Taxonomy and morphological study on the vertebrate remains of Shark and rays fauna from the Middle and Late Eocene succession, Fayoum Depression, Egypt

Abstract

Well preserved vertebrate remains of Shark and ray fauna from the Middle and Late Eocene succession of Fayoum depression, at Gebal Qasr El-Sagha area have been documented and studied in details. Four stratigraphic sections are measured, described and sampled in the field at Hussein Wally Village, Birket Qarun, Qasr el-Sagha and Wadi el-Afreet. Lithostratigraphically, the studied succession is divided into three formations arranged from ba se to top into Gehannam, Birket Qarun and Qasr el-Sagha formations. This Eocene sequence provides by far the most complete view of the endemic African vertebrate fauna. Identification of the basic pattern of fish remains and taxonomic evaluation revealed that the recorded shark and rays taxa belong to 3 classes, 7 orders, 12 families, 18 genera, and 21 species. The identified taxa are macro-scale, collected on the surface, and known either from teeth or rostral remains. A taxonomic account and detailed morphologic description of the fossils shark, rays and bony fish teeth have been achieved. The depositional environments in the studied Middle-Late Eocene age sequence are interpreted. The abundance of recognized vertebrate fauna indicates environments varying from open marine shelf with low energy conditions to restricted marine shallow water conditions. However, the frequent distribution of macrofauna with intense bioturbation in sandstones of Birket Qarun Formation is a good indicator of restricted shallow water conditions.

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Copyright @ by Tanta University, Egypt
Delta J. Sci. 2017 Vol. 38 ; (202 -217 )
GE OL OGY
Taxonomy and morphological study on the vertebrate remains of Shark and rays
fauna from the Middle and Late Eocene succession, Fayoum Depression, Egypt
Abdelfattah A. Zalat, Hamza M. Khalil, Mohamed S. Fathy and Rana M. Tarek
Department of Geology, Fac. of Science, Tanta University, Tanta, Egypt
(E-mail: abzalat@science.tanta.edu.eg)
Abstract: Well preserved vertebrate remains of Shark and ray fauna from the Middle and Late Eocene succession of
Fayoum depression, at Gebal Qasr El-Sagha area have been documented and studied in details. Four stratigraphic sections
are measured, described and sampled in the field at Hussein Wally Village, Birket Qarun, Qasr el-Sagha and Wadi el-
Afreet. Lithostratigraphically, the studied succession is divided into three formations arranged from base to top into
Gehannam, Birket Qarun and Qasr el-Sagha formations. This Eocene sequence provides by far the most complete view of
the endemic African vertebrate fauna. Identification of the basic pattern of fish remains and taxonomic evaluation revealed
that the recorded shark and rays taxa belong to 3 classes, 7 orders, 12 families, 18 genera, and 21 species. The identified
taxa are macro-scale, collected on the surface, and known either from teeth or rostral remains. A taxonomic account and
detailed morphologic description of the fossils shark, rays and bony fish teeth have been achieved. The depositional
environments in the studied Middle-Late Eocene age sequence are interpreted. The abundance of recognized vertebrate
fauna indicates environments varying from open marine shelf with low energy conditions to restricted marine shallow
water conditions. However, the frequent distribution of macrofauna with intense bioturbation in sandstones of Birket Qarun
Formation is a good indicator of restricted shallow water conditions.
Key words: Taxonomy, Vertebrates remains, Eocene, Stratigraphy, Fayoum depression
Introduction:
The Fayoum depression was subjected to many
geological and archaeological studies over 200 years, as
it holds a rich heritage of best paleontological and
archeological sites in the world. Among these studies
(Said et al. 1972; Butzer 1976; Wendorf and Schild
1976; Kozlowski 1983; Hassan 1985, 1986, 1997; Said
1990; Hendrickx and Vermeersch 2000). The depression
hosts an unique heritage in terms of vertebrates and
mammals fossils. It has been known as a repository of
Paleogene vertebrates and has attracted the attention of
scientists worldwide (e.g., Kirk and Simons 2001;Simons
2005; Lewis and Simons 2007;Simons et al. 2007; Antar
2011; Adnet et al. 2011; Underwood et al. 2011) . The
Middle and Late Eocene represented an important period
in the evolution of sharks and rays and saw the
establishment of ‘modern-type’ trophic systems
(Underwood et al. 2011).
The Eocene rocks around and to the west of the
Fayoum depression comprise a thick succession of shelf
marine rocks representing environments from open shelf
to restricted lagoon. The stratigraphy of the area has been
documented by several authors (e.g. Beadnell 1905; Said
1962; Iskander 1943; Allam et al. 1991; Gingerich 1992;
Shama and Shided 1994; Abdullah et al. 1997; Dolsonet
al. 2002; Ismail and Abd El-Azeam 2008; Abu El Ghar
2012). However, a considerable lateral variation within
parts of the succession has caused problems with
applying a lithostratigraphical scheme (Strougo 2008).
The Fayoum depression has a fundamentally
important role to play in developing an understanding of
Egyptian prehistory, in particular during the
Epipalaeolithic – Neolithic time. This depression has
been inhabited by humans as early as 8500 BP. The first
significant occupation by humans, which is documented
archaeologically, occurred during the epipalaeolithic and
Neolithic Age, around 8200 BP. (Hassan 1986). During
this time clear traces of human impact on the Fayoum
depression could be observed in the old Fayoum lake
sediments (Zalat 2015).
Description of study area
The Fayoum depression occupies nearly triangular
depression, located between latitudes 29° 00’ – 29° 45’
N and longitudes 30° 00’ – 31° 10’ E, immediately to the
west of the Nile Valley and about 90 km to the south
west of Cairo (Fig.1).The depression covers an area of
approximately 1700 km2 and it lies below sea level. The
Fayoum is separated from the Nile Valley in the east by a

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ridge running south from the Giza plateau. The ridge is
approximately 8-10 km wide at its northern boundary,
but narrows to 2.5km in the south. Its surface is flat and
slopes downward northwesterly from 32 m above mean
sea level to 45 m below mean sea level at Qarun Lake,
which is the lowest part of the depression.The Fayoum
depression is carved out of the Middle- Upper Eocene
bedrock of the Western Desert of Egypt, and is
surrounded by escarpments from all sides . The Eocene
sequence is differentiated into four rock units including
Wadi El-Rayan, Gehannam, Birket Qarun and Qasr El-
Sagha Formations.
Material and Methods
Four stratigraphic sections are measured, described
and sampled in the studied area. The first section (I) is
located in south west Qarun Lake at Hussein Wally
Village, between Longitude29°27′ 56′′N and Latitude
30°23′36.8′′ E. The second section (II) is obtained from
the cliffs bordering the south western shore of lake
Qarun and far about 4Km east from the first section,
between longitude 29°27`47.1′′ N and Latitude
30°26`1.0′′ E. The third section (III) is measured at Qasr
El-Sagha Temple, between Longitude 29°36`3.2′′ N and
Latitude 30°40`13.9′′ E. The last section (IV) is located
far about 3.5 Km N-E Qasr El-Sagha Temple (Wadi El-
Afreet Section), between Longitude 29°37`10.3′′ N and
Latitude 30°41`54.8′′ E. A total of 130 samples are
collected from the studied sections and investigated in
details for paleontological and microfacies analyses.
Lithostratigraphy
The general lithostratigraphic units for Middle-
Upper Eocene sequences proposed by Beadnell (1905)
and modified later by Said (1962) are the most suitable
and will be followed herein. The Umm Rigl Member of
Gingerich (1992) is used herein to define the lowermost
strata of the Qasr El-Sagha Formation in the Hussein
WalleyVillage and Birket Qarun sections. Moreover, the
Temple and the Dir Abu Lifa members of Qasr El-Sagha
Formation suggested by Bown and Kraus (1988) in the
vicinity of Qasr El-Sagha Temple are adopted. Three
rock units are recognized as follows from the base to the
top; Gehannam, Birket Qarun and Qasr El-Sagha
Formations. The last formation is included Umm Rigl,
Temple and Dir Abu Lifa members.
Gehannam Formation
Author: Said (1962)
Type locality and Type section: The Garret
Gehannam, south west Fayoum Provence. It is contained
about 35m of gypsiferous claystone, marly limestone,
marly sandstone, glauconitic sandstone and marls (Said
1962).
Measured sections: The base of the formation is not
exposed, but it is likely that the lowest sample was from
a level quite high in the formation. About 15 to 20m of
this formation are exposed in the base of Hussein Walley
Village and Birket Qarun sections respectively.
Boundaries: In the west Fayoum area, the Gehannam
Formation conformably overlies El Gharaq Formation
and underlies the Birket Qarun Formation.
Description: The exposed section of upper part
Gehannam Formation in the west south Qarun Lake is
distinguished mainly by gypsiferous calcareous
claystones and shale with marl intercalations (Fig. 3).
The beds are bioturbated, sometimes glauconitic and
contain a diverse, open marine fauna. The upper part of
the Gehannam Formation passes upwardly into the
mudstones and sandstones of the Birket Qarun
Formation.
Birket Qarun Formation
Author: Beadnell (1905)
Type locality and type section: It is represented in
cliffs bordering the northern shore of lake Qarun (Said
1962), and the type-section is thus the steep-faced
precipitous escarpment described by Beadnell (1905).
Measured sections: This formation is well developed
in the two studied sections at south-western end of Lake
Qarun. Its thickness of about 38m at Hussein Wally
Village section and increase to about 52.5m at Birket
Qarun section (Fig. 3).
Boundaries: The Birket Qarun Formation conformably
overlies the Gehannam Formation by stratigraphic
gradual contact and underlies the Qasr El Sagha
Formation.
Description: In the studied sections, the Birket Qarun
Formation consists mainly of mudstone, siltstones and,
heavily bioturbated calcareous sandstones, and trace
fossil banks at its top part. The lower part of this
formation is arranged in coarsening-upward sequences
starting with claystones, siltstones and ending with fine
to medium sized sandstone. This sandstone characterizes
by golden yellow color, carbonate concretions,
calcareous, fossiliferous thin beds with trace fossils in
parts. The size and numbers of carbonate concretions
increase upwardly and to north direction in the studied
area. The upper part of the Formation consists mainly of
brownish yellow sandstone with some shale layer
intercalations. The shale change in thickness laterally
and characterizes by Thalassinoides trace fossils. On
other hand the top most sandstone bed at the contact with
superimposed Qasr El-Sagha Formation is highly
bioturbated by Rhizoliths trace fossils.

