny); Aly et al., p. 46, pl. 3,figs. 2e3; text-fig. 3 (2).

Fig. 3. Lithostratigraphic correlation of the measured sections arranged on the CenomanianeTuronian datum line (C/T).

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Material. 10 moderately preserved, incomplete internal moulds(WG-10-1e9, 080216-8.5-3) from the Wadi Ghonima section (15 mbelow the top of Galala Formation), two moderately preserved,incomplete internal moulds (080217-11-3, 080213-3-3) from theEast Wadi Ghonima section (at 58.5 m and 84 m), and oneincomplete internal mould (WA-9-1) from theWadi Askhar section(at 60 m). All specimens were collected from the lower UpperCenomanian.

Measurements.

Specimen No. D Wb Wh Wb/Wh U

WG-10-1 69.6 (100) 23.3 (33.4) 46.4 (66.6) 0.50 6.4 (9.1)WG-10-2 106.6 (100) 26.0 (24.3) 59.4 (55.7) 0.43 8.1 (7.5)

Description. Very involute, small- to medium-sized. Whorlsstrongly compressed, elevated, with lanceolate shape and flatventer. Greatest breadth around mid-flank. Flanks slightly convexwith ventrolateral shoulders ornamented by delicate clavi,approximately 40 clavi/whorl in specimen WG-10-1. Umbilicussmall (about 9% of total diameter), shallow, with low androunded umbilical wall. Very weak ribs are noted on the innerparts of flanks. The suture line has broad saddles, narrow andlanceolate lobes, with flattened ends of all suture elements. Thelobes become shallower in the direction of the umbilicus. Inaddition, some specimens show overlapping suture lines(Fig. 4A).

Remarks. N. vibrayeanus can be recognized by its compressedwhorls, distinctive suture line, flat venter and the small umbilicus.The specimens studied herein closely correspond to the holotypeas figured by Kennedy and Juignet (1981) and the specimensdescribed by Luger and Gröschke (1989) from Egypt. Most of ourspecimens lack ornament, but some show delicate ventrolateralclavi and very weak ribbing on the inner flanks. In addition, theyshow considerable variation in the suture line due to taphonomicconditions such as compaction. Differences to other Neolobitesspecies have been discussed by Kennedy and Juignet (1981), Lugerand Gröschke (1989), and Wiese and Schulze (2005). The spec-imen described by El-Hedeny and Nafee (2001, p. 119, pl. 1, fig. 1)and also the specimen figured by Kora et al. (2001, pl. 2, fig. 9, 10)from Egypt have suture lines which differ from the suture ofN. vibrayeanus. Furthermore, the venter of the latter authors’

Fig. 4. Suture lines and whorl section of Neolobites vibrayeanus (d’Orbigny, 1841), Upper CenWG-10-6; D and E, specimen WG-10-2. All specimens from the Wadi Ghonima section (15

specimen seems to be sharp which is incompatible with thetabular venter of N. vibrayeanus. Therefore, their specimens arenot regarded here as N. vibrayeanus. Also, the suture line of thespecimen figured by Abdel-Gawad et al. (2004b, pl. 6, fig. 1a, 1b)as N. vibrayeanus is completely different from the suture line ofthe species; the specimen in question may be belong to theAcanthoceratidae.

Occurrence. According to Kennedy and Simmons (1991, p. 136)and Wiese and Schulze (2005), N. vibrayeanus (d’Orbigny) isa widespread and characteristic lower Upper Cenomanian species,occurring commonly in western Europe (France, Spain, andPortugal), northern Africa and the Middle East (Morocco, Algeria,Tunisia, Niger, Egypt, Israel, Lebanon, Jordan, Syria, Saudi Arabia,and Oman), as well as South America (Bolivia, Colombia, Peru andVenezuela). In Egypt, it has been recorded from the Upper Cen-omanian of the north Eastern Desert, from the Gulf of Suez region,and from Sinai. In the study area, N. vibrayeanus is recorded fromthe lower Upper Cenomanian N. vibrayeanus Zone, which is (asdefined by Nagm et al., 2010) equivalent to the standard ammoniteZone of C. naviculare (Gradstein et al. 2004).

Superfamily: Acanthoceratoidea Grossouvre, 1894Family: Acanthoceratidae Grossouvre, 1894Subfamily: Acanthoceratinae Grossouvre, 1894Genus Thomelites Wright and Kennedy, 1973

Thomelites sp.Figs. 5GeH, 7AeB

Material. One moderately preserved, incomplete compositemould (WA-7-1) from the Upper Cenomanian of the GalalaFormation in the Wadi Askhar section (at 50 m).

Description. Medium-sized, with compressed whorl section(Fig. 7AeB). Maximum width at the umbilical shoulders. Ventertabular, compressed, and relatively narrow. Umbilicus moderatelywide with very strong umbilical tubercles. Ribs arise singly or inpairs from umbilical tubercles and are intercalated. The ventrolat-eral shoulders have strong inner tubercles and outer clavi.

Occurrence. According to Wright and Kennedy (1981), the genusThomelites is recorded from the Upper Cenomanian of England,France, Israel, (?) southern India, Texas, South Dakota, and Brazil.This is thefirst recordof the genus Thomelites fromEgypt, it has beencollected from the lower Upper Cenomanian (N. vibrayeanus Zone).

omaninan Galala Formation. A, specimen WG-10-1; B, specimen WG-10-3; C, specimenm below the top of Galala Formation). Not to scale.


080217-26-1 118.21 (100) 42.74 (36.1) 54.15 (45.8) 0.78 43.56 (36.8)080217-26-2 57.88 (100) 19.35 (33.4) 23.51 (40.6) 0.82 22.96 (39.6)


Subfamily: Euomphaloceratinae Cooper, 1978Genus Euomphaloceras Spath, 1923

Euomphaloceras septemseriatum (Cragin, 1893)Figs. 5IeK, 7C

1893 Scaphites septem-seriatum Cragin, p. 240.1981 Euomphaloceras septemseriatum (Cragin); Wright and

Kennedy, p. 55, pl. 12, figs. 1e8; pl. 13, figs. 1e6; pl. 14, figs.5e9 (with full synonymy).

2003 Euomphaloceras septemseriatum (Cragin); Kennedy et al.,p. 10, pl. 2, figs. 9, 10, 13, 15, 18e23.

Material. Two poorly preserved incomplete composite moulds(080215-6-1, 6-2) from the Upper CenomanianM. geslinianum Zoneof the Maghra el Hadida Formation in the Wadi Ghonima section(24.7 m above the base of the formation).

Description. Medium-sized, with subtrapezoidal, depressedwhorl section (Fig. 7C). Maximum width at the position of theumbilical tubercles. Venter wide, with weak keel ornamented byweak siphonal tubercles equal in number to the ventrolateraltubercles. Flanks short, ornamented with weak ribs that are bentforward at the ventrolateral shoulder.

Remarks. Euomphaloceras septemseriatum has a depressed whorlsection and a wide venter with siphonal tubercles and two rows ofobliquely oriented ventrolateral tubercles on the two sides.Kennedy et al. (2003) recorded wide morphological and ontoge-netic variations. However, the present material is too poorlypreserved to comment on this, and most characters have beendocumented by previous authors (e.g., Wright and Kennedy, 1981).

Occurrence. E. septemseriatum has been recorded from the UpperCenomanianof England, France, Angola, theUnited States, andTunisia(e.g., Wright and Kennedy, 1981; Kennedy et al., 2003). This is thefirst record of E. septemseriatum from Egypt, where it has beenrecorded fromtheUpperCenomanianassociatedwithM. geslinianum.

Genus Kamerunoceras Reyment, 1954

Kamerunoceras turoniense (d’Orbigny, 1850)Figs. 6AeF, 7DeE

1850 Ammonites turoniensis d’Orbigny, p. 190.1969 Protexanites salmuriensis (Courtiller); Freund and Raab,

p. 69, pl. 5, figs. 4e6; text-fig. 14cej.1979a Kamerunoceras turoniense (d’Orbigny); Kennedy and

Wright, p. 1170, pl. 2, figs. 1e11; pl. 3, figs. 1e2; pl. 4, figs.1e3; text-fig. 2e3 (with full synonymy).

1994 Kamerunoceras turoniense (d’Orbigny); Chancellor et al.,p. 26, pl. 4, figs. 1e3; pl. 5, figs. 1e3; pl. 6, figs. 6e7; pl. 7,figs. 3e4; pl. 8, figs. 8e9; text-fig. 11a, g.

