An essay on

Paleontological studies on some whale fossils

from the Eocene of Wadi El-Hitan in the
Fayum Depression of Egypt.

Submitted by

Maamoun Adel Shalaby

Under Supervision of

Prof. Adel Mohamed El-Sayed Genedi

Full professor of geochemistry of sedimentary
rocks, Geology Department, Faculty of Science,
Mansoura University.

2024
 ACKNOWLEDGMENTS
I would like to express my gratitude to several people who have supported me throughout this

project.

First, I want to thank my professor, Prof. Adel Genedi, for his constant support and

encouragement. His guidance has been invaluable to me, and I'm grateful for the opportunity to

pursue my passion in geology.

I'm also deeply thankful to Prof. Hesham Sallam for his dedication and unwavering support.

His mentorship has been instrumental in helping me complete this project.

I'd like to extend special thanks to Abdullah Gohar for his ongoing assistance and for providing

valuable information related to the project.

I'm also grateful to everyone in the Sallam Lab (MUVP) for creating an excellent working

environment and providing support that has been crucial to the project's success.

Finally, I want to express my deepest appreciation to my family for their unwavering support,

encouragement, and motivation. Their belief in me has been a constant source of strength during

the most challenging times.

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 CONTENTS

LIST OF FIGURES ....................................................................................................................3

INTRODUCTION.......................................................................................................................6
Cetacean Paleobiology ..........................................................................................................10
The Fayum Depression .........................................................................................................12
New specimens .....................................................................................................................18
GEOLOGY AND STRATIGRAPHY .....................................................................................23
Geology of the Fayum Depression and surrounding areas ...................................................24
Stratigraphy ...........................................................................................................................27
Stratigraphic Levels Yielding Cetacea in the Fayum Depression ........................................34
Wadi El-Hitan .......................................................................................................................35
METHODOLOGY ...................................................................................................................50
1- Techniques in the Field at Wadi El-Hitan ........................................................................50
2- Laboratory Preparation at MUVP lab ...............................................................................53
SYSTEMATIC PALEONTOLOGY .......................................................................................79
Species: Dorudon atrox (Andrews, 1906). ............................................................................79
Skull ......................................................................................................................................80
Mandible ...............................................................................................................................85
Dentition ...............................................................................................................................85
Axial skeleton (postcranial) ..................................................................................................86
SUMMARY AND CONCLUSIONS .....................................................................................101
REFERENCES........................................................................................................................103
‫ الملخص العربي‬..............................................................................................................................125

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 LIST OF FIGURES

Figure 1.1. Cranial telescoping and posterior migration of bony nares in extinct and extant
cetaceans…………………………………………………………………………….…………19

Figure 1.2. Geographical map of Northern Egypt showing the location of the Fayum Depression
and the Wadi El-Hitan UNESCO World Heritage Site in the Western Desert of
Egypt………………………………………………………...…………………………………20

Figure 1.3. Mollweide (oval globe) projection map showing global paleogeography for the
Paleogene period (45 million years ago)....................................................................................21

Figure 1.4. The ankle, foot, and toes of Basilosaurus isis were excavated in Wadi El-Hitan,
Egypt………………………………………………………………………………………..….22

Figure 2.1. Panorama showcasing the geomorphological features of Wadi El-Hitan……..….41

Figure 2.2. Isolated hill located to the north of Wadi El-Hitan………………………...….…..42

Figure 2.3. View of sand dunes (Ghard Gehannam) east of Garet Gehannam………...……...43

Figure 2.4. Rocky sandy bajada in the northeastern region of Wadi El-Hitan……….....……..44

Figure 2.5. Location map of the Fayum Depression, Egypt. ……...……………………….…45

Figure 2.6. Correlation of stratigraphic units of the Cairo and Fayum areas…………………46

Figure 2.7. Wadi El-Hitan from 40 million years ago.………………………………….……..47

Figure 2.8. wadi El-Hitan now.………………………………………………………….…….47

Figure 2.9. Skeletons and reconstructions of Wadi El-Hitan archaeocetes Basilosaurus isis and
Dorudon atrox mounted in the University of Michigan Natural History Museum………...….48

Figure 3.1. Prof. Hesham Sallam explaining the anatomical features of Dorudon atrox at the
Open-Air Museum of Wadi El-Hitan.……………………………………………………….…57

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Figure 3.2. Principal prospecting for fossils at the Wadi El-Hitan.………………………..….58

Figure 3.3. Transporting tools needed for fossil collecting……………………………………59

Figure 3.4. Transporting equipments to the excavation site………………………...………...60

Figure 3.5. Collecting small-sized fossils using sieves.……………………………………….61

Figure 3.6. Detecting the external boundaries of the specimen.……………………………....62

Figure 3.7. Using chisels and rock hammers for the fossil excavation.…………………...….63

Figure 3.8. Single bone fragment found during prospecting.…………………...…………….64

Figure 3.9. A broken rib on the surface, with another still in situ.……………...…………….65

Figure 3.10. Partial skeleton of Dorudon atrox specimen ready for jacketing.……………….66

Figure 3.11. The trenching process in preparation for jacketing.………………………...…...67

Figure 3.12. Preparing the remains of Dorudon atrox at the Open-Air Museum, Wadi El-
Hitan………………………………………………………………………………………..68

Figure 3.13. MUVP team at the Dorudon atrox site at the end of the fieldwork.………….….69

Figure 3.14. Preparing a layer of gypsum for jacketing……………………………………….70

Figure 3.15. Cleaning and lab preparing the specimen…………………………………...…...71

Figure 3.16. CT scanning at Al-Mawji Radiology Center, Mansoura……………………...…72

Figure 3.17. 3D laser scanning with the Leo Micro 3D scanner……………………………...73

Figure 3.18. Photographing the specimen at MUVP………………...………………………..74

Figure 3.19. Fine preparation using an air scribe pen under the stereozoom microscope at
MUVP……………………………………………………………………………………….…75

Figure 3.20. Specimen preparation under the stereozoom microscope at MUVP……...……..76

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Figure 3.21. Investigating the specimen through measurements and descriptions at
MUVP....77

Figure 4.1. Dorudon atrox skeletal restoration.……………….…………...……………….…90

Figure 4.2. Dorudon atrox skull…………………………………………………………..…..91

Figure 4.3. Dorsal view of the skull of Dorudon atrox…………….………..………………..92

Figure 4.4.Ventral view of the skull of Dorudon atrox……………………………………….93

Figure 4.5.Left first upper molar (M1) ………….…………………………….………………94

Figure 4.6. Right tympanic bulla of Dorudon atrox in dorsal, lateral and medial
views…...…95

Figure 4.7. Labial view of A,the right dentary; B,the left dentary of Dorudon atrox……….....96

Figure 4.8.Lingual view of A,the right dentary; B,the left dental of Dorudon atrox ……….…97

Figure 4.9. Atlas of Dorudon atrox ……………………………………….………………….98

Figure 4.10. Cervical vertebrae of Dorudon atrox …..……………….………….……………99

Figure 4.11. second right rib of Dorudon atrox ……………………...………….…………..100

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 Chapter One

INTRODUCTION

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 INTRODUCTION

Cetacea is one of the most ecologically and morphologically diverse groups of

mammals.Along with dolphins and sirenians (sea cows), cetaceans have fully adapted to life in
water (Gingerich, 2003; Thewissen et al., 2007, Uhen, 2010; Marx et al., 2016).Cetaceans are
carnivorous, feeding on fish and plankton, whereas sirenians are strictly herbivorous (Marx et
al., 2016).Unlike pinnipeds (seals, sea lions, and walruses),cetaceans conduct all life activities—
sleeping, mating, birthing, and nursing—entirely in water (Marx et al., 2016). Instead of fur, they
use a thick layer of blubber for insulation.Their bodies are streamlined, without external features
such as ears or genitals that could increase drag (Gingerich 2003). Their forelimbs have evolved
into flippers used for steering, while propulsion is achieved by moving their powerful tails,
ending in distinctive horizontal flukes (Uhen, 2010).