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Qasr El Sagha Formation
Author: Beadnell (1905)
Type locality and Type section: It is located at Qasr
El Sagha Temple with a thickness175 m of alternating
series of clays and limestone with sands and sandstone in
the upper beds.
Measured sections: The Qasr El Sagha Formation is
recorded in all studied sections and distinguished by
three members. Um Rigl Member, the lower part of the
Qasr El Sagha Formation attains 51m at Birket Qarun
section and 31m at Hussein Walley Village section. In
addition to the two upper members (Temple and Dir Abu
Lifa) are measured at Qasr El Sagha and Wadi El Afreet
sections by 88m and 95 respectively (Fig. 3).
Boundaries: In the studied area, the Qasr El Sagha
Formation conformably overlies the Birket Qarun
Formation and conformably and/or unconformably
underlies the Gabel Qatrani Formation.
Description and subdivisions: The Qasr El Sagha
Formation consists mainly of a thick succession of dark
gypsiferous mudstones with thin hash fossiliferous
reddish bed intercalations, and overlain by friable,
cleaned washed, cross bedded sandstones and ends with
an extensively hard yellow carbonate layer known as the
Bare Limestone (very hard dolomitic limestone).
Results and Systematic description
The recorded vertebrate remains from the studied Eocene
successions are investigated in detail. The results of
vertebrate identification show 21 species belong to 18
genera, 12 families, 7 orders and 3 classes. The
recognized fish assemblage has clear ecological affinities
with Eocene Tethyan fauna and also has common
elements with Eocene myliobatids, Carcharhinidae,
Triakidae and Hemigaleidaefishes. The most commmon
taxa are included Hexanchus agassizi Cappetta, Pristis
lathami Galeotti, Otodussokolovi (Jaekel),
Macrorhizoduspraecursor (Leriche), Cretolamna
twiggsensis (Case), Galeocerdo latidens (Agassiz),
Negaprion frequens (Dames)Rhinoptera sherburni
Arambourg, Carcharodon carcharias Linnaeus,
Galeocerdo cuvieriPéron & Lesueur 1822,
Xiphiorhynchus aegyptiacusWeiler, Rhynchobatus sp.,
Carcharias sp., Negaprion sp., Aetobatus sp.,
Galeocerdo sp., Rhizoprionodon sp., Leidybatis sp., and
Eutrichiurides sp.
The classification followed here after Compagno et al.
2005 Class Chondrichthyes Huxley 1880
Subclass Elasmobranchii Bonapart 1836
Cohort Euselachii Hay 1902
Subcohort Neoselachii Compango 1977
Super order Squalomorphii Compagno 1973
Order Hexanchiformes Buen 1926
Suborder Hexanchoidei Garman 1913
Family Hexanchidae Gray 1851
Genus Hexanchus Rafinesque1810
Hexanchus agassizi Cappetta 1976
Pl. 1, fig. 15.
2013 Hexanchus agassizi, Otero et al., Fig. 5: 1-6
2014 Hexanchus cf. agassizi, Carlsen & Cuny, Fig. 16
E–F
Description: The best preserved tooth measures 11 mm
mesio-distally, 6 mm apico-basally and 1.7 mm labio-
lingually. The tooth is worn. It is comb-shaped with a
main cusp and seven accessory cusps regularly
decreasing in size distally. All cusps are leaning
apicodistally. The mesial cutting edge of the main cusp
has no serration and no heel. It is convex in the basal two
thirds and straight near the apex. The distal cutting edge
is concave in the basal part and convex in the apical part.
The lingual face is convex, the labial face almost flat.
Geological distribution: The genus is known since the
Early Jurassic to Recent (Cappetta, 1987), with a
cosmopolitan distribution. Ward (1979) describes the
teeth of Hexanchus agassizi, from the Lower Eocene of
England. Hexanchus agassizi is known from south-
western France and from the London Clay of England
(Adnet 2006). Carlsen & Cuny Fig (2014) 16E–F (Early-
Middle Eocene Denmark).Ypresian of England, Eocene
of New Jersey, and the Oligocene of Australia and the
the Lower Oligocene of ex-U.S.S.R. (Cappetta, 1987).
Eocene Hexanchus has been described from Seymour
Island, Antarctica (Cione and Reguero, 1994).
Known geologic range: Cretaceous – Oligocene (Otero et
al. 2012)
Hexanchus sp.
2006 Hexanchus sp. Adnet, Fig.3
2006 Hexanchus griseus Adnet, Fig. 2 (A-L),
Remarks: Small teeth, with very distinct cutting edges
and wide crown. The recorded taxon is similar to
Hexanchus sp. and Hexanchus griseus (Bonnaterre,
1788) that reported by Adnet (2006) from late
Ypresian/early Lutetian, south−western France.
Order: Rajiformes Berg, 1940
Suborder Pristioidei CAPPETTA 1980
Family: Pristidae BONAPARTE 1838
Genus Pristis Latham, 1794
Pristis lathami Galeotti, 1837
Pl.1, fig. 18
2012 Pristis lathami Galeotti, Diedrich, Fig. 14 (18).
2012 Pristis lathami Galeotti, Zalmout et al., Fig. 5O-P
Description: The tooth is 45 mm long, 15 mm wide and 7
mm thick at the base of the crown. The proximal half of
the tooth is thickest, and the tooth tapers distally to a
much thinner 3 mm. The anterior edge of the tooth is
thick and blunt, while the posterior edge is concave, a
shallow gutter that runs from the base of the tooth to the