2001 Protexanites cf. salmuriensis (Courtiller); Galal et al., pl. 5,figs. 8e9.

2007 Kamerunoceras turoniense (d’Orbigny); Barroso-Barcenilla, p. 153, pl. 12, figs. dee; pl. 13, figs. aef (withadditional synonymy).

Material. Three moderately preserved, incomplete compositemoulds (080217-26-1e3) from the Lower Turonian Choffaticerasspp. Zone of the Maghra el Hadida Formation in the East WadiGhonima section (at 132 m).

Fig. 5. AeF, Neolobites vibrayeanus (d’Orbigny, 1841), Upper Cenomanian Galala Formation. Ax0.75; WG-10-2; E: lateral view, x1; F: ventral view, x1; WG-10-3. All specimens from WadiCenomanian Galala Formation. G: lateral view, x1; H: ventral view, x1; WA-7-1 from WadCenomanian Maghra el Hadida Formation. I: lateral view, x1; J: ventral view, x1; 080215-6-above the base of the formation).

Measurements.

Description. Evolute, with large umbilicus, reaching up to 40% oftotal diameter. Medium- to large-sized, compressed, with greatestbreadth at the outer ventrolateral tubercles. Flanks narrow, flat-tened, with broadly rounded ventrolateral shoulders. Whorlsection oval to subrectangular. Venter slightly convex with ridgemarking the site of the siphuncle. Ornamentation consists of weakribs, bearing strong and well developed inner and outer ventro-lateral tubercles as well as weak tubercles near the umbilicalshoulders. The smallest specimen (080217-26-2) has 7 ribs per halfwhorl. Suture line characterized by a narrow, deep external lobe,with a narrow, high external saddle, a broad asymmetrically bifidlateral saddle, a moderately wide, deep lateral lobe, and a lowumbilical saddle with small branches (Fig. 7DeE).

Remarks. The wide and evolute coiling, flank ribbing with rowsof tubercles on umbilical shoulder or on mid-flank, inner and outertubercles on ventrolateral shoulders, and the wide venter witha ridge marking the site of the siphuncle are the features thatcharacterize Kamerunoceras turoniense. Kennedy and Wright(1979a) revised the abundant material of K. turoniense from Tour-aine in France. They described this species in detail and discussedits relationship to other species of Kamerunoceras. Chancellor et al.(1994) mentioned that K. turoniense extended well into the MiddleTuronian, although they recorded it from the Lower Turonian (theirThomasites rollandi Zone). The specimens studied herein are nearlyidentical to Tunisian material described by Pervinquière (1907),Robaszynski et al. (1990), and Chancellor et al. (1994), and co-occur with T. rollandi in the upper part of the Choffaticeras spp. Zoneof the Lower Turonian.

Occurrence. According to Kennedy and Wright (1979a), Wrightand Kennedy (1981) and Chancellor et al. (1994), K. turoniensehas been recorded from the Lower Turonian of Tunisia, Israel,France, England, Spain and Venezuela. In Egypt, Galal et al. (2001)recorded the species from the Eastern Desert while Abed et al.(1996) and Bauer et al. (2001) mentioned (without figuring)K. turoniense from the Lower Turonian of Sinai.

Genus Pseudaspidoceras Hyatt, 1903

Pseudaspidoceras sp.Fig. 6G

Material. One poorly preserved incomplete composite mould(080215-6-3) from the Upper Cenomanian M. geslinianum Zone ofthe Maghra el Hadida Formation in the Wadi Ghonima section(24.7 m above the base of the formation).

Description.Medium-sized, evolute, with large umbilicus. Whorlsection compressed with maximum breadth at the outer ventro-lateral tubercles. Flanks narrow, with flat sides. Ornamentationconsists of weak ribs, bearing weak inner and outer ventrolateraltubercles with umbilical bullae.

Remarks. The present specimen is similar to Pseudaspidoceraspaganum Reyment (1954) as described by Zaborski (1995, p. 63,

: lateral view, x1; B: ventral view, x1; WG-10-1; C: ventral view, x0.75; D: lateral view,Ghonima section (15 m below the top of Galala Formation). GeH, Thomelites sp., Upperi Askhar section (at 50 m). IeK, Euomphaloceras septemseriatum (Cragin, 1893), Upper1; K: ventral view, x1; 080215-6-2. All specimens from Wadi Ghonima section (24.7 m

Fig. 7. Whorl section and suture line of AeB, Thomelites sp., Upper Cenomanian Galala Formation; C, Euomphaloceras septemseriatum (Cragin, 1893), Upper Cenomanian Maghra elHadida Formation; DeE, Kamerunoceras turoniense (d’Orbigny, 1850), Lower Turonian Maghra el Hadida Formation; F, Metoicoceras geslinianum (d’Orbigny, 1850), Upper Cen-omanian Maghra el Hadida Formation. AeB, specimen WA-7-1 from the Wadi Askhar section (at 50 m); C, specimen 080215-6-1 from the Wadi Ghonima section (24.7 m above thebase of the formation); DeE, specimen 080217-26-1 from the East Wadi Ghonima section (at 132 m); F, specimen 080215-3-1 from the Wadi Ghonima section (24.7 m above thebase of the formation). Not to scale.


080215-3-1 50.2 (100) 11.2 (22.3) 25.8 (51.3) 0.43 9.1(18.1)


figs. 15e16, 22e23) from Nigeria and by Meister and Abdallah(2005, p.127, pl. 6, fig. 2) from Tunisia. However, the presentspecimen is unfortunately too poorly preserved to permit a specificidentification.

Occurrence. According to Zaborski (1995), the genus Pseudaspi-doceras has a stratigraphical range from the Upper Cenomanian tothe Lower Turonian. It occurs in Texas, New Mexico, Arizona andColorado (USA), Mexico, Brazil, Germany, southern England,Portugal, Tunisia, Egypt, Algeria, Angola, Niger, Nigeria,Madagascar, southern India and (?) Japan. The present specimen isrecorded from the Upper Cenomanian in association withM. geslinianum.

Subfamily: Mammitinae Hyatt, 1900Genus Metoicoceras Hyatt, 1903

Metoicoceras geslinianum (d’Orbigny, 1850)Figs. 7F, 8AeC

1841 Ammonites catillus Sowerby; d’Orbigny, p. 325, pl. 97,figs. 1e2.

1850 Ammonites Geslinianus d’Orbigny, p. 146.1981 Metoicoceras geslinianum (d’Orbigny, 1850); Wright and

Kennedy, p. 62, pl. 17, fig. 2; pl. 18, figs. 1, 2; pl. 19, figs. 1, 2;pl. 20, figs. 1e3; pl. 21, figs. 1, 2; text-figs. 19cee, 20,21aed (see for synonymy).

1981 Metoicoceras geslinianum (d’Orbigny); Kennedy andJuignet, p. 39, text-figs. 7dee, 8aec, 9a, e, 10a.

1996 Metoicoceras geslinianum (d’Orbigny); Kassab, pl. 1, figs.4e6.

Fig. 6. AeF, Kamerunoceras turoniense (d’Orbigny, 1850), Lower Turonian Maghra el HadidaD: ventral view, x1; 080217-26-2; E: lateral view, x0.75; F: ventral view, x0.75; 080217-26-Upper Cenomanian Maghra el Hadida Formation. lateral view, x1; 080215-6-3 from Wadi G

?2001 Metoicoceras geslinianum (d’Orbigny); Aly and Abdel-Gawad, p. 35, pl. 4, fig. 2.

2005 Metoicoceras geslinianum (d’Orbigny); Meister andAbdallah, p. 128, pl. 8, fig. 1 (with additional synonymy).

2009 Metoicoceras geslinianum (d’Orbigny); Lehmann andHerbig, p. 69, pl. 1, figs. teu.

Material. Two relatively poorly preserved composite moulds(080215-3-1, 3-2) from the Upper CenomanianM. geslinianum Zoneof the Maghra el Hadida Formation in the Wadi Ghonima section(24.7 m above the base of the formation).

Measurements.

Description. Involute, small- to medium-sized. Whorls stronglycompressed (Fig. 7F). Greatest breadth around mid-flank. Venternarrow, with ventrolateral shoulders. Umbilicus medium-sized,shallow, with low umbilical wall and rounded umbilical shoulder.Ornamentation consists of weak umbilical bullae giving rise tomoderately strong and straight ribs with additional short inter-calatories. All ribs bear slightly clavate ventrolateral tubercles.Suture line poorly preserved.