Taxonomically, cetaceans fall into three major groups (suborders): 1) the toothed whales
(Odontoceti) of the Oligocene to Recent age range; (2) the baleen whales (Mysticeti) of the late
Eocene to Recent; and (3) the ancestral archaic whales (Archaeoceti) of the Eocene, which most
likely evolved into odontocetes and mysticetes (Berta and Sumich 1999; Gingerich et al. 2001;
Uhen 2007), each of which comprises a range of families (Marx et al., 2016; Figure 1.1).

Archaeocetes are identified by primitive traits such as (1) well-developed hind limbs, (2) a
limited number of morphologically distinct teeth (heterodont) that are replaced once in their
lifetime (diphyodont), and (3) a strong connection to land, often for resting or giving birth
(Thewissen et al., 2007; 2009). In contrast, mysticetes and odontocetes are fully aquatic, with no
external hind limbs and the inability to support themselves on land. Both groups have undergone
significant reorganization of their facial bones, known as telescoping, to aid in breathing
(Fitzgerald, 2012).Modern odontocetes are characterized by (1) a single blowhole, (2) numerous
simple, conical teeth (polydont and homodont), and (3) the ability to echolocate, using sound for
navigation and prey detection (Au, 1993).Mysticetes, on the other hand, (1) are often extremely
large, (2) lack teeth as adults, and (3) possess rows of keratinous baleen plates in their upper
jaws, serving as sieves (Werth, 2000).

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Incidentally, the term "whale" lacks precise biological meaning unless used to encompass all
cetaceans. Commonly, it refers to larger species and their presumed relatives, including some
dolphins like the killer whale (Orcinus orca).

The oldest whales exhibit morphological similarities to terrestrial mammals, complicating the
recognition of their evolutionary links. Diagnostic features often involve skull morphology, such
as early telescoping and tooth shape and arrangement—specifically, the alignment of the tooth
row and the absence of crushing basins on the cheek teeth (Thewissen et al., 2007; Uhen, 2010).
However, these traits can be difficult to identify across all cetaceans and may appear in other
non-cetacean mammals.

The most distinct trait uniting cetaceans is the increased thickness and density
(pachyosteosclerosis) of the medial wall of the tympanic bulla, an ear bone at the skull base. This
feature was once considered unique to cetaceans until a similar morphology was found in extinct
artiodactyls known as raoellids (Thewissen et al., 2007; Patel et al., 2024).This discovery is not
problematic, as raoellids are now recognized as more closely related to cetaceans than to other
artiodactyls, effectively categorizing them as cetaceans (Thewissen et al., 2007; Patel et al.,
2024).

Modern whales and dolphins play crucial roles in ocean ecosystems as top predators, nutrient
distributors, and a food source for deep-sea organisms (Croll et al., 2006; Nicol et al., 2010;
Smith and Baco, 2003; Willis, 2014; Wing et al., 2014).They hold several records, often due to
their immense size: the basilosaurid Perucetus colossus, estimated at 340 tonnes (Bianucci et al.,
2023); the sperm whale (Physeter macrocephalus), with the largest brain (up to 8 kg) (Marino,
2009); right whales (Eubalaena spp.), with the largest testes (about 1 tonne) (Brownell and Ralls,
1986); and the longest-lived mammal, the bowhead whale (Balaena mysticetus), living over 200
years (George et al., 1999).All current species are either mysticetes or odontocetes, as
archaeocetes went extinct around 25 million years ago. The Society of Marine Mammalogy
currently recognizes 90 living species, with 84% being odontocetes.

Overall, the modern cetacean population is notably concentrated in three families: the rorquals
(Balaenopteridae), which comprise about 60% of all living mysticetes, and the oceanic dolphins
(Delphinidae) and beaked whales (Ziphiidae), making up roughly 50% and 30% of all
odontocetes, respectively.Interestingly, almost all balaenopterids and about two-thirds of

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ziphiids belong to a single genus (Balaenoptera and Mesoplodon).This uneven taxonomic
distribution likely reflects relatively recent diversification, possibly influenced by the
development of larger brains or specific feeding and mating strategies.Cetacean taxonomy is still
evolving, with new species (including large ones) being discovered relatively frequently. For
instance, a new beaked whale was identified as recently as 2014, and at least one new rorqual
awaits formal description (Dalebout et al., 2014; Sasaki et al., 2006).Living cetaceans vary in
size from about 1 meter to over 30 meters and inhabit oceans and seas worldwide.

Geographically, the highest diversity of modern cetaceans is found at mid-latitudes with sea
surface temperatures around 21°C (Whitehead et al., 2008).Mysticetes migrate long distances
between low-latitude breeding grounds and high-latitude feeding areas (Stern, 2009).Cetacean
feeding strategies are generally categorized into (1) filter feeding, characteristic of mysticetes,
which targets large quantities of small prey; and (2) targeting individual prey using suction,
raptorial feeding, or a combination of both, as seen in odontocetes (Pivorunas, 1979; Werth,
2000).Most species feed primarily on fish and cephalopods, although mysticetes also consume
tiny crustaceans (mainly copepods and krill).The killer whale (Orcinus orca) regularly preys on
other marine mammals and occasionally turtles and seabirds.The false and pygmy killer whales
(Pseudorca and Feresa) may also hunt marine mammals, but less frequently (Werth, 2000).
Feeding occurs at various depths, with sperm whales and beaked whales diving the deepest (over
2.9 km for Ziphius) and the longest, typically 40–70 minutes (Aoki et al., 2007; Hooker and
Baird, 1999; Schorr et al., 2014).In contrast, dolphins, porpoises, and mysticetes generally dive
shorter (up to 10 minutes) and shallower (100–150 m) (Stewart, 2009).

Most odontocetes are highly social. Species like sperm, killer, and pilot whales form matrilineal
family groups, while others have more fluid fission-fusion societies.Group living may protect
against predators (e.g., sperm whales), facilitate cooperative feeding, and serve mating purposes
(Trillmich, 2009).Older females in species like killer and pilot whales experience menopause,
allowing them to support their descendants through daily assistance or allomaternal care (Foster
et al., 2012).In contrast, mysticetes are generally more solitary but gather during migration,
breeding, and cooperative feeding (Brown and Corkeron, 1995). Large groups of pygmy right
whales have been observed (Matsuoka et al., 1996), and individual humpbacks have been noted
to form long-term associations across feeding seasons (Ramp et al., 2010).Both mysticetes and

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odontocetes exhibit cultural behaviors and engage in complex social interactions, requiring
flexible communication and advanced cognitive abilities, which likely contribute to their
intricate vocalizations (May-Collado et al., 2007) and the development of large brains (Marino
et al., 2007).

Cetacean Paleobiology
In discussions of cetacean origins, molecular biologists focus on living cetaceans, or crown
group Cetacea, which includes the two existing clades: Mysticeti and Odontoceti.This group is
also referred to as Neoceti (Fordyce, 2008). Paleontologists, however, often include a variety of
additional forms classified within the paraphyletic suborder Archaeoceti.These extinct taxa form
the stem group leading to crown Cetacea, known as stem Cetacea (Fordyce, 2008). Extensive
evidence over recent decades suggests that both Odontoceti and Mysticeti originated from the
archaeocete family Basilosauridae (Fordyce, 2008; Uhen, 2004).The taxon Pelagiceti formally
recognizes this lineage, encompassing the last common ancestor of Basilosauridae and Neoceti,
along with all their descendants (Uhen, 2008). Here, "archaeocete" refers to Cetacea members
outside of Neoceti, while "Cetacea" broadly includes both archaeocetes and Neoceti.

Initially, the term "Cetacea" was used for living cetaceans but later expanded to include fossil
forms such as Basilosaurus and other archaeocetes due to their close ties to modern cetaceans
(Fordyce, 2008). Similarly, "Neoceti" could be expanded to include forms sharing characteristics
with both subclades of modern Cetacea.Although not yet described, it is probable that fossil
forms exist within Neoceti that are neither Mysticeti nor Odontoceti.

The traits that define the base of the cetacean clade are found in the head, while significant
changes to the body and limbs occur at various stages of cetacean evolution. Early cetaceans
exhibit several synapomorphies (evolutionary innovations) shared with later cetaceans: a
pachyosteosclerotic bulla with a prominent involucrum and sigmoid process (Thewissen et al.,
2001); incisors and canines aligned with the cheek teeth (Uhen, 2007); lower molars lacking
trigonid and talonid basins, and upper molars with a small or absent trigon basin (Thewissen et
al., 2007); cheek teeth adapted for shearing with reentrant grooves on the anterior molar margins,
and a long, narrow postorbital/temporal skull region (Thewissen et al., 2007). Thewissen et al.
(2007) also noted that the raoellid artiodactyl Indohyus possesses an enlarged tympanic bulla

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with an involucrum, indicating that this feature may have originated within traditional
artiodactyls rather than at the base of Cetacea (Patel et al., 2024).