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distal end. The vertebrae is rounded and has concentric
lines in internal view and longitudinal lines in side view.
Geographic distribution: Pristis lathami is known from
lower and upper Eocene deposits in Africa and North
America (Cappetta, 1987). Eocene of the Paris Basin
(Casier, 1949). In Egypt the species is known from the
Eocene of the Fayoum and Bahariya regions (Stromer,
1905B; Case and Cappetta, 1990; Underwood et al.,
2011; Murray et al., 2011; Adnet et al., 2011). Late
Eocene Qattara depression, Egypt (Zalmout et al. 2012),
late Eocene of South Carolina and North America
(Cicimurri 2007), Southern North Sea (Diedrich 2012).
Farther east, Pristis lathami is known from the Eocene of
Qatar (Casier, 1971). Middle–Late Eocene, southwestern
Morocco (Adnet et al. 2010)
Known Geologic range: Early to Late Eocene.
Pristis aquitinicus Delfortrie 1872
1959 Pristis aquitinicus, Delfortrie, Ghosh , p. 67, pl. 88,
Figs. 7, 8.
2013 Pristis aquitinicus Delfortrie, Sharma and Patnaik,
Fig. 5: J, K.
Description: The anterior surface is tapering gently
towards the tip thereby increasing the curvature while the
posterior surface is straight and has a broad sulcus.
Length of about 22 mm and width 21 mm.
Geographic distribution: The fossil remains of the genus
Pristis ranging in age from Lower Eocene to Recent
(Sharma and Patnaik2013). It is reported from Miocene
of India (Sharma and Patnaik2013). Cappetta (1970) also
reported P. aquitanicus from the Middle Miocene of
Southern France
Known Geologic range: Early to Late Eocene.
Suborder Rhynchobatoidei Flower 1941
Family: Rhynchobatidae Garman, 1913
Genus: Rhynchobatus Müller and Henle, 1837
Rhynchobatus sp.
2010 Rhynchobatus sp.Adnet et al., Fig. 4 E.
2011 Rhynchobatus sp.Underwood et al., Fig. 7 (X).
2011 Rhynchobatus sp.Antar Fig. 29 (O)
2013 Rhynchobatus sp.Sharma and Patnaik, Fig. 5: F.
Description: A small tooth; crown enamel is having a
granular texture, the lingual face is flat, concave on each
side separated by a moderate medial uvula, the terminal
end of the median uvula is pointed, lateral uvulae absent;
root massive and extends beyond the lingual face of the
crown; root lobes divided on both the sides by a deep
nutritive groove and a small foramina.
Geographic distribution: It is recorded from Middle to
Late Eocene of the Fayoum area (Underwood et al.,
2011), Late Eocene of the western Desert, Egypt (Adnet
et la., 2010), Langhian of Loupian, Herault, Southern
France (Cappetta, 1987) and from Oligocene Chandler
Bridge Formation of South Carolina, U.S. (Cicimurri and
Knight, 2009). Rhyncobatus teeth have also been
reported from the Maastrichtian of Morocco
(Arambourg, 1952), the Miocene of Japan (Itoigawa et
al., 1985) and Miocene of India (Sharma and Patnaik,
2013).
Known geologic range: Maastrichtian to Miocene.
Suborder Myliobatoidei Stingrays
Super family Myliobatoidea Compango 1973
Family: Myliobatoidea Bonaparte 1838
Genus: Myliobatis Cuvier 1817
Myliobatis sp.
1972 Myliobatis sp., Welton, Plate 1, p. 167, Fig. 8.
2011 Myliobatis sp., Antar, Fig 29 J-J1
2011 Myliobatis sp., Underwood et al., Fig. 4 (G).
2012 Myliobatis sp. Otero et al ., Fig. 3: T
2013 Myliobatis sp., Sharma and Patnaik, Fig. 3(A).
Description: Teeth broader than length, with hexagonal
contour in shape and rectilinear outline; the length is
about 28 mm and the width of about 7-9 mm, the crown
is as thick as the root, flat and quite convex, being
displaced anteriorly with respect to the root. The crown
and the root are well separated by a minor groove. The
basal surface of the root is flat with 23 root lobes
separated by grooves. The tooth is much broader than
long.
Geographic distribution: The genus Myliobatis is known
since the Early Paleocene to Recent, with a cosmopolitan
distribution (Cappetta, 1987). Myliobatis sp. has been
described from the Paleocene to Eocene of western
Africa including Angola, Egypt, Morocco, Togo and
Nigeria (Cappetta 1987; Cook et al. 2010). Middle
Eocene of Kutch (Mishra, 1980), Miocene Coaledo
Formation along the Oregon coast Welton (1972),
Miocene of Baripada Beds, Orissa (Sahni and
Mehrotra,1981), Eocene of Subathu Formation, Bilaspur
area and Himachal Pradesh (Singh, 1985, Kumar and
Loyal, 1987), Eocene Vastan lignite Mine of Cambay
Shale, Gujarat (Rana et al., 2004), Lower Eocene
Panandro lignite field, Gujarat (Bajpai et al., 2002),
Early Eocene Kapurdi Formation of Rajasthan (Rana et
al., 2006), Eocene deposits of Fayoum and western
Desert, Egypt (Case and Cappetta, 1990; Antar, 2011;
Underwood et al., 2011),Miocene of India (Sharma and
Patnaik, 2013). The genus also has been recognized on
Seymour Island, Antarctica (Kriwet, 2005).
Known geologic range: Campanian to Recent.
Genus: AetobatusBlainville1816
Aetobatus sp.
2011 Aetobatus sp. Antar, Fig. 29 (M).as
2011 Aetobatus sp. Underwood Fig. 4 (F)
2013Aetobatus sp. Sharma and Patnaik Fig. 4: E, F, G

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Description: One sample is collected and well preserved.
Root is quite distinctive with small vertical lines along it
and has smooth edges. The crown lacks the hexagonal
shape which is usually associated with myliobatids. The
contact between root and crown is straight. Crown is
slightly serrate and less thickness than root. The occlusal
surface is smooth. The root is nearly as high as the crown
and is divided into longitudinal ridges and grooves. The
length is about 6mm and width of about 6mm.
Geographic distribution: Aetobatus teeth have been
described from the Lower Miocene of the western coast
of India (Sahni and Mishra, 1975).
Known geologic range: Middle Eocene to Lower
Miocene
Genus: Leidybatis Cappetta, 1986
Leidybatis sp.
Pl.1, fig.17
Description: Teeth broad with rectangular shape. The
length is about 28-32 mm and the width of about 7-9
mm, Root is thick and distinctive with small serration
along it and has smooth edges. The contact between root
and crown is straight. Crown is slightly serrate and less
thickness than root. The root is nearly as high as the
crown and is divided into longitudinal ridges and
grooves.
Known geologic range: Middle to Late Eocene
Order: Myliobatiformes Compagno, 1973
Family: Rhinopteridae Jordan and Evermann, 1896
Genus: Rhinoptera Cuvier, 1829
Type species: Myliobatis marginata Saint−Hillaire,
1817, Recent, Mediterranean Sea.
Remarks: The Rhinopteridae are comprised of a single
genus, Rhinoptera (Cappetta 2004)..
Geographic distribution: The genus is known from the
Paleocene in North and West Africa, and Europe
(Cappetta 2004), and Lower Miocene of Switzerland and
France and most other marine Neogene deposits (Leriche
1927; Cappetta 1970).
Rhinoptera sherburni Arambourg 1952
Pl.1, fig.16
1952 Rhinoptera aff.sherburniArambourg, Plate XXXII,
Figs. 15-24.
2013 Rhinoptera sherburni Arambourg, Sharma and
Patnaik(Fig.4: H, I)
Description: The teeth are medium size, with 8-10mm
length, and 4-6 mm width, hexagonal in shape; teeth
possess a prominent but thin lingual shelve; labial face is
more upright, the lingual root overhangs the crown by a
distinct margin, the crown is thick. Upper surface of root
is slightly curved and smooth. The contact between root
and crown is very slightly curved. The crown is serrate
and curved in middle area and become straight towards
edges.
Geographic distribution: Rhinoptera sherborni has been
reported from the early Eocene of Virginia (Kent, 1999);
the Middle Eocene of England (Kemp et al., 1990),
Eocene Nigeria (White, 1926), Morocco (Arambourg,
1952) and Uzbekistan (as Rhinoptera cf. sherborni, Case
et al., 1996) and the late Eocene of Egypt (Murray et al.,
2011). Fossil record of Rhinoptera extends back upto the
Late Palaeocene (Cappetta, 1987, 2006), Middle–Late
Eocene, southwestern Morocco (Adnet et al. 2010),
Miocene of India (Sharma and Patnaik2013),
Madagascar (Wallett, 2006).
Known geologic range: Paleocene to Miocene
Order Lamniformes BERG 1958
Family: Odontaspididae Müller and Henle, 1839
Genus: Carcharias Rafinesque, 1810
Carcharias sp.
2012Carcharias sp.Zalmout et al. Fig. 3B-E
2012 Carcharias sp. Diedrich Fig. 14 (7-8)
2013Carcharias sp. Otero et al., Fig. 3: 1-4
Description: The recorded taxon is small-sized not larger
than 2 cm, with length 15-18 mm, and width 5-8mm. The
crown sharp having smooth enamel in both labial and
lingual surfaces; the cutting edge is complete and reaches
the base of the crown; slender root branches with medial
groove and deep depression between them; In some
specimens the root of tooth is partially broken and
arched. The contact between root and crown is arched.
Crown is thin and long and has smooth edges and surface
in lingual and labial view are ended with very acute end.
Geographic distribution: Fossils of Carchariashave been
found all over the world, especially in the Cretaceous,
Eocene, Miocene and Oligocene sediments of Europe,
United States, Australia and Africa (Cappetta and Case,
1975a,b, Cappetta, 1987, Otero et al., 2013, Zalmout et
al., 2012, Diedrich 2012).Southern North Sea (Diedrich
2012),Late Eocene Qasr El Sgha, Fayoum (Underwood
et al. 2011), Late Eocene of Qattara depression, Egypt
(Zalmout et al. 2012), Pliocene Farol das Lagostas
locality, Angola (Antunes 1978).
Known geologic range: Maastrichtian to Pliocene.
Family Otodontidae Gückman 1964
Genus Otodus Jordan and Hannibal, 1923
Otodus sokolovi (Jaekel, 1895)
Pl.1, figs.1-4
2010 Carcharocles sokolovi Jaekel, Adnet et al. Fig.3
a1,2
2011 Carcharocles sokolowi Jaekel, Antar, Fig.24
2011 Otodus (Carcharocles) sokolowi Jaekel,
Underwood et al. Fig.4 A
2012 Otodus cf. sokolovi Jaekel, Zalmout et al. 2012,
Fig. 3F-BB
2012Carcharocles sokoloviJaekel, Diedrich, Fig.10 (9-
10)
Description: Large teeth that may reach 60 to 80 mm in
height and 45-65 mm in width. Central blades are large