Remarks. Even if their preservation is poor, the studied speci-mens are fairly similar to the lectotype of M. geslinianum asdescribed by Kennedy and Juignet (1981), differing only in having

Formation. A: ventral view, x1; B: lateral view, x1; 080217-26-1; C: lateral view, x1;3. All specimens from East Wadi Ghonima section (at 132 m). G, Pseudaspidoceras sp.,honima section (24.7 m above the base of the formation).


a narrower venter (which may be related to compaction). Thespecimen described by Aly and Abdel-Gawad (2001) from Sinai hasa more inflated whorl section and the whorl height is very low.Furthermore, its ornamentation on the ventrolateral shoulders isvery weak or absent. Therefore, their specimen is only questionablyincluded here in the synonymy.

Occurrence. M. geslinianum occurs in the Upper Cenomanian ofEngland, France, Spain, Germany, the United States, Mexico, Brazil,Colombia, Niger, Angola, Tunisia and Morocco (e.g., Wright andKennedy, 1981; Kennedy and Juignet, 1981; Meister and Abdallah,2005; Lehmann and Herbig, 2009). It is the index of the epony-mous mid-Late Cenomanian ammonite standard zone. In Egypt, ithas been recorded from the Upper Cenomanian of the northEastern Desert and the Gulf of Suez region. The record from Sinai(Aly and Abdel-Gawad, 2001) is questionable.

Family: Vascoceratidae Douvillé, 1912Genus Vascoceras Choffat, 1898

Vascoceras adonense Choffat, 1898Figs. 8DeE, 10A

1898 Vascoceras adonense Choffat, p. 59, pl. 9, fig. 3; pl. 21,fig. 12a, b.

1969 Vascoceras cf. adonense Choffat; Freund and Raab, p. 32,pl. 5, fig. 1; text-fig. 7aeb.

2001 Vascoceras adonense Choffat; Kora et al., pl. 2, fig. 3.2003 Vascoceras adonense Choffat; Hewaidy et al., p. 343, pl. 1,

figs. 14e15.

Material. Three poorly preserved complete internal moulds(WG-18-1e3) from the Wadi Ghonima section (33.5 m above thebase of Maghra el Hadida Formation) and one poorly preservedcomplete internal mould (WA-15-1) from the Wadi Askhar section(at 106 m). All specimens come from the Lower Turonian V. prop-rium Zone of the Maghra el Hadida Formation.

Measurements.


WG-18-1 101.4 (100) 69.9 (68.9) 35.1 (34.6) 1.99 29.4 (28.9)WA-15-1 113.2 (100) 62.3 (55.0) 40.2 (35.5) 1.54 30.2 (26.6)


080215-8-1 116.0 (100) 42.0 (36.2) 59.1 (50.9) 0.71 21.5 (18.5)WG-15-1 127.1 (100) 49.0 (38.5) 64.7 (50.9) 0.75 26.3 (20.6)

Description. Medium-sized, slightly inflated, asymmetricallydistorted ammonites. Early whorls involute, later whorls moreevolute.Whorl section rounded and semilunar (Fig.10A). Umbilicuswide (about 29% of total diameter), relatively deep, with flat walls,inclined, and distinctive margin. Venter wide, rounded, withmoderately narrow flanks. Ornamentation and sutures absent dueto poor preservation.

Remarks. The studied specimens correspond closely to thosedescribed by Freund and Raab (1969) from Israel. However, they aresmaller and the umbilical tubercles are not seen due to poor preser-vation. Freund andRaab (1969) stated that their specimens are similarto the Portuguese holotype in dimensions, having weaker umbilicaltubercles, and considered the asymmetric shape of their specimens asan original feature that cannot be a result of deformation.

Occurrence. Vascoceras adonense has been recorded from theLower Turonian of Portugal and Israel (Choffat, 1898; Freund and

Fig. 8. AeC, Metoicoceras geslinianum (d’Orbigny, 1850), Upper Cenomanian Maghra el Hadispecimens fromWadi Ghonima section (24.7 m above the base of the formation). DeE, Vascoview, x1; E: lateral view, x1; WG-18-1 from Wadi Ghonima section (33.5 m above the baseLower Turonian Maghra el Hadida Formation. F: apertural view, x1; G: lateral view, x1; WA1904), Lower Turonian Maghra el Hadida Formation. H: lateral view, x1; I: ventral view, x1

Raab, 1969). In Egypt, it has been recorded from the Lower Turo-nian of the north Eastern Desert and the Gulf of Suez region.

Vascoceras cauvini Chudeau, 1909Figs. 9AeB, 10B

1909 Vascoceras cauvini Chudeau, p. 67, pl. 1, figs. 1a, 2a; pl. 2,figs. 3, 5; pl. 3, figs. 1b, 2b, 4.

1969 Paravascoceras cauvini (Chudeau); Freund and Raab,p. 20, pl. 3, figs. 1e3; text-fig. 5aeb.

1987 Vascoceras cauvini Chudeau; Kora and Hamama, pl.1, fig. 6.1989 Vascoceras cauvini Chudeau; Luger and Gröschke, p. 374,

pl. 40, figs. 3, 6, 8, 9; pl. 41, figs. 1e4; pl. 42, fig. 1; text-figs. 6g,h, 8c (with synonymy).

1996 Vascoceras cauvini Chudeau; Kassab, pl. 2, figs. 1, 2.1996 Paravascoceras cauvini (Chudeau); Zaborski, p. 65,

figs. 2e8 (with synonymy).2001 Vascoceras cauvini Chudeau; Aly and Abdel-Gawad, p. 40,

pl. 5, fig. 3aec.2001 Vascoceras cauvini Chudeau; El-Hedeny and Nafee, p.125,

pl. 2, figs. 3e4.2002 Vascoceras cauvini Chudeau; El-Hedeny, p. 406, figs.

4bec, 7f.2003 Vascoceras pioti (Peron and Fourtau); Hewaidy et al.,

p. 344, pl. 2, figs. 2e5.2004 Vascoceras cauvini Chudeau; Abdel-Gawad et al., pl. 4,

figs. 2e3, 5.2006 Vascoceras cauvini Chudeau; El Qot, p. 117, pl. 25,

figs. 2e3, 5.2007 Vascoceras cauvini Chudeau; Abdel-Gawad et al., pl. 2,

fig. 4.2008 Vascoceras cauvini Chudeau; Aly et al., p. 47, pl. 4,

figs. 3e4; text-fig. 3 (3).2009 Vascoceras cauvini Chudeau; Lehmann and Herbig, p. 74,

pl. 2, figs. aeb, men.

Material. Four moderately preserved incomplete internalmoulds (WG-15-1e3, 080215-8-1) from the Upper CenomanianV. cauvini Zone of the Maghra el Hadida Formation in the WadiGhonima section (29 m above the base of the formation).

Measurements.

Description. Involute, medium-sized. Whorl section compressed(Fig. 10B), higher than wide, with slightly inclined flanks andmaximum breadth close to the umbilical shoulder. Umbilicusrelatively wide (about 20.5% of total diameter), deep, with feeblyrounded shoulders and vertical walls. Venter well rounded. Orna-mentation consists of faint, weak marginal ribs crossing the venter.Sutures not preserved.

Remarks. The studied specimens resemble the material of Lugerand Gröschke (1989) from the Egyptian Eastern Desert, but arelarger in size. They are also similar to the material of Freund andRaab (1969) from Israel, being slightly more evolute with weakerornament. However, all these differences are small considered the

da Formation. A, C: lateral view, x1.5; 080215-3-1; B: lateral view, x1.5; 080215-3-2. Allceras adonense Choffat, 1898, Lower Turonian Maghra el Hadida Formation. D: aperturalof Maghra el Hadida Formation). FeG, Vascoceras durandi (Thomas and Peron, 1889),

-14-4.5 from Wadi Askhar section (at 105 m). HeI, Vascoceras pioti (Peron and Fourtau,; WG-17-4 from Wadi Ghonima section (at 32 m).

Fig. 10. Whorl sections of A,Vascoceras adonenseChoffat,1898, Lower TuronianMaghra elHadida Formation; B,Vascoceras cauviniChudeau,1909,Upper CenomanianMaghra elHadidaFormation; C, Vascoceras durandi (Thomas and Peron, 1889), Lower Turonian Maghra el Hadida Formation; D, Vascoceras pioti (Peron and Fourtau, 1904), Lower Turonian Maghra elHadida Formation; E-H, Vascoceras proprium (Reyment, 1954), Lower Turonian Maghra el Hadida Formation. A, specimenWG-18-1 from theWadi Ghonima section (33.5 m above thebase ofMaghra elHadida Formation); B, specimenWG-15-1 fromtheWadiGhonima section (29mabove the base of the formation); C, specimenWA-14-4 fromtheWadiAskhar section(at 105m);D, specimenWG-17-4 from theWadiGhonima section (at 32m); E, specimenWG-17-1; F, specimenWG-17-2;G, specimenWG-17-3; specimens E, FandGare from theWadiGhonima section (31 m above the base of Maghra el Hadida Formation); H, specimen WA-14-1 from the Wadi Askhar section (at 104 m). Not to scale.