The origin of Cetacea, both broadly and specifically for modern cetaceans (Neoceti), marks
significant transitions in their morphology and ecology. The earliest known whale,
Himalayacetus, dates back approximately 52.5 million years ago (Mya) from the Ypresian
Subathu Formation in Kuthar Nala, India (Bajpai and Gingerich, 1998; Figure 1.3). Fossils like
this provide a minimum age estimate for the emergence of these organisms. While this constraint
doesn't prevent estimates of their origin time, it does present challenges.

One approach to estimating this origin time involves analyzing genetic differences among living
members and using models of gene evolution. However, challenges arise because genes may
evolve at varying rates across different branches, necessitating more complex evolutionary
models.Additionally, precise divergence times from the fossil record are crucial (Uhen, 2008),
but these can be uncertain due to issues with calibration nodes and the accuracy of fossil age
determinations.Molecular dating typically only provides insights into the origins of crown
clades, excluding stem taxa known only from fossils.Another method uses statistical properties
of fossil distributions over time to construct confidence intervals for the group's origin.
Typically, the earliest fossil provides the best estimate, with the confidence interval based on
available fossil evidence.Although these methods rely on assumptions about the fossil record
(Uhen, 2010), they are robust when the record is well-sampled.

Gingerich and Uhen (1998) estimated the origin of Archaeoceti (and thus Cetacea) at 49.5 Mya,
with a 95% confidence interval extending to 51.64 Mya. This estimate is slightly younger than
the Himalayacetus fossil at 52.5 Mya (Bajpai and Gingerich, 1998), primarily due to revised age
estimates for the Subathu Formation rather than methodological flaws. Utilizing data from 160
collections in the Paleobiology Database (Uhen, 2005), the classic confidence interval for
archaeocetes suggests an origin for the earliest cetaceans around 54 Mya. Advances in fossil
discoveries and dating techniques have pushed back the origin time while refining the confidence
interval. Improved sampling of archaeocete fossils has reduced uncertainty regarding when this
group first appeared. Recent molecular clock estimates suggest the split between
Hippopotamidae and Cetacea occurred approximately 54.9 Mya.

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The earliest cetacean fossils are found in the Indo-Pakistan region, which was part of the ancient
Tethys Sea during the early stages of the collision between Asia and India. While there is some
debate about the exact age of Pakicetus and Himalayacetus, several other fossils from the early
Eocene (Ypresian) have been discovered. For example, a lower molar fragment initially
classified as Ichthyolestes from the Subathu Formation of Babbian Gala was later reassigned to
Pakicetus (Thewissen and Hussain, 1993). Another fossil, possibly a canine of Cetacea
(Kutchicetus minimus?), was found in the Ypresian Panandhro lignite of Kachchh, India. Despite
being fragmentary, these specimens provide evidence of cetaceans in Indo-Pakistan during the
late early Eocene.

The geological formations yielding Ypresian and early Lutetian whale fossils vary widely.
Pakicetus, often considered the oldest whale, was initially discovered in the fluvial red beds of
the Kuldana Formation, alongside other fossils of freshwater vertebrates and invertebrates. This
initially supported the idea that early whales evolved in freshwater continental habitats before
expanding into coastal and open marine environments. However, this narrative became more
complex with the discovery of Himalayacetus in the marine Subathu Formation, which predates
the Kuldana Formation (Bajpai and Gingerich, 1998). Another ancient whale specimen from the
Panandhro lignite is described as from "backswamp deposits," indicating a clearly freshwater
origin.

Other Lutetian deposits in Indo-Pakistan where whales have been found range from freshwater
fluvial deposits to coastal and more open marine settings. Analysis of oxygen isotopes in dental
enamel suggests that early cetaceans like Himalayacetus and Pakicetus inhabited freshwater
environments, despite Himalayacetus being found in marine deposits (Clementz et al., 2006).
This suggests either that Himalayacetus lived primarily in freshwater with occasional forays into
the marine environment, or that its remains were transported into marine settings post-mortem.
Later Lutetian whales such as Babiacetus, Dalanistes, and Remingtonocetus show dental oxygen
isotope values similar to those of modern semiaquatic pinnipeds, indicating a comparable
lifestyle.

The Fayum Depression

The Fayum Depression (Figure 1.2) is an oasis situated about 80 km southwest of Cairo in
Egypt's Western Desert (El-Shabrawy and Dumont, 2009). This region is celebrated for its

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numerous fossil-rich sites that yield a diverse range of vertebrate remains spanning the Eocene
to Oligocene periods (Andrews, 1906a; Seiffert et al., 2006: 2008; Simons and Rasmussen,
1990).Since the nineteenth century, prominent paleontologists like Charles W. Andrews,
Eberhard Fraas, Ernst Stromer von Reichenbach, and Henry F. Osborn have extensively
excavated and studied these sites. Their research, alongside subsequent studies, has greatly
enhanced our understanding of the early mammalian communities in Africa during this time,
particularly the origins and evolution of groups such as Proboscidea, Cetacea, and Primates (e.g.,
de Vries et al., 2021; Simons, 2008; Simons and Rasmussen, 1990).

History of the study of Fayum Cetacea

Cetaceans are one of the most extensively studied and best-known fossil vertebrate groups from
the Paleogene of Egypt. Complete skeletons have been recovered from the Gehannam Formation
and Birket Qarun Formation (from the Wadi El-Hitan or Valley of Whales); however, they occur
only sparsely in the Qasr El-Sagha Formation (Gameil et al., 2016; Gingerich, 1992, 2010). The
first fossil whales from Africa were found by Georg Schweinfurth in 1879, representing the first
archaeocete cetaceans known from the eastern hemisphere. The remains were found in the Birket
Qarun Formation on the island of Geziret el Qarn in the center of Lake Birket Qarun (Dames,
1883a; Gingerich, 2010). These fossils include vertebrae (Dames, 1883a, 1894) and skull
fragments (Dames, 1883b) of “Zeuglodon” sp. as well as the holotype of “Zeuglodon'' (now
Saghacetus osiris), an almost complete dentary (Dames, 1894)

At the beginning of the twentieth century, Beadnell collected a considerable amount of

archaeocete remains from the Birket Qarun Formation, which were briefly described by Andrews
(1901c, 1904f). Andrews (1904f) established the species “Zeuglodon” isis (now Basilosaurus
isis). The name of this species was originally proposed by Beadnell in a memoir about the
geology of the Fayum Province (Beadnell, 1905) that was, however, published after Andrews
(1904f). The holotype is a complete mandible containing the entire dentition. Beadnell also
collected fossil archaeocetes from an area west of Garet Gehannam. The fossils were so abundant
there that he called this area “Zeuglodon Valley” (Beadnell, 1905), which was later named Wadi
El-Hitan or Valley of Whales. During this phase, he collected a complete skull of an archaeocete
that was later described as the holotype of “Prozeuglodon” atrox (now Dorudon atrox), by
Andrews (1906a). All fossil archaeocetes collected by Beadnell and Andrews at the beginning

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of the twentieth century were described in detail by Andrews (1906a) in his monograph about
the vertebrate fauna of the Fayum.

Andrews (1906a) described the holotype skull of “Prozeuglodon'' as an intermediate between

Protocetus and “Zeuglodon”, but in 1908 he revised the specimen, concluding that it instead
represented a juvenile with deciduous teeth. Therefore, “Prozeuglodon” atrox was long thought
to be the juvenile form of “Zeuglodon” isis (e.g. Barnes and Mitchell, 1978; Kellogg, 1936).
However, Gingerich et al., (1990) studied complete skeletons of both species and showed that
“Prozeuglodon” atrox is not a juvenile of “Zeuglodon” isis and thus belongs to a separate genus
and species. The archaeocete fossil collected by Andrews and Beadnell was furthermore studied
by Dart (1923), who established three new species of “Zeuglodon” (Zeuglodon sensitives,
Zeuglodon elliotsmithii, and Zeuglodon intermedius) based on natural endocasts of the brain
cavity.