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and bear fine and regular strong serrations along both
cutting edges. The teeth have two divergent lateral cusps
which are triangular in shape, serrated and wide.
Geographic Distribution: This species is recorded from
middle Eocene of Syria, Jordan (Cappettta et al., 2000,
Mustafa and Zalmout, 2002; Smadi et al., 2003; Mustafa
et al., 2005), middle Eocene of Fayoum, Egypt (Stromer,
1905A; Case and Cappetta, 1990; Vliet and Abu el
Khair, 2010; Underwood et al., 2011), Middle–Late
Eocene, southwestern Morocco (Adnet et al., 2010), and
Nigeria, Congo, and Angola (White, 1926; Dartevelle
and Casier, 1942, 1943, 1949, 1959),Late Eocene Qattara
depression (Zalmout et al. 2012), In North America
Otodus (Carcharocles) is known from North Carolina
(Bourdon and Chandler, 2007), the Barnwell Formation
in Georgia (Case and Borodin, 2000), and middle Eocene
of Alabama (White, 1956), middle and late Eocene
Tethyan deposits in North Africa, the Middle East, and
North America (Zalmout et al. 2012).
Known geologic range: Middle Eocene - Lower
Oligocene.
Family: Lamnidae Müller & Henle 1838
Genus Carcharodon Müller & Henle, 1838
Carcharodon carcharias Linnaeus, 1758
Pl.1, fig. 9
1973 Carcharodon carcharias Linnaeus, Mehrotra et al.,
p. 191-192, pl. 2, figs. 2, 6 a-b.
1981 Carcharodon carcharias Linnaeus, Sahni and
Mehrotra, p. 112-113.
1998 Carcharodon carcharias Linnaeus, Tiwari et al., p.
12, pl. 1, fig. 5, 6.
2009 Carcharodon carcharias Linnaeus, Mondal et al.,
pl. II, figs. 1-4.
2010 Carcharodon carcharias Linnaeus, Andreev and
Motchurova, Fig. 2 a, b
Description: The tooth is having height equal or greater
than the width. Length 22 mm and width 16 mm. Tooth
composed of root and crown. Root is broken except
small part and appeared slightly curved and crown is
broad in upper part and become narrow towards lower
part and has sharp end and serrate edges and has
longitudinal lines in ligual and labial view. Serrations of
the cutting edges are uniform, and relatively coarser
towards the base in larger specimens. Mesial edge is
more concave than distal edge, mesial concavity shows
wide variations, some are feebly concave or straight.
Labial surface is flat, however at the tip it may be
slightly inwardly curved. The base of the root may be
feebly bifurcated or flat.
Geographic distribution: This species is recorded from
Farol das Lagostas, Angola. (Andreev and Motchurova
2010), Miocene Chesapeake Bay at Calvert Cliffs in
Maryland. USA, Lowery et al. (2011),Early Pliocene of
Spain, Early – Middle Miocene of Greens mill run, north
carollina, western and Eastern USA, South Africa,
Australia and Late Eocene Qasr El-Sagha Formation
Egypt (Zalmout et al. 2012).
Known geologic Range: Eocene to Early Pliocene.
Genus Macrorhizodus Glikman, 1964
Type species: Isurus praecursor (Leriche, 1905).
Macrorhizodus praecursor (Leriche, 1905)
Pl.1, fig.5
2011 Cosmopolitodus praecursor (Macrorhizodus
praecursor ), Leriche, Antar, Fig.25: A-B7.
2012 Macrorhizodus praecursor Leriche, Zalmout et al.,
Fig. 4A-V
2012 Isurus praecursor Leriche,Diedrich, Fig.11 (1-10).
2012Macrorhizodus (Isurus) praecursor Leriche,Otero et
al., Fig. 3: L, M
2013 Macrorhizodus praecursor Leriche, Otero et al.,
Fig. 3: 28-34
Description: Medium to relatively large sized lamniform
teeth with high, slender and triangular crown that
becomes broader toward the base, being labio-lingually
compressed; flat labial face with soft enamel and smooth
folds near the root; lingual face is convex with some
cracks in the enamel; root is slightly massive, basally
flattened. The teeth may reach 40-44 mm in total height
anteriorly, and 30-32 mm in total width laterally.
Geographic distribution: Macrorhizodus praecursor is
widely known from all middle and late Eocene Tethyan
deposits of Europe, North America and some late Eocene
marine vertebrate localities in Egypt (Adnet et al., 2011),
Middle to Late Eocene of the Seymour Island in
Antarctica (Cione and Reguero, 1994), Priabonian of the
southernmost Chile (Otero et al., 2012b). Eocene
deposits in Belgium, Syria, Nigeria, Togo, Guinea Bissau
and England (Cappetta, 1987). Middle-Late Eocene of
Chesapeake Bay, U.S. (Kent, 1994),southwestern
Morocco (Adnet et al. 2010) and Southern North Sea
(Diedrich (2012).Late Eocene of Qattara depression,
Egypt (Zalmout et al., 2012).
Known Geologic range: Middle - late Eocene
Family: Cretoxyrhinidae Gluckman 1958
Genus: Cretolamna GLUCKMAN 1958
Cretolamna twiggsensis Case, 1981
Pl.1, figs.11-12
1981 Lamna twiggsensis Case, pl. 3, Figs. 3–8
1990 Cretolamna twiggsensis Case, Case & Cappetta, pl.
3, Figs. 40–55
2007 Cretolamna twiggsensis Case, Adnet et al., Figs.
6.21 and 6.22
2011 Brachycarcharias twiggsensis Case, Underwood et
al., Fig. 4L–M
2011 Cretolamna twiggsensis Case, Adnet et al, Fig. 3A
2012 Brachycarcharias twiggsensis Case, Zalmout et al.,
Fig. 4W-X

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Description: Teeth are relatively large; with height up to
20-25mm and 15-18 mm width, the cusp is high, quite
triangular in upper teeth and not very thick. The root
lobes are elongated and their ends are often rounded. The
upper anterior tooth has two divergent lobes with a
height less than the crown, the anterior –lateral teeth
have vertical central blade, slightly sigmoidal in shape
and had a wide base, with pointed edge and having two
small diverted lateral cusps.
Geographic distribution: the genus is nearly worldwide,
Early Cretaceous-Early Eocene. African distribution and
occurrence: Morocco, Angola, Democratic Republic of
the Congo, Niger, Nigeria and Egypt; Late Cretaceous-
Eocene (Cappetta 1987). The species extends to
palaeotropical seas between the Caribbean, western
Tethys (Case, 1981; Case & Borodin, 2000) and oriental
Tethys (Casier, 1971; Case & Cappetta, 1990; Case &
West, 1991; Adnet et al. 2007). It is recorded from Egypt
(Mokkatam to Wadi Hitan areas), both in Bartonian and
Priabonian deposits (Case and Cappetta, 1990 and
Underwood et al., 2011), middle and late Eocene of the
Fayoum and Qattara depression, Western Desert of
Egypt (Case and Cappetta 1990; Strougo et al., 2007;
Underwood et al., 2011; Adnet et al., 2011; Zalmout et
al. 2012), the species is recorded elsewhere from
Pakistan (Adnet et al., 2007), Middle–Late Eocene of
southwestern Morocco (Adnet et al., 2010) and Georgia,
USA (Case, 1981; Case and Borodin, 2000).
Known geologic range: Middle–Late Eocene
Order Carcharhiniformes
Family:Carcharhinidae Jordan & Evermann, 1896
Genus Galeocerdo Müller and Henle, 1837
Type species: Galeocerdo cuvieri Peron and Le Sueur,
1822
Galeocerdo cuvieriPéron & Lesueur, 1822
Pl.1, fig. 14
1973 Galeocerdo cuvieri Péron & Lesueur, Mehrotra et
al., p. 184, pl. 1, fig. 3.
1981 Galaeocerdo cuvieri Péron & Lesueur, Sahni and
Mehrotra, p. 109, pl.2, fig. 12.
2009 Galeocerdo cuvieri Péron & Lesueur, Mondal et
al., Pl.I, figs. 11-12.
Description: Tooth large with 23-30 mm long and 18-21
mm width, sub-triangular, irregular in shape, broader
than high and highly oblique. Crown is labio-lingually
thickened; the mesial cutting edge has a deep notch,
distal margin strongly convex; both cutting margins are
serrated; strength of serrations becomes weak to obsolete
near the apex; at distal cutting edge strength of serrations
are coarser at the middle, decreases towards the base and
the apex, whereas mesial edge serrations are finer and
uniform. Mesial heel is extended, with numerous
denticles, size of the denticles increases proximally. On
the labial surface a prominent triangular pit is present at
the middle near the root; labial surface bears few
longitudinal striations; crown-root boundaries at both
faces are convex, convexity more pronounced in lingual
side. Root higher than crown; in cross -section it is labio-
lingually arched and thicker at the middle part; in profile
the tooth deflects outward.
Geographic distribution: G. cuvieri has been reported
from the lower Miocene beds of different localities in
Gujarat (Mehrotra et al. 1973). Miocene from baripada,
Orissa (Mondal et al 2009). The type species,
Galeocerdo cuvier(Peron and Le Sueur 1822), is extant
and can be found in all tropical and temperate seas,
including those of Madagascar (Smale 1998; Cappetta
2004). Pliocene of Italy (Lawley 1876), South Africa
(Davies 1964), and North Carolina (Cappetta 2004).
Known geologic range: Middle Eocene to Pliocene.
Galeocerdo sp.
1990 Galeocerdo sp. Case and Cappetta, 1990, plate 5,
figs 92-95.
2010 Galeocerdo sp. Andreev and Motchurova, fig. 5 a-
c; fig. 6
2002 Galeocerdo sp. Bajpai and Thewissen, txt-fig.2f.
Description: Root of the tooth is arched and elongate and
has linear groove in mid line in lingual view and smooth
in labial view. The contact between root and crown is
slightly arched.
Geographic distribution: Late Eocene of El-Sagha
Formation (Case and Cappetta, 1990). Teeth of
Galeocerdo have previously been reported in Egypt by
Stromer (1905a) and Priem (1897b). Eocene of Minqar
Tabaghbagh, western Desert, Egypt (Vliet and Abu El-
Khair 2010). This species is recorded from Upper
Miocene of Angola. . (Andreev and Motchurova2010).
Known Geologic range: Eocene to Upper Miocene.
Genus Carcharhinus Blainville, 1816
Type Species: Carcharias melanopterusQuoy and
Gaimard, 1824
Carcharhinus sp.
2010 Carcharhinus sp. AndreevP, Motchurova, Fig. 5 c,
d
2010 Carcharhinus sp. Adnet et al. Fig.3 G.
2011 Carcharhinussp. 1, Antar, Fig. 28 C
2012 Carcharhinus sp1.Zalmout et al., Fig. 5E-F.
Description: The teeth are relatively small up to 11 mm
height and 11-12mm width. Cusp is flattened labially and
triangular in shape , extends mesiodistally into shoulders;
the shoulders are sharp and separated from the main
blade by slightly developed notches. Lingually, it is
convex and the apex curved labially and has no lateral
cusplets. The roots have a shallow furrow in the middle
of the lingual portion.
Geographic distribution: Middle–Late Eocene,
southwestern Morocco (Adnet et al. 2010), Middle and