WA-14-4 126.1 (100) 56.6 (44.8) 61.3 (48.6) 0.92 38.1 (30.2)


intraspecific variability inwhorl shape and ornament as pointed outby Schöbel (1975). Further detailed treatments of V. cauvini fromNiger and Nigeria can be found in Meister et al. (1992) and Zaborski(1996).

Occurrence. According to Luger and Gröschke (1989, p. 375),V. cauvini is widely known fromUpper Cenomanian strata of centraland northern Africa, the Middle East, and southern America withrecords from Peru, Niger, Nigeria, Angola, Algeria, Morocco, Israel(e.g., Meister et al., 1992; Lehmann and Herbig, 2009). It also occursin France. In Egypt, the species has been recorded from the UpperCenomanian of the north Eastern Desert, Sinai and the Gulf of Suezregion. In the study area, V. cauvini is recorded from strata of latestCenomanian age, below the entry of V. proprium and above the lastoccurrence of M. geslinianum (Nagm, 2009; Nagm et al., 2010).The V. cauvini Zone is, thus, equivalent to the N. juddii Zone in thestandard ammonite zonation (cf. Gradstein et al., 2004).

Vascoceras durandi (Thomas and Peron, 1889)Figs. 8FeG, 9CeD, 10C

1889 Pachydiscus durandi Thomas and Peron, p. 27, pl. 18,figs. 5e8.

1969 Vascoceras durandi (Thomas and Peron); Freund andRaab: p. 29, text-fig. 6hei.

1989 Vascoceras durandi (Thomas and Peron); Luger andGröschke: p. 376, pl. 43, figs. 1, 2, text-fig. 8a.

Fig. 9. AeB, Vascoceras cauvini Chudeau, 1909, Upper Cenomanian Maghra el Hadida Form(29 m above the base of the formation). CeD, Vascoceras durandi (Thomas and Peron, 1889),x0.75; WA-14-4 from Wadi Askhar section (at 105 m).

1994 Vascoceras durandi (Thomas and Peron); Chancelloret al., p. 48, pl. 2, fig.1; pl. 10, figs. 1e4; pl. 11, figs. 1e2; pl.12, figs. 1e3; pl. 13, figs. 3e4; pl. 14, figs. 2, 5 (see forsynonymy).

2003 Vascoceras durandi (Thomas and Peron); Hewaidy et al.,p. 343, pl. 2, figs. 1e2.

?2004 Vascoceras cf. durandi (Thomas and Peron); Abdel-Gawad et al., pl. 4, figs. 4.

2006 Vascoceras cf. durandi (Thomas and Peron); El Qot, p. 118,pl. 25, fig. 4; pl. 26, fig. 1aeb.

2009 Vascoceras durandi (Thomas and Peron); Lehmann andHerbig, p. 73, pl. 2, figs. geh.

Material. Two moderately preserved complete internal moulds(WA-14-4, -4.5) from the Lower Turonian Maghra el HadidaFormation of Wadi Askhar section (at 105 m).

Measurements.

Description. Involute, medium-sized, and compressed. Whorlshigher than wide, body chamber large. Whorl section becomestriangular at the body chamber (Fig. 10C). Flanks narrow and slightlyconvex. Umbilicuswide (about 30% of total diameter) andmoderately

ation. A: apertural view, x1; B: lateral view, x1; WG-15-1 from Wadi Ghonima sectionLower Turonian Maghra el Hadida Formation. C: lateral view, x0.75; B: apertural view,

Fig. 12. Whorl sections of AeB, Neoptychites cephalotus (Courtiller, 1860); C, Fagesia cf. peroni Pervinquière,1907. A, specimenWG-23-1; B, specimenWG-23-2; C, specimenWG-23-3.All specimens from the Lower Turonian Maghra el Hadida Formation of the Wadi Ghonima section (44 m above the base of Maghra el Hadida Formation). Not to scale.


deep, with rounded shoulders and vertical walls. Venter narrowlyrounded.Ornamentationandsutures absentdue topoorpreservation.

Remarks. Vascoceras durandi can be distinguished by its largesize, absence or early loss of umbilical tubercles and a subangularumbilical shoulder with subtriangular whorl section at maturity(Chancellor et al., 1994). The studied specimens closely correspondto those described from Israel (Freund and Raab, 1969), Tunisia(Chancellor et al., 1994) and the north Eastern Desert of Egypt(Luger and Gröschke, 1989).

Occurrence. V. durandi occurs in the Upper Cenomanian to LowerTuronian of Portugal (Berthou et al., 1985) and in the Lower Turo-nian of Israel, Tunisia, Algeria and Morocco (Meister and Abdallah,2005; Lehmann and Herbig, 2009). Furthermore, it is known fromSpain, Angola(?), Japan, Mexico, and Brazil (Berthou et al., 1985). InEgypt, it has been recorded from the Lower Turonian of the northEastern Desert and from Sinai.

Vascoceras pioti (Peron and Fourtau, 1904)Figs. 8HeI, 10D

1904 Ammonites pioti Peron and Fourtau in Fourtau, p. 275, pl.1,figs. 4e5.

1969 Vascoceras pioti (Peron and Fourtau); Freund and Raab,p. 28, pl. 4, figs. 1e9; text-fig. 6deg.

Material.One incomplete,butwellpreservedspecimen(WG-17-4)and one poorly preserved incomplete internal mould (WG-17-5)from the Lower Turonian Maghra el Hadida Formation of the WadiGhonima section (at 32 m).

Measurements.


WG-17-4 47.6 (100) 17.2 (36.1) 30.0 (63.0) 0.57 4.7 (9.8)WG-17-5 57.1 (100) 22.4 (39.2) 36.3 (63.5) 0.61 5.6 (9.8)

Description. Very involute, relatively small. Whorls stronglycompressed (Fig. 10D). Maximum whorl breadth near the

Fig. 11. AeF, Vascoceras proprium (Reyment, 1954), Lower Turonian Maghra el Hadida Formatview, x1; WG-17-1; E: lateral view, x1; F: apertural view, x1; WG-17-3. All specimens fromNeoptychites cephalotus (Courtiller, 1860), Lower Turonian Maghra el Hadida Formation. G: lathe base of Maghra el Hadida Formation).

umbilicus. Umbilicus small (about 10% of diameter), with verticalwalls and well defined borders. Venter flat to slightly convex,clearly separated from the flanks. The body chamber occupiesalmost one complete whorl. Ornamentation consists of numerousribs, weak at the umbilicus and strong on the middle and outerflanks, each rib forming small clavi at the ventrolateral shoulder.Sutures not seen in the present specimens.

Remarks. The studied material is very close to that described byFreund and Raab (1969) from Israel. These authors stated that theirspecimens are identical to the holotype as described and figured byPeron and Fourtau (in Fourtau 1904, p. 275, pl. 1, figs. 4e5). Thespecimens described by Hewaidy et al. (2003, p. 344, pl. 2, figs. 2e5)as Vascoceras pioti have completely rounded venters, withoutventrolateral shoulders, and with wide, deep umbilici, while V. piotihas a flat venter, clearly separated from the flanks, with very smalland shallow umbilici. Therefore, their specimen are not consideredhere as V. pioti, but may be related to V. cauvini.

Occurrence. V. pioti has been recorded from the Lower Turonianof Israel and Egypt (Freund and Raab, 1969; Nagm, 2009).

Vascoceras proprium (Reyment, 1954)Figs. 10EeH, 11AeF

1954 Pachyvascoceras proprium Reyment, p. 258, pl. 5, fig. 1;text-fig. 3d.

1987 Vascoceras proprium (Reyment); Kennedy et al., p. 46,pl. 4, figs. 1e15, 18e19; pls. 5e6; text-figs. 8aec, 9 (withfull synonymy).