Besides Beadnell and Andrews, Stromer and Fraas also collected archaeocete remains from the
Fayum region, north of Birket Qarun. They discovered a new skull and lower jaw of
“Zeuglodon”osiris from the Qasr El-Sagha Formation (Stromer, 1902) and established the
species “Zeuglodon” zitteli based on a natural endocast of the nasal region, the skull remains, as
well as vertebrae (Stromer 1903). More recently, Gingerich (2007) established a new species of
archaeocete whale from that region based on the vertebrae collected by Stromer and additional
vertebrae from Garet el Esh, which he named in honor of Stromer, Stromerius nidensis.

During the same period, Fraas and Markgraf collected a cranium and associated postcranial
remains of a small archaeocete from Gebel Mokattam. The specimen was described by Fraas
(1904) as Protocetus atavus, a new genus and species, which became the type of genus of the
new family Protocetidae (Stromer, 1908). Another skull from the same formation was named by
Fraas (1904, 1906) as “Mesocetus” schweinfurthi (now known as Eocetus schwinfurthi). In 1906,
Fraas and Markgraf excavated a large skeleton of “Zeuglodon” isis west of Birket Qarun
comprising a skull with a length of 1.3 m and a sequence of vertebrae and ribs about 10 m in
length (Fraas, 1906; Gingerich, 2010). This skull was described by Stromer (1908) and Slijper
(1936) and was later depicted by Heizmann (1991). After the excavations of Fraas and Markgraf,
Stromer together with Markgraf went to the Fayum Depression to collect fossil whales
(Gingerich, 2010). They recovered two large archaeocete vertebrae from Gebel Mokattam, an

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archaeocete skull with jaws, and some vertebrae from the Birket Qarun Formation (Stromer,
1904).

After the expeditions of Fraas and Stromer, several scientists and fossil collectors from the USA,
such as Henry F. Osborn and Walter Granger (in 1907), as well as Robert H. Denison and Paules
E. P. Deraniyagala (in 1947) visited the areas around Qasr El-Sagha, Birket Qarun, and Wadi
El-Hitan (Gingerich, 2010). They collected many large zeuglodonts from Wadi El-Hitan
(Deraniyagala, 1948; Kellogg, 1936; Osborn, 1907) and partial skulls of “Zeuglodon” isis, as
well as endocranial casts of “Prozeuglodon” atrox (Pilleri, 1991). Kellogg (1936) was the first
to mention the use of Basilosaurus over Zeuglodon, as he regarded them as synonyms. Later,
Gingerich et al., (1990) and Uhen (2004) supported this taxonomic assignment. Furthermore,
Kellogg (1936) placed “Zeuglodon” osiris in Dorudon, and later Uhen (2004) included
“Prozeuglodon” atrox in Dorudon as well. The species “Zeuglodon” osiris was later placed into
Saghacetus by Gingerich (1992).

Moustafa (1954) described a partial skull of a subadult “Zeuglodon” isis in the Birket Qarun
Formation that was collected by him in 1950. In the following two decades, paleontologists
Simons and Meyer visited the area around Birket Qarun, Qasr El-Sagha, and Wadi El-Hitan
(Gingerich, 2010). They found skulls of “Zeuglodon” isis in the Birket Qarun Formation and
some archaeocete remains in the Qasr El-Sagha Formation (Simons and Wood, 1968). Barnes
and Mitchell (1978) provided a review of African cetaceans, in which they included
Prozeuglodon atavus, Eocetus schweinfurthi, and Pappocetus lugardi in Protocetidae. They also
placed among others “Zeuglodon” osiris and “Zeuglodon” isis into Basilosauridae.

Shortly after, Gingerich and Simons excavated in the Fayum region, uncovering new archaeocete
remains from the Birket Qarun Formation (Wadi El-Hitan) and Qasr El-Sagha Formation in 1983
(Gingerich, 2010). They found many well-preserved skeletons of archaeocete whales. The most
common taxa in Wadi El-Hitan were Basilosaurus isis and Dorudon atrox, both represented by
equal numbers of individuals (Gingerich, 2010; Figure 1.4). Both species are described to be
morphologically quite similar with slight osteological differences (Gingerich and Smith, 1990;
Gingerich et al., 1990; Luo and Gingerich, 1999). Furthermore, recent studies using stable
isotopes confirmed that both Basilosaurus and Dorudon were fully aquatic (Clementz et al.,
2006). Moreover, Gingerich et al. (1990) focused on the development of the hindlimbs in

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archaeocetes based on fossils from Wadi El-Hitan ultimately providing insights into archaeocete
evolution (Figure 1.4). In addition to Basilosaurus isis and Dorudon atrox, further archaeocete
whales are known from Wadi El-Hitan such as Ancalecetus simonsi (Gingerich and Uhen, 1996).

In a short period, Gingerich described two new genera and species of basilosaurid archaeocete
whales. The first, Stromerius nidensis, was based on an articulated set of thoracic, lumbar, and
caudal vertebrae extracted from the Temple Member of the Qasr El Sagha Formation in the
Fayum Depression. Stromerius is distinguished by its lumbar vertebrae, which have longer and
more anteriorly directed metapophyses. The second, Masracetus markgrafi, is notable for its
large lumbar centra that are relatively short anteroposteriorly (Gingerich, 2007).

In subsequent years, Gingerich provided a detailed review of cetacean discoveries in Egypt over
the past decades, summarizing African cetacean localities, including Egyptian sites. He
described several cetaceans from Egypt and Africa, such as Protocetus atavus, Eocetus
schweinfurthi, Dorudon atrox, Basilosaurus isis, Masracetus markgrafi, Ancalecetus simonsi,
Saghacetus osiris, Stromerius nidensis, Schizodelphis aff. sulcatus, Delphinus vanzelleri, and
Cyrtodelphis aff. sulcatus (Gingerich, 2008; 2010).

In 2011, Giovanni Bianucci and Philip Gingerich named a new protocetid whale, Aegyptocetus
tarfa, discovered in a commercial rock quarry in Egypt. The quarry, located on the northern flank
of Wadi Tarfa, yielded a unique specimen with thin dentary lateral walls, wide mandibular
canals, and thin anterolateral walls of the tympanic bullae, suggesting an increased ability to hear
in aquatic environments. The specimen also had well-developed ethmoidal turbinal bones,
indicating a functional sense of smell (Bianucci and Gingerich, 2011; Peri et al., 2019).

In 2012, Fahlke studied bite marks on juvenile Dorudon atrox specimens described by Uhen
(2004) using CT scans, digital surface scanning, and 3D reconstruction. She found that the bite
marks matched the dentition of Basilosaurus isis, suggesting that B. isis preyed on juvenile D.
atrox. This made B. isis the only known archaeocete to prey on other cetaceans (Fahlke, 2012;
Voss et al., 2019).

In 2016, Ryan M. Bebej and colleagues described a new genus and species of remingtonocetids,
Rayanistes afer, from the Midawara Formation in Egypt. This discovery extended the known
range of Remingtonocetidae. The fossils included a sacrum with four vertebral centra, several

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lumbar and caudal vertebrae, an innominate with a complete ilium, ischium, and acetabulum,
and a nearly complete femur. The morphology suggested a specialized mode of locomotion, with
a larger ischium supporting powerful hind limb retraction and posteriorly curved neural spines
on the lumbar vertebrae enhancing flexibility (Bebej et al., 2016).

In early 2019, Manja Voss and colleagues confirmed Fahlke's findings and showed from stomach
contents that B. isis from Wadi El-Hitan preyed on juvenile D. atrox and giant fishes, providing
clear evidence of its diet and supporting the predator-prey relationship between B. isis and D.
atrox (Voss et al., 2019).

Recently, Gingerich and his colleagues described a new African protocetid whale, Aegicetus
gehennae, based on a partial skull and associated postcranial skeleton from the Gehannam
Formation near the Wadi El-Hitan World Heritage Site. The elongated vertebrae, loss of
sacroiliac articulation, and reduced hind limbs suggest A. gehennae was more aquatic and less
skilled at foot-powered swimming than earlier protocetids. A. gehennae is the latest surviving
protocetid known, surviving until about 38 million years ago (Gingerich et al., 2019).