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Late Eocene deposits of Fayoum and Qattara
depressions, Egypt (Underwood et al. 2011, Zalmout et
al., 2012)
Known Geologic range: Middle and Late Eocene.
Genus Negaprion Whitley, 1940
Negaprion frequens (Dames, 1883)
Pl.1, fig.10
1908 Carcharias (Aprionodon) aff. frequens Priem,
pl.15, Fig. 6–7
1971 Aprionodon frequens Casier, pl. 1, Fig. 6
1990 Carcharhinus frequens Dames, Case & Cappetta,
pl. 5, Figs. 102– 107; pl.7, Figs. 143–148 and 151–159
1990 Negaprion frequens Dames, Case and Cappetta,
Plate 7, 147–147.
2011 Carcharhinus aff. frequens Dames, Adnet et al.,
Fig. 3G–H
2011 Negaprion frequens Dames, Underwood et al., Fig.
5V
2012 Negaprion frequens, Dames, Zalmout et al., Fig. 5,
A-D
Description: Teeth of are relatively small, ranging from 7
to 12 mm in total height, from 5 to 12 mm in total width
(mesiodistally), and from 2 to 3 mm in lingolabial
thickness in the middle of the root. The upper teeth have
a crown of triangular outline with a cusp slightly slanted
distally, while the lower teeth generally have a slender
cusp with smooth cutting edges than never reach the
heels, except on some lateral teeth. The cutting edges are
always unserrated and are not disconnected from the
cutting edges of lateral heels which are totally smooth.
The root lobes have enlarged and rounded ends.
Geographic distribution: It is recorded from the Miocene
of Europe (Leriche 1926; Leriche 1957; Antunes and
Jonet 1970; Cappetta 1970), North Africa (Arambourg
1952), North and South America (Longbottom 1979),
and India (Mehrotra et al 1973; Sahni and Mehrotra
1981), Angola (Antunes 1978), middle Eocene of the
Midra and Saila shales of Qatar (Casier, 1971), Late
Eocene of Birket Qarun and Qasr el-Sagha formations,
Fayoum area (Case and Cappetta, 1990; Underwood et
al., 2011), late Eocene of Jordan (Cappetta et al., 2000;
Mustafa and Zalmout, 2002), and the late middle Eocene
to late Eocene of southwestern Morocco (Adnet et al.,
2010), Miocene fishes from baripada beds, Orissa
(Patnaik et al., 2014).
Known geologic range: Eocene to Pleistocene.
Genus: Rhizoprionodon Whitley 1929
Type species: Carcharias (Scoliodon) crenidens
Klunzinger, 1880.
Rhizoprionodon sp.
Pl.1, figs. 7-8
1990 Rhizoprionodon sp., Case& Cappetta, pl.7. Figs.
160–163
1991 Rhizoprionodon sp., Case & West, pl. 3. Figs. 2–4
2011 Rhizoprionodon sp., Adnet et al., Fig. 3P–Q
2012 Rhizoprionodon sp., Zalmout et al., Fig. 5K
2013 Rhizoprionodon sp., Otero et al., Fig. 6: 3-8
2014 Rhizoprionodon sp., Sharma and Patnaik, Pl. 4,
figs. 9, 10
Description:Teeth are small, up to 7-8mm height and 8-
10mm width, and 2-3mm thick, cusp small and is bent
backward with extended crown at the base. Both the
cutting edges are sharp, smooth without any serration.
The mesial cutting edge is long slightly concave and
recurved towards the apex and the distal edge is short;
the mesial cutting edge is sigmoidal. The lingual face is
convex and the labial face is flat. The distal heels are
rounded with cusplet. The crown overhangs the root. The
root low and broad, median nutritive groove is present in
the lingual face of the root. The basal margin of the root
concave to straight.
Geographic distribution: This genus is relatively
common in worldwide Eocene marine deposits, Fayoum
and Qattara depression, Egypt (Case and Cappetta, 1990;
Strougo et al., 2007; Underwood et al., 2011), from
Pakistan (Case and West, 1991; Adnet et al., 2007),
Middle to Late Eocene. Río Baguales Formation (Otero
et al. 2012).Rhizoprionodon sp. is recorded from the
Eastern Coast of India (Patnaik et al., 2014).
Known geologic range: Middle Eocene to Miocene.
Class: Osteichthyes Huxley, 1880
Order: Perciformes Bleeker, 1859
Family: Trichiuridae Rafinesque, 1810
Genus: Eutrichiurides Casier 1944
Eutrichiurides sp.
Pl.1, fig.13
1966 Eutrichiurides sp. pl.3, fig. 30
2004 Eutrichiurides sp. Rana et al., Fig. 3 (41)
Description: Teeth long and slender with conical shape;
height up to 33 mm and width of about 8 mm. Apex
small chisel-like, about one-tenth of basal part; slight
keel along the edges extend towards base; basal part
laterally compressed with vertical ridges and grooves. It
has irregular surface and longitudinal lines on lingual
and dorsal view.
Geographic distribution:Eutrichiurides is an extinct
genus of prehistoric bony fish. This genus is recorded
from the Cretaceous age at the Khouribga Plateau,
Morocco, the Early-Mid Eocene, Monmouth County, the
Early Eocene (Ypresian) of the lower part of the
Jaisalmer basin (Cappetta 1987).
Known geologic range: Cretaceous -Eocene
Class Actinopterygii (sensu Nelson, 1994)
Division Teleostei Muller, 1844
Order Perciformes Bleeker, 1859
Suborder Xiphioidei Swainson, 1839