1994 Vascoceras proprium (Reyment); Kassab, p. 120, fig. 5(1e4).

1994 Vascoceras proprium (Reyment); Kassab and Ismael,p. 226, fig. 2 (10e12).

1996 Vascoceras proprium (Reyment); Kassab, pl. 2, figs. 5e7.2001 Vascoceras proprium (Reyment); Aly and Abdel-Gawad,

p. 39, pl. 5, fig. 4aec.2001 Vascoceras proprium (Reyment); Kora et al., pl. 2, fig. 3.2002 Vascoceras proprium (Reyment); El-Hedeny, p. 407,

figs. 5aeb, 7c.

ion. A: lateral view, x1; B: apertural view, x1; WG-17-2; C: lateral view, x1; D: aperturalWadi Ghonima section (31 m above the base of Maghra el Hadida Formation). GeH,teral view, x1; H: ventral view, x1; WG-23-1 from Wadi Ghonima section (44 m above


2003 Vascoceras proprium (Reyment); Hewaidy et al., p. 344,pl. 2, figs. 3e5.

2008 Vascoceras proprium (Reyment); Aly et al., p. 49, pl. 5,figs. 1e2; text-fig. 3 (4).

Material. Three moderately preserved incomplete internalmoulds (WA-14-1e3) fromtheWadiAskhar section (at 104m), threemoderately preserved complete internalmoulds (WG-17-1e3) fromtheWadiGhonima section (31mabove the base ofMaghra elHadidaFormation), and two poorly preserved incomplete internal moulds(080217-23.5-1e2) from the East Wadi Ghonima section (at 117 m).All specimens come fromthe LowerTuronianV. propriumZoneof theMaghra el Hadida Formation.

Measurements.


WA-14-1 71.3 (100) 41.8 (58.6) 36.6 (51.3) 1.14 15.6 (21.8)WA-14-2 83.5 (100) 48.5 (58.0) 41.5 (49.7) 1.16 24.6 (29.4)WA-14-3 62.8 (100) 32.6 (51.9) 27.5 (43.7) 1.18 13.6 (21.6)WG-17-1 50.8 (100) 44.3 (87.2) 28.6 (56.2) 1.54 7.1 (13.9)WG-17-2 66.6 (100) 47.8 (71.7) 35.0 (52.5) 1.36 11.8 (17.7)WG-17-3 69.3 (100) 46.5 (48.2) 40.0 (57.7) 1.16 7.7 (11.1)


WG-23-1 110.3 (100) 32.7 (29.6) 54.1 (49.0) 0.60 6.4 (5.8)WG-23-2 96.6 (100) 55.1 (57.0) 53.5 (55.3) 1.02 9.3 (9.6)WG-23-3 109.2 (100) 60.9 (55.7) 48.1 (44.0) 1.26 10.2 (9.3)WG-23-4 93.3 (100) 43.1 (46.1) 47.0 (50.3) 0.91 9.6 (10.2)WG-23-5 95.2 (100) e 45.9 (48.2) e 10.6 (10.7)

Description. Involute, medium-sized, varying from slightlycompressed tomoderately inflated.Whorl sectionvaries fromslightlyrounded to triangular (Fig.10EeH). Shape of venter broadly arched tofairly narrow. Flanks short and convex. Umbilici relatively deep, withvertical to slightly undercut walls and well rounded umbilicalshoulder. Umbilical width ranges from 11% to 29% of diameter.Ornamentation and suture line absent due to poor preservation.

Remarks. V. proprium (Reyment) was discussed in detail byKennedy et al. (1987). They studied a large number of specimensfrom a single horizon and the same locality and concluded thattheir intermediate specimens match well with the holotype ofReyment (1954). They placed a range of associated forms separatedby Reyment (1954, 1955) and Barber (1957) in the synonymy of thisspecies. The present material also shows variations in whorlsection, thus supporting the opinion of Kennedy et al. (1987) ofa considerable morphological variation within this species.

Occurrence. V. proprium has been recorded from the LowerTuronian of Nigeria, northern Mexico and Texas (Kennedy et al.,1987) as well as Jordan (Aly et al., 2008). In Egypt, it has beenrecorded from the Lower Turonian of the north Eastern Desert, theGulf of Suez region and from Sinai. It characterizes the lowermostTuronian ammonite zone of the study area (Nagm et al., 2010).

Genus Neoptychites Kossmat, 1895

Neoptychites cephalotus (Courtiller, 1860)Figs. 11GeH, 12AeB, 13AeB

1860 Ammonites cephalotus Courtiller, p. 248, pl. 2, figs. 1e4.1979b Neoptychites cephalotus (Courtiller); Kennedy and

Wright, p. 670, pl. 82, figs. 3e5; pl. 83, figs. 1e3; pl. 84,fig. 3; pl. 85, figs. 1e5; pl. 86, figs. 4e5; text-fig. 2 (withfull synonymy).

1979b Neoptychites xetriformis Pervinquière; Kennedy andWright, p. 679, pl. 84, figs. 1e2; pl. 86, figs. 1e3.

Fig. 13. AeB, Neoptychites cephalotus (Courtiller, 1860), Lower Turonian Maghra el Hadidasection (44 m above the base of Maghra el Hadida Formation). CeD, Fagesia cf. peroni PerD: apertural view, x0.75; WG-24-1 from Wadi Ghonima section (at 39.5 m above the baseLower Turonian Maghra el Hadida Formation. E: lateral view, x1; F: ventral view, x1; G: ap

1990 Neoptychites cephalotus (Courtiller); Robaszynski et al.,p. 266, pl. 20, figs. 2e3; pl. 21, fig. 3.

1994 Neoptychites cephalotus (Courtiller); Chancellor et al.,p. 70, pl. 16, figs. 1e9; pl. 17, figs. 1e5; pl. 18, figs. 1e3;pl. 26, figs. 2e4.

1994 Neoptychites xetriformis Pervinquière; Kassab, p. 121,fig. 5 (8).

2001 Neoptychites cephalotus (Courtiller); Aly and Abdel-Gawad, p. 39, pl. 6, fig. 1.

2008 Neoptychites cephalotus (Courtiller); Aly et al., p. 49, pl. 6,fig. 1aeb.

2008 Neoptychites cephalotus (Courtiller); Kennedy et al.,p. 159, pl. 3, figs. 6e8; pl. 6, figs. 1e5.

2009 Neoptychites cephalotus (Courtiller); Barroso-Barcenillaand Goy, p. 34, fig. 9 (4e6), fig. 10 (1e3) (see forcomplete synonymy).

Material. Three moderately preserved complete internal moulds(WG-23-1e3) and two well preserved incomplete shells (WG-23-4e5) from the Wadi Ghonima section (44 m above the base ofMaghra el Hadida Formation) as well as two poorly preservedincomplete internal moulds (080217-26-3e5) from the East WadiGhonima section (at 132 m). All specimens come from the LowerTuronianChoffaticeras spp. Zone of theMaghra el Hadida Formation.

Measurements.

Description. There are two forms ofNeoptychites cephalotus in thematerial studied, compressed and depressed ones (Fig. 12AeB). Inthe medium-sized compressed form, coiling is involute, whorls arestrongly compressed, with rectangular whorl section and small,shallow umbilicus (about 6% of total diameter) with low androunded umbilical shoulder. Flanks are vertical on the inner andslightly convex on the outer part, venter narrowly rounded. Orna-mentation consists ofweak foldingon theouterflank. Thedepressedform is likewisemedium-sized,with involute coiling, butwhorls aredepressed, with rounded whorl section, the umbilicus being wider(up to about 10.5% of diameter), walls are convex, venters are morewidely rounded. Ornamentation consists of strong folding on theouter flank. Suture lines not seen in both forms.

Remarks. Kennedy andWright (1979b) revised different forms ofNeoptychites from Touraine, France, and recognized two species,N. cephalotus and Neoptychites xetriformis, considering the latterspecies different in having stronger ribs. However, according toChancellor et al. (1994, p. 72), N. xetriformis is the microconch of thespecies and thus a junior synonym of N. cephalotus. Kennedy andWright (1979b, p. 674) demonstrated that N. cephalotus passesthough three distinctive ontogenetic stages. The third stage is wellrepresented in the studied specimens, where compressed, smoothdepressed, and ribbed depressed forms have been recorded. Thecompressed form (Fig. 11GeH) is identical to the material ofKennedy and Wright (1979b, pl. 83) and Chancellor et al. (1994,

Formation. A: lateral view, x1; B: apertural view, x1; WG-23-2 from Wadi Ghonimavinquière, 1907, Lower Turonian Maghra el Hadida Formation. C: lateral view, x0.75;of Maghra el Hadida Formation). EeG, Thomasites rollandi (Thomas and Peron, 1889),ertural view, x1; WG-23-6 from Wadi Ghonima section (at 44 m).

Fig. 15. Whorl sections and suture lines of AeD, Thomasites rollandi (Thomas and Peron, 1889); EeF, Wrightoceras munieri (Pervinquière, 1907). A, D, specimen 080217-25.5-3 fromthe East Wadi Ghonima section (at 127 m); B, WG-23-13; C, specimen WG-23-6; B and C from the Wadi Ghonima section (at 44 m); EeF, specimen 080216-18-3 from the WadiGhonima section (at 47 m). All specimens from the Lower Turonian Maghra el Hadida Formation. Not to scale.