Recently, Gohar et al. (2021) described Phiomicetus anubis, a medium-sized protocetid whale
discovered in the Middle Eocene Midawara Formation in the southwest part of the Fayum
Depression. The holotype, which includes a partially preserved skull, mandibles, vertebrae, and
ribs, allowed the researchers to identify Phiomicetus as the most primitive African protocetid,
noted for its adept hunting abilities targeting elusive prey (Gohar et al., 2021).

More recently, Anter et al. (2023) documented Tutcetus rayanensis, a basilosaurid whale from
the middle Eocene (early Bartonian) of the Sath El-Hadid Formation in the Fayum Depression,
Egypt. The holotype, comprising an incomplete skull with mandibles, the hyoid apparatus, and
the atlas vertebra of a small-sized subadult, was found in a limestone block. This whale,
estimated at 2.5 meters in length and approximately 187 kilograms in body mass, represents the
smallest known basilosaurid species and one of the oldest globally. The discovery significantly
broadens our understanding of basilosaurid size diversity and underscores the substantial
evolutionary development of whales during the middle Eocene.

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New specimens
A new fossil basilosaurid whale, Dorudon atrox, in this project includes a skull, mandibles, and
associated vertebrae and ribs collected from Wadi El-Hitan of the Fayum Depression in Egypt.
These fossils provide information about the paleoecology, evolution, and systematics of Eocene
basilosaurids in Egypt because they include cranial and postcranial elements found in
association.

Of all the contributions towards understanding the natural history of the Fayum cetacean fossils,
only very few published reports discussed the structure, systematics, habitats, and secondary
adaptation of the basilosaurid whales (Uhen, 2004).

Recovery of such Dorudon specimens contributes to a better understanding of the paleobiology

and paleoecology in the basilosaurid archaeocetes.

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Figure 1.1. Cranial telescoping and posterior migration of bony nares in extinct and extant
cetaceans. A. Early Eocene remingtonocetid archaeocete, the skull of Andrewsiphius sloani
(modified from Bajpai et al., 2011). B. Middle Eocene archaeocete (Basilosauridae) skull of
Dorudon atrox (modified from Uhen, 2004). C. Recent odontocete, the skull of Tursiops
truncatus (modified from Mead and Fordyce, 2009). D. Recent mysticete, the skull of
Balaenoptera sp. (modified from Berta et al., 2006). Thick arrows indicate the location of bony
nares and similar coloration of bones among skulls indicates homology. Life restorations of taxa
illustrated by Carl Buell.

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Figure 1.2. Geographical map of Northern Egypt showing the location of the Fayum Depression
and the Wadi El-Hitan UNESCO World Heritage Site in the Western Desert of Egypt (from Voss
et al., 2019).

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Figure 1.3. Mollweide (oval globe) projection map showing global paleogeography for the
Paleogene period (45 million years ago) © Ron Blakey, NAU Geology.

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Figure 1.4. The ankle, foot, and toes of Basilosaurus isis were excavated in Wadi El-Hitan,
Egypt. This find was described by Gingerich et al. (1990). Photograph ©1991 Philip Gingerich.

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 Chapter Two

GEOLOGY AND STRATIGRAPHY

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 GEOLOGY AND STRATIGRAPHY

The geology of the Fayum Depression, in northeastern Egypt has been studied by a
long list of geologists from the late nineteenth century to the present (Seiffert, 2008). Here we
will review some of these major studies of geology: The distribution of Eocene deposits yielding
sirenian fossils in Egypt was largely controlled by the tectonic history of the African continent.
Major rifting events during the Late Jurassic and Early Cretaceous along the North African-
Arabian margin of Tethys caused a series of pull-apart basins to form in northern Egypt and
Sinai. This system, the Syrian Arc system of Krenkel (1924), extended as far north as western
and central Jordan and most of Syria. The continuous movement of the Afro-Arabian plate to
the northeast imposed severe compressional forces through the Late Cretaceous and middle to
late Eocene, which gave final shape to this system and its basins (Guiraud et al., 2005). Two
aspects are of interest here: first, the geology of the Fayum and surrounding areas, and second,
the stratigraphy of the cetacean-bearing formations in northern Egypt.

Geology of the Fayum Depression and surrounding areas

The Fayum, or Fayum Basin (also known as the Fayum-Gindi Basin), spans 120 kilometers wide
and is characterized by a graben filled with 2 kilometers of Eocene sediments (Salem, 1976).
These sediments overlay Upper Cretaceous strata unconformably. The basin is flanked by two
major highlands: El Kattaniya-Abu Rosh to the north and Nashfa to the south of Wadi El Rayan,
with the latter marking the southern boundary of the Syrian Arc system.

Within this graben, the Fayum Basin exemplifies a classic model of marine progradation. The
lower and lower-middle parts of the Eocene are dominated by carbonate shelf facies, while the
upper-middle and upper Eocene feature siliciclastic estuarine, lagoonal, and deltaic facies
(Salem, 1976). Northern Egypt, including the Fayum Basin, was a broad, stable marine platform
during the middle and late Eocene, following Syrian Arc rifting and compression but preceding
Red Sea rifting. The platform experienced passive subsidence with minimal tectonic influence,
preserving a rich record of shallow marine strata.

In this geological review, I focus on Wadi El-Hitan as representative of the Fayum Depression,
renowned globally for its significant concentration of cetacean fossils.

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Geomorphology
The geomorphology of the Fayum region, including the Fayum Depression, has been extensively
studied by Beadnell (1905) and Said (1962). Tectonic movements, as well as physical and
chemical weathering processes driven by water and wind actions, have shaped the landscape of
the area.

The Fayum Depression floor exhibits low elevations, ranging from a maximum of +25 meters
above sea level to a minimum of 45 meters below sea level, covered predominantly by Nile
alluvium. On the western side of the Fayum Depression, there are deposits of gravel and coarse
sands interspersed with oblate, rounded pebbles of chert and shells dating back to the Pleistocene
era. Dune are prevalent in both the southwestern and northwestern parts of the depression.

To the north of the depression lies the Libyan plateau, gently sloping towards the north and
northwest, rising to approximately +200 meters above sea level. This plateau features prominent
escarpments such as Jebel Qatrani and Qasr El-Sagha escarpments, situated north of Qarun Lake
(Figure 2.1). Cuesta terraces are found within the escarpments in the northern part of the Fayum
Depression.

To the southwest of the Fayum Depression are the Wadi El Rayan Lakes, with the renowned
Wadi El-Hitan area located northwest of these lakes. The topographic and geomorphological
maps illustrate that Wadi El-Hitan is characterized by various landforms including tablelands,
escarpments, pediplains, cuestas, isolated hills, rocky sandy bajadas, Wadi plains, sand dunes,
and sand sheets.

Tableland: Located north and northwest of Wadi El-Hitan, this area is characterized by Birket
Qarun beds topped with terraces of Qasr El-Sagha beds. These terraces form due to differences
in rock composition and structure, with gently to moderately steep slopes formed by erosion.

Escarpments: These formations result from back erosion of various beds, where softer layers
erode more quickly, leaving harder layers as cliffs. The high northern escarpments bound the
study area, extending from northeast to southwest of Wadi El-Hitan. They consist of alternating
sandstone and shale from the Birket Qarun Formation, and calcareous sandstone and banks of
oyster and Carolio beds from the Qasr El-Sagha Formation, featuring steep cliffs and structural
benches with elevations around 50 meters.

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Pediplains: Found in the middle to north and northeast of Wadi El-Hitan, pediplains feature a
rocky surface of yellowish-white sandstone (the mangrove layers) with whale skeletons,
representing the top of Gehannam on the Wadi El-Hitan floor. These areas are occasionally
covered by weathered polished gravel and sand sheets.

Cuesta: Asymmetric ridges with a short, steep escarpment on one side and a gentle slope on the
other. Cuestas are located in the southern borders of Wadi El-Hitan, cut by small tectonically
originated valleys, dividing them into smaller cuestas. Main stratigraphic sections such as Garet
Gehannam and Mingar El-Hut cuestas are found here.

Isolated hills: Scattered across the middle and western sides of Wadi El-Hitan, these hills vary
in size with steep sides and small flat tops. Composed of sandstone from the Birket Qarun
Formation (Figure 2.2).