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Family Xiphiidae Swainson, 1839
Subfamily Xiphiorhynchinae Regan, 1909
Genus Xiphiorhynchus van Beneden, 1871
Type Species-Xiphiorhynchus elegans van Beneden,
1871
Xiphiorhynchus aegyptiacusWeiler, 1929
1929 Xiphiorhynchus aegyptiacus.Weiler, Taf. I , Fig. 4.
Description: Elongate vertebrae, thin thickness in the
middle that become broad towards lower and upper
edges and has knobs in labial view. Height of about 12-
15 mm and width 8-13 mm.In cross section, the rostrum
contains two types of longitudinal canals, an unpaired
central canal and two pairs of lateral nutrient canals, and
both types vary as to how far they extend distally. The
dorsal pair of lateral nutrient canals is positioned closer
to the mid-line than the ventral pair of lateral nutrient
canals.
Known Geologic range: Eocene
Summary and conclusion
The studied Eocene succession exposed at the
northern part of Lake Qarun is differentiated into three
rock units arranged from base to top: Gehannam, Birket
Qarun and Qasr El-Sagha formations. Qasr El-Sagha
Formation is distinguished into three members: Um
Reigl, Temple and Abu Lifa members. Paleontologically,
the vertebrate faunal content of the studied sections is
investigated. A total of 21 sharks species are recorded
and suggest that they occupied a wide range of
ecological niches. Some of the recorded species are
limited in their stratigraphical range and show potential
to be used, as biostratigraphical indicators through the
Eocene sedimentary sequences. The upper part of
Gehannam Formation yields vertebrate remains such as
the Shark and ray teeth, marine mammal skeletons.
Vertebrate remains of great white shark Macrorhizodus
praecursor (Leriche), Otodus sokolovi, Pristis
lathami,Pristis aquitinicus, andCarcharias sp., in
addition to some traces fossils, Thalassinoides and
Rhizolithsare common in Birket Qarun and Qasr El-
Sagha Formations. The marked variations in abundance
of the recognized vertebrate faunas reflect distinct
environmental conditions that control the distribution of
many species. The water depth is considered one of the
important ecological factors beside the eutrophic state of
the water.
The depositional environments in the studied area
during Middle-Late Eocene age are interpreted. The
abundance of the vertebrate faunas indicates
palaeoenvironments varying from open marine shelf with
low energy conditions to restricted marine shallow water
conditions. However, the abundance of microfauna
within calcareous shale and malrs of Gehannam
Formation indicate an open shallow marine shelf with
low energy conditions. The frequent distribution of
macrofauna with shallow water shark taxa and intense
bioturbation in sandstones of Birket Qarun Formation is
a good indicator of restricted shallow water conditions.
Horizontal laminations consisting of alternating shale,
siltstone and sandstone with dolostones sheets that show
cyclic changes in layer thickness of Temple Member
resemble tide-dominated estuaries environment, which
cut off by incised valley fill heterolithic sediments of
Abu Lifa Member.
References
Abdallah, A., Abuol Ela, N., Shama, K. and Abdel Aziz,
S.(1997). Biostratigraphy of the Temple Member
(Qasr El-Sasgha Formation, Late Eocene) north El
Fayum depression: Egypt. J. Geol. 41(1): 541-578.
Abu El Ghar, M. S. (2012). Sequence stratigraphy and
cyclicity in the Middle Eocene of the Fayoum
ranges, Western Desert, Egypt: Implications for
regional sea level changes. Marine and Petroleum
Geology, 29 (1): 276-292.
Adnet, S. (2006).Nouvelles faunes de sélaciens
(Elasmobranchii, Neoselachii) de l'Éocène des
Landes (Sud-Ouest, France).Implication dans les
connaissances des communautés d'eaux
profondes.Palaeo Ichthyologica, 10: 1-128
Adnet, S., Cappetta, H. and Nakaya, K. (2006).Dentition
of etmopterid shark Miroscyllium (Squaliformes)
with comments on the fossil record of
lanternsharks. Cybium, 30 (4): 305-312.
Adnet, S., Antoine, P.O., Hassan Baqri, S.R., Crochet J.-
Y., Marivaux L.,JL. Welcomme J-L., and Métais,
G. (2007). New tropical carcharhinids
(chondrichthyes, Carcharhiniformes) from the late
Eocene–early Oligocene of Balochistan, Pakistan:
Paleoenvironmental and paleogeographic
implications Journal of Asian Earth Sciences 30:
303–323.
Adnet, S., Cappetta, H. and Tabuce, R. (2010). A
middle-late Eocene vertebrate fauna (marine fish
and mammals) from southwestern Morocco;
preliminary report: age and palaeobiogeographical
implications. Geological Magazine, 147: 860-870.
Adnet, S., Cappetta, H., Elnahas, S. and Strougo, A.
(2011). A new Priabonian Chondrichthyans
assemblage from the Western desert, Egypt:
Correlation with the Fayum Oasis. Journal of
African Earth Sciences, 61: 27–37.
Allam, A., Shama, K. and Zalat, A.
(1991).Biostratigraphy of the Middle Eocene
succession at Mishgigah, Wadi El Rayan, Libian
Desert, Egypt. Journal of the African Earth
Sciences; 12(3): 449-9.
Andreev, P. and Motchurova, N. (2010).Fossil
Elasmobranchii and Actinopteryg II. Bulletin of the
Natural History Museum, 3: 115-129

Zalat et al.
Taxonom y and morphol ogical study on the vertebrate remains o f Shark and rays f auna from the Middle and Late Eocene success ion, Fayoum Depress ion, Egypt

211
Antar, M.S.M. (2011). Paleo-environments of the
exposed Eocene Sediments between Wadi El-Hitan
and east Siwa in the Egyptian Western Desert based
on their faunal content especially the vertebrates.
Ph.D Thesis, Zagazig University, Faculty of
Science, Geology Department, 257 pp.
Antunes, M.T.(1978). Faunes ichthyologiques du
Neogene superieur d’Angola, leur age, remarques
sur Ie Pliocene marin en Afrique australe. Ciencias
da Terra (UNL), Lisboa, 4: 59-90.
Antunes, M.T. and Jonet, S. (1970). Requins de l’
Helvétien supérieur du Tortonien de
Lisbonne.Revista da Faculdade deCiências da
Universidade de Lisboa 2, C, 16(1): 119-280.
Arambourg, C. (1952). Les Vertébrés fossiles des
gisements de phosphates (Maroc, Algérie, Tunisie).
Service Géologique du Maroc, Notes et Mémories
92: 1-372.
Bajpai, S. and Thewissen, J.G.M. (2002).Vertebrate
fauna from Panandhro lignite field (Lower Eocene),
District Kachchh, western India.Curr. Sci., v. 82
(5): 507–509.
Beadnell, H.J.L.(1905).The topography and geology of
the Fayum Province of Egypt.Survey Department
of Egypt, Cairo, 101 pp.
Bourdon, J. and Chandler,R. (2007).The Castle Hayne
Formation. Middle Eocene Sharks and Rays of
North Carolina. http://www.elasmo.com.
Bown, T.M. and Kraus M. J.(1988). Geology and
paleoenvironment of the Oligocene Jebel Qatrani
Formation and adjacent rocks, Fayum Depression,
Egypt: U.S. Geological Survey Professional Paper,
1452: 1-60.
Butzer, K.W.(1976). Early Hydraulic Civilisation in
Egypt, a Study in Cultural Ecology. The University
of Chicago Press, Chicago.
Cappetta, H. (1970).Les sélaciens du Miocène de la
région de Montpellier.Palaeovertebrata, mém, ext.,
1- 139.
Cappetta, H. (1976). Selaciens nouveaux du London
Clay del'Essex (Ypresien du Bassin de Londres):
Geobios, 9 (5): 551-574.
Cappetta, H. (1987). Chondrichthyes II. Mesozoic and
Cenozoic Elasmobranchii, in Schults, H.P., Kuhn,
O.(eds.), Handbook of Paleoichthyology: v.3B,
193pp., Stuttgart, New York: Gustav Fischer
Verlag,
Cappetta, H. (2004).Handbook of Paleoichthyology
3B.Chondrichthyes 2 Mesozoic and Cenozoic
Elasmobranchii. 193 pp., ISBN 10: 3899370465
ISBN 13: 9783899370461
Cappetta, H. (2006). Elasmobranchii Post-Triadici,
(Index specierum et generum). In: W. Riegraf (ed.).
Fossilium Catalogus, I: Animalia, v.142, pp. 1-472.
Backhuys Publishers, Leiden.
Cappetta, H., Pfeil, F. and Schmidt-Kittler, N.
(2000).New biostratigraphical data on the marine
Upper Cretaceous and Palaeogene of Jordan.
Newsletters on Stratigraphy 38: 81–95.
Carlsen, A.W. and Cuny, G. (2014).A study of the sharks
and rays from the Lillebælt Clay (Early–Middle
Eocene) of Denmark, and their palaeoecology.
Bulletin of the Geological Society of Denmark,
Vol. 62: 39–88.
CASE, G.R. (1981).Late Eocene selachian from south-
central Georgia. Palaeontographica, Abteilung
A176: 52–79.
Case, G.R. and Borodin, P.D.(2000).Late Eocene
selachians from the Irwinton Sand Member of the
Barnwell Formation (Jacksonian), WKA mines,
Gordon, Wilkinson County, Georgia. Münchner
Geowissenschaftliche Abhandlungen, (A: Geologie
und Paläontologie) 39: 5–16.
Case, G.R. and Cappetta, H.(1990).The Eocene selachian
fauna from the Fayum depression of Egypt.
Palaeontographica, Abteilung A 212 (1–6): 1–30.
Case, G.R. and West, R.M.(1991).Geology and
paleontology of the Eocene Drazinda shale member
of the Kirthar formation, central Western Pakistan.
Part II: late Eocene fishes. Tertiary Research 12 (3–
4): 105–120.
Case, G.R., Udovichenko, N., Nessow, L., Averianov, A.
and Borodin, P.(1996).A Middle Eocene Selachian
Fauna from the White Mountain Formation of the
Kizylkum Desert, Uzbekistan. C.I.S.
Palaeontographica Abteilung A 242: 99–126.
Casier, E. (1949).Contributions à l’étude des poissons
fossiles de la Belgique.VIII. Les Pristidés
éocènes.Bulletin de l’Institut Royal des Sciences
Naturelles de Belgique, 25: 1-52.
Casier, E.(1971). Sur un matériel ichthyologique des
‘‘Midra (and Saila) Shales’’ du Qatar (Golfe
Persique). Bulletin de l’Institut Royal des Sciences
Naturelles de Belgique 47 (2): 1–9.
Cicimurri, D.J. (2007). A partial rostrum of the sawfish
Pristis lathami Galeotti, 1837, from the Eocene of
South Carolina. Journal of Paleontology, 81: 597-
601.
Cicimurri, D.J. and Knight, J.L.(2009). Late Oligocene
sharks and rays from the Chandler Bridge
Formation, Dorchester County, South Carolina,
USA. Acta Palaeontologica Polonica 54 (4): 627–
647.