WG-24-1 123.1 (100) 89.1 (72.3) 56.2 (45.6) 1.58 38.9 (31.6)


pl. 18). The depressed form (Fig. 13AeB) is identical to specimens ofKennedy and Wright (1979b, pl. 86, fig. 1, 2).

Occurrence. According to Chancellor et al. (1994, p. 72) andKennedy et al. (2008, p. 160), the total range of N. cephalotus is mid-Lower Turonian to lower Middle Turonian, with records fromFrance, Spain, Morocco, Algeria, Tunisia, Egypt, Israel, Syria,Cameroon, Madagascar, southern India, the Western Interior of theUnited States, northern Mexico, Trinidad, Venezuela, Colombia,Brazil, Niger and Nigeria. Recently, it also has been recorded fromJordan (Aly et al., 2008). In Egypt, it has been recorded from theLower Turonian of Sinai, the Gulf of Suez region and the northEastern Desert (see Nagm, 2009).

Genus Fagesia Pervinquière, 1907

Fagesia cf. peroni Pervinquière, 1907Figs. 12C, 13CeD

cf. 1907 Fagesia Peroni Pervinquière, p. 329, pl. 20, figs. 7e8.cf. 1994 Fagesia peroni Pervinquière; Chancellor et al., p. 66,

pl. 14, figs. 6e10.cf. 2008 Fagesia peroni Pervinquière; Kennedy et al., p. 157, pl. 3,

figs. 1e2; pl. 4, figs. 1e3; pl. 5, figs. 1e2; pl. 7, figs. 4e6;pl. 9, figs. 7e9; text-figs. 5e6.

Material. Two poorly preserved incomplete internal moulds(080217-25.5-1e2) from the East Wadi Ghonima section (at 127 m)

Fig. 14. AeD, Thomasites rollandi (Thomas and Peron, 1889), Lower Turonian Maghra el HadGhonima section (at 127 m); C: lateral view, x1; D: apertural view, x1; WG-23-13 from WTuronian Maghra el Hadida Formation. E: lateral view, x1; F: ventral view, x1; 080216-18-2;x1; 080216-18-3. All specimens from Wadi Ghonima section (at 47 m).

and one poorly preserved complete internal mould (WG-24-1) fromthe Wadi Ghonima section (at 39.5 m above the base of Maghra elHadida Formation). All specimens come from the Lower TuronianChoffaticeras spp. Zone of the Maghra el Hadida Formation.

Measurements.

Description. Involute, large-sized, and inflated. Whorl section(Fig. 12C) very depressed (Wb/Wh¼ 1.58), with very narrow flanks.Umbilicus wide (31.6% of total diameter), fairly deep, withdistinctive rounded umbilical shoulder and traces of umbilicaltubercles. Venter broadly rounded. Ornamentation and suture lineabsent due to poor preservation.

Remarks. The two studied specimens are larger than the lecto-type (see Chancellor et al. 1994, pl. 14, figs. 7, 8, 10) and the orna-mentation is not visible due to poor preservation. Thus, they arekept in open nomenclature. Well preserved material fromMoroccohas recently been illustrated and described by Kennedy al. (2008).

Occurrence. Fagesia peroni has been recorded from the LowerTuronian of Tunisia, Algeria and Morocco (Pervinquière, 1907;Chancellor et al., 1994; Kennedy et al., 2008). According toChancellor et al. (1994), there are further records from Romania,Colombia, Venezuela and Central Asia of the former Soviet Union.This is the first record of Fagesia cf. peroni from Egypt.

ida Formation. A: lateral view, x1; B: ventral view, x1; 080217-25.5-3 from East Wadiadi Ghonima section (at 44 m). EeJ, Wrightoceras munieri (Pervinquière, 1907), LowerG: lateral view, x1; H: ventral view, x1; 080216-18-1; I: lateral view, x1; J: ventral view,

Fig. 17. AeB, whorl section and suture line of Coilopoceras requienianum (d’Orbigny,1841), WG-37-1 from the Upper Turonian Maghra el Hadida Formation of WadiGhonima section. Not to scale.


Family: Pseudotissotiidae Hyatt, 1903Subfamily: Pseudotissotiinae Hyatt, 1903Genus Thomasites Pervinquière, 1907

Thomasites rollandi (Thomas and Peron, 1889)Figs. 13EeG, 14AeD, 15AeD

1889 Pachydiscus rollandi Thomas and Peron, p. 25, pl. 17, figs.1e3.

1969 Thomasites rollandi (Thomas and Peron) and varieties ofPervinquière; Freund and Raab, p. 43, text-fig. 9fek.

1969 Thomasites jordani and varieties of Pervinquière; Freundand Raab, p. 45, text-figs. 10aec.

1994 Thomasites rollandi (Thomas and Peron); Chancellor et al.,p. 75, pl. 19, figs. 1e2; pl. 20, figs. 1e12; pl. 21, figs. 1e9;pl. 22, figs. 1e6; pl. 23, figs. 1e9; text-fig. 14aef (see forcomplete synonymy).

2004 Thomasites rollandi (Thomas and Peron); Abdel-Gawadet al., pl. 3, fig. 1.

2006 Thomasites rollandi (Thomas and Peron); El Qot, p. 122,pl.27, figs. 4,5; pl. 28, fig. 1.

2007 Thomasites rollandi (Thomas and Peron); Abdel-Gawadet al., pl. 3, fig. 1.

2009 Thomasites rollandi (Thomas and Peron); Lehmann andHerbig, p. 75, pl. 2, figs. eef (with additional synonymy).

Material. 9 complete and incomplete, moderately to wellpreserved specimens (partially with shell preservation, butcommonly internal moulds, WG-23-6e14) from theWadi Ghonimasection (at 44 m) and three moderately preserved incompleteinternal moulds (080217-25.5-3e5) from the East Wadi Ghonimasection (at 127 m). All specimens come from the Lower TuronianChoffaticeras spp. Zone of the Maghra el Hadida Formation.

Measurements.


WG-23-6 38.8 (100) 28.7 (73.9) 20.6 (53.0) 1.93 11.6 (29.8)WG-23-7 56.1 (100) 37.1 (66.1) 32.6 (58.1) 1.13 7.1 (12.6)WG-23-8 48.7 (100) 39.8 (81.7) 31.1 (63.8) 1.27 6.8 (13.9)WG-23-9 51.4 (100) 33.6 (65.3) 30.4 (59.1) 1.10 7.4 (14.3)WG-23-10 37.1 (100) 27.0 (72.7) 18.8 (50.6) 1.43 5.7 (15.3)WG-23-11 92.0 (100) 54.2 (58.9) 50.9 (55.3) 1.06 10.3 (11.1)WG-23-12 60.7 (100) 41.4 (68.2) 36.1 (59.4) 1.14 7.5 (12.3)WG-23-13 55.6 (100) 29.0 (52.1) 31.2 (56.1) 0.92 5.4 (9.7)WG-23-14 51.7 (100) 29.9 (57.8) 30.3 (58.6) 0.98 7.2 (13.9)080217-25.5-3 52.1 (100) 22.2 (42.6) 27.5 (52.7) 0.80 5.1 (9.7)

Description. Medium-sized, very involute. Whorl sectioncompressed to globose and inflated. Umbilicus moderately wide tosmall (about 30e10% of total diameter), deep, with a high, sub-vertical wall; umbilical shoulder narrowly rounded, with distinc-tive tubercles. Greatest width at the umbilical shoulder. The innerflanks are strongly inflated, but the outer flanks are flattened andconvergent. The ventrolateral shoulders are rounded, with a row ofsmall bullae on each side (19 bullae/whorl in specimen WG-23-6)which form coarse, blunt ribs that are strong on the outer flank andweaker on the inner flank. Venter rounded to nearly flat. Suture linecomposed of four saddles, the first one being the broadest(Fig. 15AeD).

Remarks. According to Chancellor et al. (1994) who examinedPeron’s types plus 150 other specimens of Thomasites from Tunisiathere is considerable intraspecific variation in T. rollandi. They thus

Fig. 16. AeC, Coilopoceras requienianum (d’Orbigny, 1841), Upper Turonian Maghra el Hadidathe Wadi Ghonima section (at 210.5 m).

also included the numerous varieties and new species’ ofPervinquière (1907) and Douvillé (1928) in T. rollandi. The speci-mens described herein are very similar to the Tunisian materialdescribed by Chancellor et al. (1994) and Meister and Abdallah(1996, 2005), and nothing needs to be added here.