Dunes: A longitudinal dune east of Garet Gehannam extends NW-SE for about 4 kilometers in
length and 200 meters in width (Figure 2.3), consisting of well-sorted quartz sand grains. Smaller
longitudinal dunes are also found south of Wadi El-Hitan.

Rocky sandy bajada: Broad, flat, gently sloping areas surrounding the high cuestas of the
southern borders of Wadi El-Hitan and Garet Gehannam. Composed of gravely, pebbly sand,
some alluvial mud, and exposed hard rocks of the Gehannam Formation (Figure 2.4).

Sand sheets: Covering the north and southeast borders of Wadi El-Hitan around Garet
Gehannam and Ghard Gehannam, these sheets consist of fine to moderately coarse, well-sorted
quartz sand grains.

Wadi Plain: Located north of the cuestas in the middle of Wadi El-Hitan, this deep and narrow
valley widens to the east. Composed of alluvial sediments of mud and sand with meandering
channels from surrounding high cuestas.

Stratigraphy
The geology of the Fayum Depression has been extensively studied by researchers over the past
two centuries, with key contributions from (Beadnell, 1905), (Bown and Kraus, 1988), and
(Gingerich, 1992). The Fayum Depression itself is characterized as a 120-kilometer-wide graben

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structure filled with Eocene–Oligocene sediments that unconformably overlay Upper Cretaceous
formations (Salem, 1976).
Within the northern part of the Fayum Depression (Figure 2.5), terrestrial vertebrate-yielding
sediments belong to three formations spanning from the Late Eocene to the Early Oligocene.
These formations are capped by the Widan El Faras Basalt, which varies in thickness from 2 to
25 meters and has been radio-isotopically dated (40Ar/39Ar) to 23.64 ± 0.035 million years ago
(Kappelman et al., 1992). Both the Paleogene sediments and the Widan El Faras Basalt are
overlain by Neogene sediments of the Khashab Formation.
The classification of Eocene rocks in Egypt lacks a unified regional consensus. Initial
classifications followed Zittel's major divisions, later formalized into the Libya and Mokattam
groups by Said (1962). Numerous new formational names have since been introduced to
differentiate specific facies or designate lithological types.
According to Beadnell (1901, 1905) and Gingerich 1992, the lowermost four formations of the
Wadi El Rayan series are Lutetian and Bartonian in age, and they are exposed in the Wadi El
Rayan valley floor, mesas, and escarpments. These formations were mapped by Beadnell (1905)
and Iskander (1943) as part of the Wadi El-Rayan Series. The entire section includes (from the
bottom):

Muweilih Formation (Iskander, 1943)

The Muweilih Formation, is named Muweilih because it forms the floor of the Muweilih Oasis
(the Salt Oasis in Arabic) south of the Wadi El Rayan. It was suggested by Iskander (1943) to
define the limestones intercalated with claystone in the southern part of the Fayum Depression.
The Muweilih Formation represents the lowest formation exposed in the area and it lies
unconformably over the Samalut Formation and conformably underlies the Midawara
Formation. It is made of about 36.5 m of shallow marine, greenish-brown to grey, gypsiferous,
saliferous compact claystone, yellow to yellowish-grey, sandy, burrowed, and fossiliferous
limestones (Helal and Holcová, 2017).

Midwara Formation (Iskander, 1943)

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This formation was introduced by Iskander (1943) to describe the Eocene fossiliferous
limestones with claystone intercalations exposed in the southern part of the Fayum Depression.
The name of the Midawara Formation is based on its characteristic topographic features. The
Midawara Formation conformably overlies the Muweilih Formation and conformably underlies
the succeeding Sath El Hadid Formation. The Midawara Formation is conformably underlined
by the nummulitic limestone of the Muweilih Formation and overlined by the snow-white
limestone of the Sath El Hadid Formation. It is mainly composed of both light and dark brown
shales, sandy shale, sandy limestone, and green glauconitic, shelly sand. It is majorly exposed
as isolated or connected conical hills or hillocks of relatively low height ranging from 10 to 20
meters above ground level.
According to the stratigraphic and biostratigraphic correlations between the Fayum Depression
and Gebel Mokattam region in Cairo found that the Midawara Formation ranges the fourth and
fifth Mokattamian stages (MK4-MK5) and this makes it middle to late Lutetian in age and this
is confirmed by the presence of Nummulites gizehensis and evidence that top of the Midawara
Fm. spans the Nanoplankton zone 16 (Strougo, 2008), which is aged at about 42 Ma
(Leuterbacher et al., 2004).
Sirenian, whale remains, Phiomicetus anubis (Gohar et al., 2021) and Rayanistes afer (Bebej et
al., 2016), and sharks were recorded in the upper one-third of this formation in the bioturbated
glauconitic sands near Dur El-Milaha east of Qusor El-Arab.

Sath El Hadid Formation (Iskander, 1943)

The Sath El Hadid Formation, "Iron surface" in Arabic, is a nummulitic limestone with both
large and small Nummulites and caps several mesas in the Wadi El Rayan area.
It conformably overlies the Midawara Formation and underlies the following El Gharaq
Formation. The contact in the base of the Sath El Hadid Formation in the type locality (Gebel
Sath El Hadid) is gradual and represented by the basal snow-white marly bryozoan limestone of
the Sath El Hadid and the topmost glauconitic sandy brown limestone of the Midawara
Formation. The bryozoan limestone of the top of the Sath El Hadid Formation is in sharp contact
with the reddish to pink limestone enriched by Nummulites lyelli of the overlying El-Gharaq
Formation. This contact with the El-Gharaq Formation can be defined by the first appearance of
brown and light brown shales above the snow-white nummulitic limestone. The top of the Sath

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El-Hadid Formation is notable in having secondary flint concretions. The basilosaurid whale,
Tutcetus rayanensis was unearthed from this Formation (Anter et al., 2023)

El-Gharaq Formation (Iskander, 1943)

The El-Gharaq Formation was named by Iskander (1943) for exposures near the village of El-
Gharaq in western Fayum. The upper part of the formation is exposed in the southeastern part of
Wadi El-Hitan, where it forms the lowest valley floor. The El Gharaq Formation is a white to
gray marine limestone rich in large Nummulites — Bartonian late middle Eocene in age.

Gehannam Formation (Beadnell, 1905)

The Gehannam Formation was initially named "Ravine Beds" by Beadnell (1905) because its
deposits are exposed in ravines and small valleys; below the alluvium succession at the deep-
water channels or Ravine in the Fayum Depression. Said (1962, 1990) changed its name and
called it the "Gehannam Formation'' as it is exposed very well, with a distinct base and top, at
Garet Gehannam, the type section of the Gehannam Formation (West of the Fayum Depression).
It consists mainly of gypsiferous claystone, with marls and argillaceous limestone beds. It was
deposited on a shallow shelf and dated to Bartonian through early Priabonian, Middle to Late
Eocene, in age (Haggag, 1985; 1990; Gingerich, 1992). It is marine, some 30 m thick in Wadi
El-Hitan, and exposed over a large area in the central and southeastern part of the protected area
where it often forms the valley floor. The Gehannam Formation is a gray calcareous shale, with
interbedded marl and glauconitic sandstone — early Priabonian in age. The El Gharaq –
Gehannam formational boundary is a paraconformity, which corresponds to the first of the
Priabonian low sea stands (Pr-1) identified by Hardenbol et al., (1998). Basilosaurid whales and
protosirenid sea cows are common in the Gehannam Formation.
The Gehannam Formation forms the floor of much of the cultivated Fayum areas and has a
changeable thickness and may reach 45 meters near the Garet Gehannam plateau (Figure 2.3).
The lower level of the formation is marked by thick deposits of glauconitic sandstone and marls
that lose large benthic forams but are rich in fish scales, shark and ray teeth, and marine mammal
skeletons. Recently, Gingerich et al., (2019) described a protocetid whale, Aegicetus gehennae,
from this level. Marl and glauconite deposits of the Gehannam Formation are substituted at the
higher stratigraphic levels with gypseous fine sands and brown shale overlined with pale

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sandstone of the Birket Qarun Formation. Gehannam Formation was divided into two members,
a lower Gehannam Marl Member which is equal to the Ravine Beds, and the upper Gehannam
Shale Member which is the Birket Qarun Formation (Ismael and Abdel-Kereem, 1971a,b).
Gingerich (1992) noted that the upper 24 m of 70 m of Beadnell’s section, the “Ravine Beds” at
Garet Gehannam, are sandstones and so he included it in the overlying Birket Qarun Formation.
East of Garet Gehannam and just west of Lake Qarun, near the village of Guta, the Gehannam
Formation is represented by limestone and marly limestone facies (Strougo, 1985a,b; 1986;
1988; 1992). In Wadi El-Hitan the middle and top parts of the Gehannam Formation are exposed
and form the base of the valley in many areas.
Gingerich (1992) marked the boundary between the two formations at the Camp White Layer
(CWL). We now follow Strougo (2008) in placing the Gehannam-Birket Qarun formational
boundary at the top of the fine-grained calcareous sandstone that is ledge-forming in the vicinity
of Garet Gehannam, at the base of Minqar el Hut, and elsewhere in Wadi El-Hitan.