Zalat et al.
Taxonom y and morphol ogical study on the vertebrate remains o f Shark and rays f auna from the Middle and Late Eocene success ion, Fayoum Depress ion, Egypt

212
Cione, A.L. and Reguero, M. (1994).New records of the
sharks Isurus and Hexanchus from the Eocene of
Seymour Island, Antarctica. Proceedings of the
Geologists Association 105: 1-14.
Compagno, L.J.V., Didier, D.A. and Burgess, G.H.
(2005). Classification of Chondrichthyan Fish In:
Fowler, S.L., Cavanagh, R.D., Camhi, M., Burgess,
G.H., Cailliet, G.M., Fordham, S.V.,
Simpfendorfer, C.A. and Musick, J.A (eds). Sharks,
Rays and Chimaeras: The Status of the
Chondrichthyan Fishes. Status Survey. IUCN/SSC
Shark Specialist Group. 461 pp. IUCN, Gland,
Switzerland and Cambridge, UK
Cook, T.D., Murray, A.M., Simons, El. L., Attia, Y.S.
and Chatrath, P.(2010).A Miocene selachianfauna
from Moghra, Egypt.Historical Biology, 22: 1-3:
78-87.
Dartevelle, E. and Casier, E. (1942). Les poissons
fossiles de l’Angola. Comunicações dos Serviços
Geológicos de Portugal, 22: 99-109.
Dartevelle, E. and Casier, E. (1943). Les poissons
fossiles du Bas-Congo et des régions voisines.
Annales du Musée du Congo Belge, Sér.A
(Minéralogie Géologie, Paléontologie), 3: 1-200.
Dartevelle, E. and Casier, E. (1949). Les poissons
fossiles du Bas-Congo et des régions voisines.
Annales du Musée du Congo Belge, Sér.A
(Minéralogie Géologie, Paléontologie), 2: 205-255.
Dartevelle, E. and Casier, E. (1959). Les poissons
fossiles du Bas-Congo et des régions voisines.
Annales du Musée du Congo Belge, Sér. A
(Minéralogie Géologie, Paléontologie),2(3): 257-
568.
Diedrich, C.G. (2012). Eocene (Lutetian) Shark-Rich
Coastal Paleoenvironments of the Southern North
Sea Basin in Europe: Biodiversity of the Marine
Fürstenau Formation Including Early White and
Megatooth Sharks. International Journal of
Oceanography Volume 2012: 1-22.
doi:10.1155/2012/565326
Dolson, J., El Barkooky, A., Wehr, F., Gingerich, P.D.,
Prochazka, N. and Shann, M.(2002). The Eocene
and Oligocene Paleoecology and Paleogeography
of Whale Valley and the Fayoum Basins:
Implications for Hydrocarbon Exploration in the
Nile Delta and Eco-Tourism in the Greater Fayoum
Basin. Cairo 2002, AAPG/EPEX/ SEG/EGS/EAGE
Field trip-guidebook: 1–79.
Ghosh, B.K. (1959).Some fossil teeth from the tertiary
deposits of Orissa. Jour. Palaeont, 33(4): 675-679.
Gingerich, P.D.(1992). Marine mammals (Cetacea and
Sirenia) from the Eocene of Gebel Mokattam and
Fayum, Egypt: stratigraphy, age and
paleoenvironments. Papers on Paleontology 30: 1–
84.
Hassan, F.A.(1985). Radiocarbon chronology of
neolithic and predynastic sites in upper Egypt and
the Delta. African Archaeological Review 3: 95-
116.
Hassan, F.A.(1986). Holocene lakes and prehistoric
settlements of the Western Faiyum. Egyptian
Journal Archaeological Science 13: 483-501.
Hassan, F.A.(1997). The dynamics of a riverine
civilization: a geoarchaeological perspective on the
Nile valley, Egypt. World Archaeology 29 (1):51-
74.
Hendrickx, S. and Vermeersch, P.(2000).Prehistory from
the Palaeolithic to the Badarian culture. In: Shaw, I.
(Ed.), The Oxford History of Ancient Egypt.
Oxford University Press, Oxford: 17-43.
Iskander, F. (1943). Geological Survey of the Gharag
elsultany sheet no.68/54: Standard Oil Company,
Egypt s. a.,reports,51:p.1-29.
Ismail, A.S.A. and Abd El-Azeam, S.
(2008).Biostratigraphy and paleoenvironment of the
Middle-Upper Eocene Ostracods from west
Fayoum province-Bulletin of the Faculty of Science
Zagazig University; 30: 261-4.
Itoigawa, J., Nishimoto, H. and Karasawa, H.
(1985).Miocene fossils of the Mizunami group,
central Japan. 3. Elasmobranchs. Monograph of the
Mizunami Fossil Museum, 5: 1–89.
Kemp, D., Kemp, L. and Ward, D. (1990).An illustrated
guide to the British Middle Eocene vertebrates, 59
pp. David Ward, London.
Kent, B.W. (1994). Fossil Sharks of the Chesapeake
Region. Egan Rees and Boyer (Eds.): 146 pp.
Maryland.
Kent, B.W. (1999).Part 3.Rays from the Fisher/Sullivan
Site.In “Early Eocene vertebrates and plants from
the Fisher/Sullivan Site (Nanjemoy Formation)
Stanford County, Virginia. Virginia Division of
Mineral Resources, Publication, 152: 39-51.
Kirk, E.C. and Simons, El.L. (2001). Diets of fossil
primates from the Fayum Depression of Egypt: a
quantitative analysis of molar shearing. Journal of
Human Evolution40: 203-229
Kozlowski, J.K.(1983). Qasr el-Sagha 1980:
Contributions to the Holocene Geology, the
Predynastic Settlements in the Northern Fayum
Desert. Panstwowe Wydawnicto Naukowe,
Warsaw.
Kriwet, J. (2005). Additions to the Eocene Selachian
fauna of Antarctica with comments on Antarctic

Zalat et al.
Taxonom y and morphol ogical study on the vertebrate remains o f Shark and rays f auna from the Middle and Late Eocene success ion, Fayoum Depress ion, Egypt

213
selachian diversity. Journal of Vertebrate
Paleontology 25 (1): 1-7.
Kumar, K. and Loyal, R.S. (1987). Eocene Ichthyofauna
from the Subathu Formation, northwestern
Himalaya, India. Jour. Palaeont. Soc. India, 32: 60-
84.
Leriche M. (1926). Les poisons Néogénes de la
Belgique. Mémoires de Museé Royal du d’Histoire
Naturelle de Belgique 32:367–472.
Leriche, M. (1927).Les poissons de la Molasse Suisse.
Mémoires de la Société Paléontologique Suisse, 46:
1-119.
Leriche, M. (1957). Les poisons neogenes de la
Bratagne, de l’Anjou et de la Touraine. Mem. Geol.
Soc. France, N. Ser., 36, Mem. 81: 1-61, Paris.
Longbottom, A.E. (1979). Miocene shark’s teeth from
Ecuador. Bulletin of the British Museum natural
History, Geology, 32 (1): 57-70.
Lowery, D., Godfrey, S.J. and Eshelman, R.(2011).
Integrated Geology, Paleontology, and
Archaeology: Native American use of Fossil Shark
Teeth in the Chesapeake Bay Region. Archaeology
of Eastern North America 39:93-108.
Mehrotra, D.K., Mishra, V.P. and Srivastava, S. (1973).
Miocene sharks from India. Rec. Res. Geol.
Hindustan Publishing Corporation, Delhi: 180-187.
Mondal, S., Das, S. Mallick,S. and Adhikary, D. (2009).
Miocene shark teeth assemblages and ancillary fish
taxa from baripada, orissa: taxonomic revision and
a global Palaeobiogeographic overview. Journal of
the Palaeontological Society of India Volume,
54(2): 135-152
Murray, A.M., Cook, T.D., Attia, Y.S., Chatrath, P. and
Simons, El.L.(2011). A freshwater ichthyofauna
from the Late Eocene Birket Qarun Formation,
Fayum, Egypt. Journal of Vertebrate Paleontology
30: 665–680.
Mustafa, H.A. and Zalmout, I.S.(2002).Elasmobranchs
from the late Eocene Wadi Esh- Shallala Formation
of Qa’ Faydat ad Dahikiya, east Jordan. Tertiary
Research 21: 77–94.
Mustafa, H.A., Zalmout, I.S., Smadi, A.A. and Nazzal,
J.S. (2005).Review of the middle Eocene (Lutetian)
selachian fauna of Jebal eth Thuleithuwat, east
Jordan. Neues Jahrbuch für Geologie und
Paläontologie, Abhandlungen, 237: 399-422.
Otero, R.A., Oyarzún, J.L., Soto-Acuña, S., Yury-Yáñez,
R.E., Pawellek, T. and Adnet, S., Cappetta, H.,
Metais, E., Salem, M., Brunet, M. and Jaeger, J.-J.
(2012). Discovery of an earliest Pliocene relic
tropical fish fauna in a newly detected cliff section
(Sabratah Basin, NW Libya). Neues Jahrbuch für
Geologie und Palaontologie, Abhandlungen, 266:
93-114
Patnaik, R., Sharma, K.M., Mohan, L., Williams, B.A.,
Kay,R. and Chatrath,P. (2014). Additional
Vertebrate Remains from the early Miocene of
Kutch, Gujarat. Special publication of the
palaeontological society of India, No. 5: 335-351.
ISBN: 978-81-926033-2-2
Priem, F.(1905).Sur les poiss ons fossiles de l’Eocène
moyen d’Egypte.Bulletin de la Société géologique
de France 5: 633–641.
Priem, F.(1908).Sur des vertébrés de l’Eocène d’Egypte.
Bulletin de l’Institut Egyptien 5 (2): 1–3.
Rana, R.S., Kumar, K. and Singh, H. (2004).Vertebrate
fauna from the subsurface Cambay Shale (Lower
Eocene), Vastan Lignite Mine, Gujarat, India.
Current Science, 87 (12): 1726-1733
Rana, R.S., Kumar, K., Loyal, R.S., Sahni, A., Rose,
K.D., Mussell, J., Singh, H. and Kulshreshtha
(2006).Selachians from the Early Eocene Kapurdi
Formation (Fuller’s Earth), Barmer District,
Rajasthan, India.Jour. Geol. Soc. India, v. 67: 509-
522.
Sahni, A. and Mehrotra, D.K. (1981).Elasmobranch from
the coastal Miocene sediments of peninsular India.
Biological Memoirs, 5 (2): 83-121.
Sahni, A, and Mishra, V.P.(1975).Lower Tertiary
vertebrates from western India. Monograph of the
Paleontological Society of India, Lucknow, 3: 1-48.
Said, R. (1962). The Geology of Egypt: 377 pp. Elsevier
Publishing Co, Amsterdam, New York.
Said, R. (1990). Cenozoic: in Said,R. (ed.), The Geology
of Egypt. pp. 451-486, Balkema, Netherlands.
Said, R., Wendorf, F., Albritton, G., Schild, R.,
Kobusiewicz, M.(1972). Remarks on the Holocene
geology and archaeology of Northern Faiyum
Desert. Archaeologica Polana 13: 7-22.
Shama, K. and Shided, A. (1994).Contributions to the
Microfiches and Paleogeological Investigation of
the Middle and Upper Eocene Sediments in
Fayoum area. Bull. Fac. of Sci., Assiut Univ.
Egypt, 23 (2-f): 381-418.
Sharma, K.M. and Patnaik, R. (2013).Additional Fossil
Batoids (Skates and Rays) from the Miocene
Deposits of Baripada Beds, Mayurbhanj District,
Orissa, India. Earth Science India, Vol. 6 (IV): 160
– 184.
Simons, El.L. (2005). Eocene and Oligocene Mammals
of the Fayum, Egypt, First International Conference
on the Geology of the Tethys, Cairo University,
November: 439-450.