Occurrence. T. rollandi has been recorded from the Lower Turo-nian of Morocco, Tunisia, Algeria, Israel, Jordan, Lebanon, Syria,France, Spain(?), and England, and it may also occur in Colombiaand Tadzhikistan (e.g., Wright and Kennedy, 1981; Chancellor et al.,1994; Lehmann and Herbig, 2009). In Egypt, it has been recordedfrom the Lower Turonian of Sinai and the Eastern Desert.

Genus Wrightoceras Reyment, 1954

Wrightoceras munieri (Pervinquière, 1907)Figs. 14EeJ, 15EeF

1907 Hoplitoides munieri Pervinquière, p. 217, pl. 10, figs. 1, 2;text-fig. 83.

1987 Wrightoceras munieri (Pervinquière); Kennedy et al., p. 58,pl. 10, figs. 9e11; text-fig. 2e.

1994 Wrightoceras munieri (Pervinquière); Chancellor et al.,p. 96, pl. 26, figs. 1, 5, 8; pl. 28, figs. 1e4; pl. 29, figs. 3e8;pl. 36, figs. 1e2; text-figs. 18geh, 19hei (with synonymy).

2007 Wrightocerasmunieri (Pervinquière); Barroso-Barcenilla andGoy, p. 480, fig. 10(3, 4) (see for further synonymy).

2008 Wrightoceras munieri (Pervinquière); Kennedy et al., p.163, pl. 2, fig. 7; pl. 3, fig. 3; pl. 7, fig. 3; pl. 9, figs. 1e6,10e11.

Material. Two well preserved incomplete shells (080216-18-1e2) and five well preserved complete and incomplete internalmoulds (080216-18-3e7) from theWadi Ghonima section (at 47 m)as well as four moderately preserved incomplete internal moulds(080217-27-1e4) from the EastWadi Ghonima section (at 143.5 m).All specimens come from the Lower Turonian W. munieri Zone ofthe Maghra el Hadida Formation.

Formation. A: apertural view, x1; B: lateral view, x1; C: ventral view, x1; WG-37-1 from


Measurements.


080216-18-1 74.7 (100) 22.1 (29.5) 45.3 (60.6) 0.48 5.4 (7.2)080216-18-3 121.9 (100) 39.4 (32.3) 76.4 (62.6) 0.51 7.9 (6.4)


WG-37-1 140.2 (100) 43.9 (39.4) 85.6 (61.0) 0.51 6.9 (4.9)WG-37-2 125.5 (100) 38.3 (30.5) 79.8 (63.5) 0.47 UnavailableWG-37-6 111.2 (100) 36.5 (32.8) 75.7 (68.0) 0.48 6.6 (5.9)

Description. Very involute, medium-sized. Whorls verycompressed. Umbilicus small, umbilical shoulder very narrowlyrounded. Maximum width at umbilical shoulder. Whorl wallsslightly convex. Venter narrow, concave, edges developed intosharp keels. All specimens lack ornamentation. The suture line iswell preserved; the first lateral saddle is the highest and the widestone, being divided by accessory lobes (Fig. 15EeF).

Remarks. The studied specimens closely correspond to theholotype as figured by Pervinquière (1907) and to those describedby Kennedy et al. (1987) as well as to the Tunisian material ofChancellor et al. (1994).

Occurrence. According to Chancellor et al. (1994, p. 100) andKennedy et al. (2008, p. 164), W. munieri is known from the LowerTuronian of Tunisia, Niger, Nigeria, Morocco, Spain, Peru, Venezuela,Colombia, Texas, and Mexico. This is the first record of W. munierifrom Egypt. It has been recorded from the upper Lower Turonianwhere it defines a total range zone (Nagm, 2009; Nagm et al., 2010).

Genus Choffaticeras Hyatt, 1903

Systematic description of the genus Choffaticeras, withcomplete discussion of its taxonomic problems as well as cali-bration of its evolutionary pattern based on high resolution stra-tigraphy of the Lower Turonian in the study area, will be treated ina separate paper. The following taxa of the genus ChoffaticerasHyatt, 1903 have been recognized: Choffaticeras (Choffaticeras)meslei (Peron, 1897), Choffaticeras (Choffaticeras) quaasi (Peron,1904), Choffaticeras (Choffaticeras) pavillieri (Pervinquière, 1907),Choffaticeras (Choffaticeras) securiforme (Eck, 1909), Choffaticeras(Choffaticeras) segne (Solger, 1903), Choffaticeras (Choffaticeras)sinaiticum (Douvillé, 1928), Choffaticeras (Leoniceras) barjonai(Choffat, 1898), Choffaticeras (Leoniceras) luciae (Pervinquière,1907), and Choffaticeras (Leoniceras.) philippii (Solger, 1904).

Family: Coilopoceratidae Hyatt, 1903Genus Coilopoceras Hyatt, 1903

Coilopoceras requienianum (d’Orbigny, 1841)Figs. 16AeC, 17, 18AeC

1841 Ammonites requienianus d’Orbigny, p. 315, pl. 93, figs. 1e4.1984 Coilopoceras requienianum (d’Orbigny); Kennedy and

Wright, p. 282, pls. 35-36, text-figs. 1e5 (see for furthersynonymy).

1989 Coilopoceras requienianum (d’Orbigny); Luger andGröschke: p. 388, pl. 46, text-figs. 6a,e, 11, 12, 13aec.

1987 Coilopoceras requienianum (d’Orbigny); Kora andHamama, pl. 1, fig. 9.

2001 Coilopoceras requienianum (d’Orbigny); Kora et al., pl. 2,figs. 11, 12.

2003 Coilopoceras requienianum (d’Orbigny); Hewaidy et al.,p. 354, pl. 6, figs. 4e5.

2004 Coilopoceras requienianum (d’Orbigny); Abdel-Gawadet al., pl. 1, figs. 3, 5.

Fig. 18. AeC, Coilopoceras requienianum (d’Orbigny, 1841), Upper Turonian Maghra el Hadidasection. DeE, Eubostrychoceras sp., Lower Turonian Maghra el Hadida Formation. D, E: x1;

2006 Coilopoceras requienianum (d’Orbigny); El Qot, p. 128,pl. 29, figs. 1aeb, 2e3.

2007 Coilopoceras requienianum (d’Orbigny); Abdel-Gawadet al., pl. 3, figs. 2, 7.

Material. Two complete and three incomplete, moderatelypreserved internal moulds (WG-37-1e5) and one moderatelypreserved, incomplete shell (WG-37-6) from the Wadi Ghonimasection as well as one complete, poorly preserved internal mould(080217-42-1) from the East Wadi Ghonima section (at 210.5 m)and one incomplete, moderately preserved internal mould(St.A-2-1) from the Saint Anthony section.

Measurements.

Description. Very involute, oxycone, medium-sized. Whorlscompressed, with lanceolate shape. Walls nearly vertical, withrounded shoulders and sharpened venter. Maximum width tendsto be situated at mid-flank. Very narrow umbilicus, with low androunded shoulders. Ornament of a few weak ribs on the earlywhorls only on specimen WG-37-1, other specimens are smooth.The suture line (Fig. 17) shows a broad and shallow external lobe,and a very broad, open and bifid lateral lobe.

Remarks. In their re-description of the type material, Kennedyand Wright (1984) suggested the existence of sexual dimorphismexpressed by smooth and ribbed forms in C. requienianum. Thespecimens studied herein closely correspond to the type material(cf. Kennedy and Wright, 1984) as well as to those described byLuger and Gröschke (1989) fromWadi Qena (Egypt). However, theyare slightly more inflated and the smoothmorphotype is dominant.

Occurrence. C. requienianum has been recorded from the UpperTuronian of Madagascar, North and South America, and fromWestern Europe (France, Spain; e.g., Collignon, 1965; Cobban andHook, 1980; Kennedy and Wright, 1984). In Egypt, it has beenrecorded from the Upper Turonian of the north Eastern Desert, theGulf of Suez region and from Sinai.

Suborder: Ancyloceratina Wiedmann, 1966Superfamily: Turrilitoidea Gill, 1871Family: Nostoceratidae Hyatt, 1894Genus Eubostrychoceras Matsumoto, 1967

Eubostrychoceras sp.Fig. 18DeE

Material. One moderately preserved incomplete compositemould (080217-27-1) from the Lower Turonian W. munieri Zone ofthe Maghra el Hadida Formation of the East Wadi Ghonima section(at 143.5 m).