Birket Qarun Formation (Beadnell, 1905)

The stratigraphically lowest formation that includes terrestrial fossils is the Birket Qarun
Formation (Birket el Qarun Series after Beadnell, 1905). The thickness of the Birket Qarun
Formation ranges from 20 to 85 m and includes mainly clays, shales, and thick fine-grained
sandstones, but also ferruginous bioclasts and calcareous grits (Anan and El Shahat, 2014.
Zalmout and Gingerich, 2012). One of its most characteristic layers is the Camp White Layer,
which can be up to 2 m thick. It is known for its richness in marine mammals (Gingerich,1992)
and for the existence of vertical, rod-like structures which had been interpreted as “mangrove
pneumatophores'' but were most recently re-evaluated and described as burrows of different
invertebrates (Gee et al., 2019).
The Birket Qarun Formation comprises an unnamed lower member and the Umm Rigl Member.
The assignment of the Umm Rigl Member to the Birket Qarun Formation has been questioned
in the past (King et al., 2014) and its attribution to the overlying Qasr El-Sagha Formation has
been proposed instead (Gingerich, 1992). (Seiffert et al. 2008) discussed the history of the Birket
Qarun Formation, including the Umm Rigl Member, and supported the original position of the
latter in the Birket Qarun Formation, because it is consistent with previous works and because
of the lack of any distinctive features that would allow its association with the Qasr El-Sagha

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Formation. Herein we follow (Seiffert et al. 2008) in regarding the Umm Rigl Member as part
of the Birket Qarun Formation.
The Birket Qarun Formation contains in total nine fossiliferous localities (Rasmussen et al.,
1992), whereas the Umm Rigl Member specifically includes one of the richest fossil sites, called
Birket Qarun Locality-2 (BQ-2). It was discovered in 2000 and has yielded a very rich fauna
with an age of 37 Ma, representing one of the most diverse vertebrate localities in the Paleogene
of Afro-Arabia (Seiffert et al., 2008). BQ-2 represents the oldest primate-bearing locality in
Egypt, with the coexistence of five primate taxa: Karanisia clarki and Saharagalago misrensis
(Seiffert et al., 2003), Biretia fayumensis and Biretia megalopsis (Seiffert et al., 2006), and
Masradapis tahai (Seiffert et al., 2018). The fauna also contains several fish taxa (El-Sayed et
al., 2020; Murray et al., 2010), snakes (McCartney & Seiffert, 2016), and a very rich mammalian
assemblage, including proboscideans, hyracoids, herodotiines, ptolemaiids, creodonts,
anomaluroid and hystricognathous rodents, chiropterans, and insectivores (Seiffert et al., 2005,
2008; Simmons et al., 2016). The paleoenvironment of the Birket Qarun Formation, including
BQ-2, has traditionally been regarded as marine (Anan and El Shahat, 2014, Beadnell, 1905;
Gingerich, 1992; Kappelman et al., 1992; Van Couvering and Harris, 1991; Wanas, 2008).
Gingerich (1992) suggested that the Birket Qarun Formation represents an offshore barrier bar
complex parallel to the Tethys shoreline. However, Seiffert et al., (2005, 2008) suggested that
BQ-2 represents a fluvial environment. The ichthyofauna and concerations from BQ-2 have
recently been studied in detail and suggest a freshwater environment, with some nearshore
marine influences (El-Sayed et al., 2020; Murray et al., 2010; Anan et al., 2024).

Qasr El-Sagha Formation (Beadnell, 1905)

The Qasr El-Sagha Formation was initially described as the “Qasr El-Sagha Series” by Beadnell
(1905) and later also used by Said (1962). It includes Late Eocene sediments that can be up to
200 m thick (the thickest Eocene formation in the Fayum Depression) and can be divided into
two to four members, depending on the authors (El-Younsy and Salman, 2021; Gingerich, 1992,
1993). As already mentioned above, herein the Umm Rigl Member will be considered as part of
the Birket Qarun Formation (following Seiffert et al., 2006; 2008). Therefore, the lowermost part
of the Qasr El-Sagha Formation is the Harab Member, which consists of 30–40 m of brown
shales (Gingerich, 1992). It is overlain by the Temple Member, which can be up to 80 m thick.

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It consists of thin layers of highly fossiliferous, glauconitic, and limonitic arenaceous limestones,
finely rippled, cross-laminated, and highly bioturbated siliceous sandstone interbedded with
thick laminated siltstones and gypsiferous sandy mudstones (El-Younsy and Salman, 2021). The
upper part of the Qasr El-Sagha Formation is composed of the Dir Abu Lifa Member, which can
be up to 80 m thick. It comprises colorful, cross-stratified sandstones, alternating with siltstone,
silty claystone, and shale, topped by a yellow, fine-to-coarse sandy limestone, with many
gradational to erosional surfaces in between (El-Younsy and Salman, 2021). The Dir Abu Lifa
Member is overlain by the Jebel Qatrani Formation with an erosional contact.
The Upper Eocene Qasr El-Sagha Formation contains eight fossiliferous localities (Rasmussen
et al., 1992; Sileem et al., 2015). The Temple Member has yielded only a few, and mostly marine,
mammals (Bown & Kraus, 1988; Holroyd et al., 1996). Most vertebrate fossils have been
recovered from the Dir Abu Lifa Member (Bown & Kraus, 1988), including fishes, crocodilians,
turtles, snakes, and some mammalian representatives, including sirenians, cetaceans, and
proboscideans. Recent re-evaluation of the sequence stratigraphy of the Qasr El-Sagha
Formation shows that it was deposited in a range of depositional environments, from shallow
marine to fluvial environments (El-Younsy and Salman, 2021).

Jebel Qatrani Formation (Beadnell, 1905)

The Jebel Qatrani Formation (also referred to as Gabal Qatrani Formation) was initially named
“Fluvio-Marine Series” by Beadnell (1905) but was later re-described by Said (1962). It lies
unconformably on the Qasr El-Sagha Formation and is topped by the Widan el Faras Basalt. The
Jebel Qatrani Formation is of Late Eocene to Early Oligocene age (Beadnell, 1905; Bown et al.,
1982; Kappelman et al., 1992; Murray, 2004) and can be up to 300 m thick. The Jebel Qatrani
Formation contains variegated sandstones and mudstones with minor carbonates and chert
pebble conglomerates and pebbly mudstones (Bown et al., 1982). It has been suggested that it
can be subdivided into three members (El-Younsy and Salman, 2021).