Zalat et al.
Taxonom y and morphol ogical study on the vertebrate remains o f Shark and rays f auna from the Middle and Late Eocene success ion, Fayoum Depress ion, Egypt

214
Simons, El.L.,Seiffer, E.R., Ryan, T.M. and Attia,
Y.(2007).A remarkable female cranium of the early
Oligocene anthropoid Aegyptopithecus Zeuxis
(Catarrhini, Propliopithecidae).National Academy
of Sciences of the USA PNAS.104 (21):8731-8736
Singh, R. (1985).Contribution to the palaeontology and
Biostratigraphy of the Subathu Formation of Simla
Hills region. Himachal Pradesh. Unpublished Ph.D.
thesis, Panjab University:1-199.
Smadi, A.A., Abu Azzam, H., Zalmout, I.S. and Mus-
tafa,H.A. (2003).Middle Eocene selachian fauna
from Al-Rijla Al-Bayda, Eastern Desert of Jordan.
Abhath Al-Yamouk Basic Science and Engineering,
12: 619-631.
Stromer, E. (1905).Sie Fischreste des mittelren und
oberen Eocäns Von Ägypten.I. Teil: Selachii, B.
Squaloidei, und II. Teil: Teleostomi, A. Ganoidei.
Beitrage zur Palaeontologie und Geologie
Ostrerreich-Ungarns und Orients, Universitats-
Buchhandler in Wein 18:163−192.
Stromer, E.(1905a). Die Fischreste des Mittleren und
Oberen Eocäns von Ägypten. I. Teil: Selachii, B.
Squaloidei, und II. Teil: Teleostomi, A. Ganoidei.
Beiträge zur Paläontologie und Geologie
Österreich-Ungarns, 18: 37-58.
Stromer, E.(1905b). Die Fischreste des Mittleren und
Oberen Eocäns von Ägypten. I. Teil: Die Selachier,
A. Myliobatiden und Pristiden. Beiträge zur
Paläontologie und Geologie Österreich-Ungarns,
18: 163-185.
Strougo, A.(2008). The Mokattamian stage: 125 years
later. Middle East Research Center. Earth Science
Series, Ain Shams University, vol. 22: 47–108.
Strougo, A., Cappetta, H. and Elnahas, S.(2007). A
remarkable Eocene ichthyofauna from the Elgedida
glauconitic sandstone, Bahariya oasis, Egypt, and
its stratigraphic implications. Earth Science Series,
Middle East Research Center, Ain Shams
University, vol. 21: 81–98.
Tiwari, R.P., Mishra, V.P. and Lyngdoh, B.C.
(1998).Lower Miocene fish teeth from Mizoram,
India. Geoscience Journal, 19: 9-17.
Underwood, C.J., Ward, D.J., King, C., Antar, S.M.,
Zalmout,I.S. and Gingerich,P.D. (2011).Shark and
ray faunas in the middle and late Eocene of the
Fayum area, Egypt. Proceedings of the Geologists’
Association, 122: 47-66.
Vliet, H.J. van and Abu El Khair,G. (2010). A new
Eocene marine mammal site in the Qattara
depression (Egypt). Cainozoic Research, 7: 73-77.
Wallett, L.A. (2006). Eocene selachian fauna from
nearshore marine deposits, Ampazony,
northwestern Madagascar. Bsc. Thesis, 59 pp.
Department of Biological Sciences, South Hadley.
Weiler, W. (1929).Ergebnisse der Forschungsreisen Prof.
E. Stromers in den Wüsten Ägyptens. V. Tertiäre
Wirbeltiere. 3. Die Mittel-und obereocäne
Fischfauna Ägyptens mit besonderer
Berücksichtigung der Teleostomie. Abhandlungen
der Bayerischen Akademie der Wissenschaften,
Mathematisch-naturwissenschaftliche Abteilung,
neue Folge, 1: 1–57.
Welton, B.J. (1972). Fossil sharks in Oregon. The Ore
Bin, v. 34: 161-170.
Wendorf, F. and Schild, R.(1976).Prehistory of the Nile
Valley. Academic Press, New York: 222-226.
White, E.I. (1926). Eocene fishes from Nigeria. Bull.
Geol. Surv. Nigeria, v. 10: 1-87.
White, E.I. (1956). The Eocene fishes of Alabama.
Bulletins of American Paleontology, 36: 123-151.
Zalat, A.A. (2015). Holocene diatom assemblages and
their palaeoenvironmental interpretations in
Fayoum depression, Western Desert, Egypt.
Quaternary International 369:86-98.
Zalmout, I.S.A., Antar, M.S.M., Abd-El Shafy, E.,
Metwally, M.H., Hatab, El.E. and Gingerich, P.D.
(2012). Priabonian sharks and rays (Late Eocene:
Neoselachii) from Minqar Tabaghbagh in the
western Qattara depression, Egypt.Museum of
Paleontology, University of Michigan.32 (6): 71-
90.

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Figure 1. Geological map of Fayoum depression showing position of the studied stratigraphic sections: I. Hussein
Wally, II. Birket Qarun, III. Qasr El Sagha area and IV. Wadi El-Afreet.
Figure 2. Field photograph of A: Hussein Walley section, B: Birket Qarun section, C: Qasr el Sagha section and D:
Wadi El-afreet section.

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Figure 3. Middle to Late Eocene studied successions A) Hussein Walley village, B) Birket Qarun, C) Qasr El Sagha,
D) Wadi El-afreet.

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Plate 1: 1-4. Otodus sokolovi (Jaekel) 1. lingual view, 3-4. labial views; 5. Macrorhizodus praecursor Leriche, lingual
view, 6.Fish-fin spine (labial view); 7-8. Rhizoprionodon sp., lingual view; 9. Carcharodon carcharias Linnaeus,
lingual view; 10. Negaprion frequens Dames, lingual view; 11,12. Cretolamna twiggsensis Case, 11. lingual view, 12.
Labial view; 13. Eutrichiurides sp., labial view; 14. Galeocerdo cuvieri Péron & Lesueur, lingual view;
15. Hexanchus agassizi Cappetta, labial view of lower lateral tooth; 16. Rhinoptera sherburni Arambourg; 17.
Leidybatis sp.; 18. Pristis lathami Galeotti.(scale bar= 10 mm).
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