Description. Small-sized, with tight coiling. Whorl sectionrounded. Ornamentation consists of simple ribs, withouttuberculation.

Remarks. The present specimen is similar to Eubostrychocerassaxonicum but differs in having coarser ribs (see Kaplan andSchmid, 1988). It also resembles Eubostrychoceras matsumotoi (see

Formation. A, C: lateral view, x1; B: apertural view, x1; WG-37-6 from Wadi Ghonima080217-27 from East Wadi Ghonima section (at 143.5 m).

Table 1Summary of the palaeogeographic distribution of CenomanianeTuronian ammonites recorded from the Wadi Araba (this study); (þ) means that the individual taxon has been recorded in the mentioned area.

Numericalsymbols ofspecies on themap (Fig. 19)

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

Locality Species

Neolobitesvibrayeanus

Thomelites(as a genus)

Euomphalocerasseptemseriatum

Kamerunocerasturoniense

Pseudaspidoceras(as a genus)

Metoicocerasgeslinianum

Vascocerasadonense

Vascocerascauvini

Vascocerasdurandi

Vascoceraspioti

Vascocerasproprium

Neoptychitescephalotus

Fagesiacf. peroni

Thomasitesrollandi

Wrightocerasmunieri

Coilopocerasrequienianum

Eubostrychoceras(as a genus)

Middle East þ þ þ þ þ þ þ þ þ þOman þSaudi Arabia þJordan þ þ þ þSyria þ þ þLebanon þ þIsrael þ þ þ þ þ þ þ þ þ

North Africa þ þ þ þ þ þ þ þ þ þ þ þTunisia þ þ þ þ þ þ þ þ þ þAlgeria þ þ þ þ þ þ þMorocco þ þ þ þ þ þ þ þ

West and South Africa þ þ þ þ þ þ þ þ þ þ þNiger þ þ þ þ þ þNigeria þ þ þ þ þCameroon þ þAngola þ þ þ þMadagascar þ þ þ þ

Europe þ þ þ þ þ þ þ þ þ þ þ þ þ þ þRomania þGermany þ þ þEngland þ þ þ þ þ þ þFrance þ þ þ þ þ þ þ þ þ þSpain þ þ þ þ þ þ þ þ þPortugal þ þ þ þ

North America þ þ þ þ þ þ þ þ þ þUnited States þ þ þ þ þ þ þ þMexico þ þ þ þ þ þ þ

South America þ þ þ þ þ þ þ þ þ þ þ þVenezuela þ þ þ þ þColombia þ þ þ þ þ þTrinidad þPeru þ þ þBrazil þ þ þ þ þBolivia þ

Asia þ þ þ þ þ þ þRussia þTadzhikistan þIndia þ þ þ þJapan þ þ þ

E.Nagm

etal./

CretaceousResearch

31(2010)

473e499

496


Cobban,1987; Küchler,1998), but differs in having finer ribs. The ribdensity and size of the present specimen can be considered asintermediate between E. saxonicum and E. matsumotoi.

Occurrence. According to Wright et al. (1996), the genus Eubos-trychoceras has been recorded from the TuronianeSantonian ofwestern Europe, northern Africa, Madagascar, southern India andJapan, as well as New Mexico and Texas (USA). The find of Eubos-trychoceras sp. from the Lower Turonian of the study area repre-sents the first record of the genus from Egypt.

4. Short palaeobiogeographic discussion

The wide distribution of early Late Cretaceous ammonites isbelieved to be a result of the major global transgression during thattime interval (Wiedmann, 1988; Lehmann and Herbig, 2009). Thegeographic distribution and taxonomic composition (predomi-nance of vascoceratids and pseudotissotiids) of the ammonitesrecorded (Table 1) confirm the strong affinity of the Wadi Arabafauna to the Mediterranean Province of the Tethyan Realm (warm-water “Vascoceratid Province” of Wiedmann, 1988). At the sametime, their distribution in different parts of the Tethys suggestsopen marine connections between Egypt and the other Tethyanregions during Late CenomanianeEarly Turonian times, related tothe general sea-level highstand during that interval (e.g.,Hardenbol et al., 1998; Sharland et al., 2001). To clarify the palae-obiogeographic affinities of the CenomanianeTuronian ammonoidsin the Egyptian Eastern Desert, their distribution is shown ona composite palaeogeographic map (Fig. 19), which is based on thereconstruction of Chancellor (1982) and Skelton (2003).

The connection between the Tethyan regions and West Africavia the Trans-Saharan epicontinental seaway appears clearly fromthis work (Table 1, Fig. 19) and has been documented by manyauthors before (e.g., Collignon and Lefranc, 1974; Reyment andChancellor, 1978; Meister et al., 1992, 1994; Kassab, 1994; Meister

Fig. 19. Palaeogeographic affinities of the ammonites recorded from the Wadi Araba area (thSkelton (2003) for the Late Cenomanian.

and Abdallah, 1996; Abdallah and Meister, 1997; Lehmann andHerbig, 2009). It is ascertained that this connection was estab-lished during the latest Cenomanian and the earliest Turonian (cf.Meister et al., 1992, 1994) and this intervals corresponds toa significant eustatic rise and maximum (Hardenbol et al., 1998;Sharland et al., 2001; Wilmsen, 2003). Interestingly, whenmoving westward into the western Tethys (Morocco), the typicalfauna of the Vascoceratid Province changes andmore cosmopolitanand temperate taxa are recorded (Wiedmann, 1988; Lehmann andHerbig, 2009). Whether or not this is related to the influence ofmore open-oceanic influences from the Central Atlantic needs to betested.

N. vibrayeanus is a typical Tethyan ammonite of wide distribu-tion, having been recorded from the Middle East, the SaharanPlatform, Portugal, Spain, and from Niger (see Wiese and Schulze,2005 for synopsis). The Late Cenomanian zonal marker M. gesli-nianum occurs very widely but not worldwide. It is knownthroughout the Boreal and Tethyan realms (Lehmann and Herbig,2009). Böse (1920) recognized that the North American fauna isconfined to a distinctive Mediterranean biofacies typified by vas-coceratid ammonites and the genus Pseudaspidoceras, which aredominant in the Turonian. He expected that this facies should bedistributed throughout northern and central Africa and intoPortugal. This view is supported here by abundant records of vas-coceratids in the ammonite collection studied. In addition, thevascoceratids (specifically V. durandi) are well represented in thesouthern Tethys from Algeria and Tunisia to Israel, and they alsooccur in the western Tethys (Lehmann and Herbig, 2009).Furthermore, V. cauvini is more widely distributed in Africa, witha few records from Europe (France) and South America (Peru).Bobkova and Luppov (1964) took the presence of tuberculate vas-coceratids in Turkestan as evidence of a connection with theMediterranean, via Iran and Syria, at peak transgression (see alsoWilmsen et al., 2005). Wiedmann (1964) and Collignon (1966) have

is study). Numbers refer to the taxa as listed in Table 1. Map after Chancellor (1982) and


stressed the links between Mexico and the Western Tethys. Theselinks relate especially to the genera Mammites, Pseudaspidoceras,Neoptychites, and Spathites.

Chancellor (1982, p. 122) pointed out that the vascoceratidsdescribed by Matsumoto and Muramoto (1978) from Japan seem torelate the Japanese fauna to Mexico, France, the Middle East, andIndia. In northwest Europe and North America, the most commonvascoceratids encountered are forms closely comparable to theMexican Fagesia. The genus Fagesia spread northwards though theWestern Interior of northern America and up the Pacific coast towhat is now almost the Canadian border. The genera Mammites,Pseudaspidoceras, Vascoceras, Neoptychites, Pseudotissotia, andWrightoceras all point to connections between Mexico and Nigeria,Niger, Gabon, Brazil, Angola and theCongo.All the above-mentionedgenera, with the exception of Neoptychites, are also known from thePacific coast of South America (Benavides-Cáceres, 1956; Etayo-Serna, 1979). This supports a direct connection between Mexicoand the Pacific via the Caribbean during the mid-Cretaceous(Chancellor, 1982). These genera, which are recorded also from thestudy area, show the open connection between the different placesmentioned above and Egypt during the early Late Cretaceous.

Acknowledgements

We thank J. Lehmann (Bremen) and C. Meister (Genève) forconstructive reviews as well as C.J. Wood (Minehead) for hiseditorial work. Many thanks also to Mrs. H. Schönig (Würzburg) forher excellent photographic work. A two-year financial support bythe Egyptian Missions Sector (High education ministry) to EN isgratefully acknowledged. This paper is a contribution to DFG(German Research Foundation) project WI 1743/6-1.

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