The Jebel Qatrani Formation has been divided into two sequences, the “Lower Fossil Wood
Zone” and the “Upper Fossil Wood Zone” (Simons and Wood, 1968). These two zones are
separated by a marker bed, consisting of cliff-forming baryte sandstones. The formation consists
mainly of alluvial sediments of meandering rivers deposited during the Oligocene (Bown et al.,

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1982). Remarkably, it represents the first major terrestrial sedimentation in Egypt since the
Cretaceous (Bown and Kraus, 1988). Some minor developments of shorelines and shallow
marine facies continuing for short periods are also present however, as indicated by thin
sandstones bearing marine mollusks (Bown and Kraus, 1988;). Bown et al., (1982) proposed that
the environment of the Jebel Qatrani Formation, in which several primate taxa coexisted,
represented a sub-tropical to tropical lowland plain, with several large meandering streams and
extensive ponds (contra Kortlandt, 1980). Murray (2004) studied the fish assemblage from the
Jebel Qatrani Formation, which supports the existence of swampy rivers with overgrown banks,
along with abundant vegetation, as also proposed by previous studies (Rasmussen et al., 1987;
Wing and Tiffney, 1982).
Fossil flora evidence outlined that during the Paleogene, a belt of tropical forests stretched along
the coast of the Tethys Ocean, surrounding the Fayum area and its vicinity, whereas some
distance further inland a belt of open woodland or even steppe-like vegetation existed during
large parts of the Paleogene (El Atfy et al., 2021).
The vast majority of the over 100 vertebrate localities of the Fayum Depression are distributed
throughout the Jebel Qatrani Formation (Rasmussen et al., 1992). Of these, eight have produced
almost 90% of the total mammalian remains (Rasmussen et al., 1992; Sileem et al., 2015); these
sites are the quarries A, B, E, G, I, M, V, and Locality 41 (L-41). Site L-41 represents, with an
age of about 34 Ma, the oldest and richest fossiliferous locality in the Jebel Qatrani Formation
(Sallam et al., 2011; Simons, 2008) and has brought to light a very diverse fauna, comprising
fishes (Murray, 2004), birds (Miller et al., 1997; Rasmussen et al., 2001), and a very rich
mammalian fauna. It consists mainly of hyracoids, as well as rodents, primates, creodonts,
macroscelideans, and anthracotheres, indicating a forested environment (Gagnon, 1997).

Widan El Faras Basalt (Bowen and Vondra, 1974)

The Widan El Faras Basalt has unconformably overlain the preceding Jebel Qatrani Formation,
the lower part of which is dated at 31 million years (Fleagle et al., 1986a,b). The basalt occurs
in the Widan El Faras in three main sheets; the lower and upper sheets are amygdaloidal,
vesicular, and intensely altered, whereas the middle sheet is massive, compact, and fresh.
Layering is present in the upper sheet. The three sheets are similar petrologically and are
tholeiitic (Heikal et al., 1983).

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Stratigraphic Levels Yielding Cetacea in the Fayum Depression
Through MUVP work in the region, MUVP work found that the sequence of faunas in the Fayum
Depression overlaps that documented in Cairo and Pakistan, with the Muweilih and Midawara
formations in the Fayum Depression being correlatives of the Mokattam Formation in Cairo and
Habib Rahi and Domanda formations in Pakistan, the Sath El Hadid in the Fayum Depression
being equivalent to Giushi Formation in Cairo and Pir Koh Formation in Pakistan, and the El-
Gharaq and possibly higher formations are being equivalent to Maadi Formation in Cairo and
Drazinda Formation in Pakistan. All these strata provide a coherent stratigraphic base for
understanding change in Eocene cetacean faunas through time.

To conclude, the following intervals in the Fayum Depression have yielded Paleogene cetacean
fossils:

● Midawara Formation, Lutetian: The protocetid whale Phiomicetus anubis (Gohar et al.,
2021), and the remingtonocetid whale, Rayanistes afer (Bebej et al. 2016).
● Sath El Hadid Formation, Bartonian: the basilosaurid whale, Tutcetus rayanensis (Anter
et al., 2023).
● Gehannam Formation, early Priabonian: abundant Basilosaurus isis and Dorudon atrox
basilosaurid whales. In addition to the protocetid whale, Aegicetus gehennae (Gingerich
et al., 2019).
● Birket Qarun and Qasr El-Sagha formations, Priabonian: abundant Basilosaurus isis,
Dorudon atrox, Ancalecetus simonsi, Stromerius nidensis and Masracetus markgrafi
basilosaurid whales (Gingerich, 2010).

Wadi El-Hitan
Wadi El-Hitan (Figure 2.6) is a UNESCO World Heritage Site in the Western Desert of Egypt
famous for the fossils it has produced, including some of the most complete skulls and skeletons
of Eocene whales known anywhere in the world. Most are from the early part of the late Eocene
epoch of Earth's history and lived in the Tethys Sea some 38 to 36 million years before the
present. Basilosaurus isis and Dorudon atrox are the most abundant and best known of archaic
whales. Sea cow skeletons are present as well, as are bones or teeth of bird, crocodiles, turtles,

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bony fishes, and a diversity of sharks and rays. The fossils are preserved in sedimentary strata
that accumulated as sea level rose and fell, producing a succession of nearshore marine
environments. Rocks exposed at the surface, now dissected by erosion, enable the study of
stratigraphic sequences and their architecture.
Wadi El-Hitan or the ‘Valley of Whales’ is an exceptionally rich Eocene paleontological site in
the Western Desert of Egypt. The site has a long history despite the difficult access that desert
isolation posed in the past. The largest and most spectacular fossils found in Wadi El-Hitan are
early sea-living whales.
Over time, rivers draining the continent carried sediment into the sea and moved the shoreline
northward. The Eocene Sea of interest in Wadi El-Hitan was the Tethys Sea, or technically the
Neo-Tethys Sea, a precursor of the modern Mediterranean. There is no evidence that Wadi El-
Hitan was known in antiquity, but this is possible. It lies some 30 kilometers north of the long-
established Darb Al-Rayan caravan route linking the Fayum Oasis, Wadi Rayan, and Bahariya
Oasis in the Western Desert southwest of Fayum (Paprocki, 2019). Wadi El-Hitan is an easy
day’s march north of the main trail. Bedouins pass through with camels seeking forage from time
to time, and it is plausible that early travelers detoured here as well. Exploration of the high
escarpment visible north of Wadi Rayan would bring a wanderer to Wadi El-Hitan with its
conspicuous 5- and 15-meter-long whale skeletons weathering in plain sight on the surface. The
quality and quantity of fossils in Wadi El-Hitan led to its 2005 inscription as a UNESCO World
Heritage Site. The site now has a visitor center, a museum, open-air fossil exhibits, and a graded
road connecting it with the Fayum Oasis, Cairo, and the wider world. As a result, Wadi El-Hitan
has become a popular national and international tourist destination.

Geography of Wadi El-Hitan

Wadi El-Hitan is a broad valley in the Western Desert of Egypt. It lies 70 km west of Fayum and
250 km southwest of Cairo. The valley itself lies in the middle of a larger 17 × 17-kilometer
Wadi El-Hitan protected area. The whole area is a dry, open desert. The Wadi El-Hitan valley is
only a kilometer wide in the southwest but opens to be 8–10 kilometers wide farther to the
northeast. Topographic highs include a northern escarpment that reaches 285 meters or more in
elevation, and southern hill masses stretching between the Visitor’s Center and Garet Gehannam

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that reach 225 meters in elevation. Drainage is internal, to points on the valley floor that are
barely 50 meters above sea level. Vegetation is sparse, and wildlife is rare.

Paleogeography of Wadi El-Hitan

The paleogeography of Wadi El-Hitan is, in some ways, more interesting than the modern
geography. Exploration for petroleum means seismic surveys have crisscrossed northern Egypt,
and many oil wells have been drilled. Combined, they enable reconstruction of the late Eocene
paleogeography Structurally, the Precambrian basement underlying the desert is a series of
parallel northeast-southwest trending faults that separate elevated horsts from intervening
grabens. This controlled erosion and deposition during the Eocene (Dolson et al., 2002). Wadi
El-Hitan is located on the northwestern margin of one of the emergent horsts, which was exposed
as land in the late Eocene. At the time erosion transported sediment to the northwest across Wadi
El-Hitan, moving it across a shallow-marine shelf from the raised area of the horst toward deeper
water in the adjacent graben. Fossils indicate that the environment was tropical and rich in marine
life. Rare fossils of land mammals show that land lay nearby)Figure 2.7-8)

Geology of Wadi El-Hitan

The first geological investigation of Wadi El-Hitan was by Hugh J. L. Beadnell (1874–1944)
and a team from the Egyptian Geological Survey. They worked during the winter field season of
1902–1903, carrying out a general survey of the topography and geology of the desert north and
west of the Fayum oasis (Beadnell, 1905; discussed above). Topography was important because
one question of interest was the feasibility of storing Nile floodwaters in a Lower Egypt reservoir
closer to Cairo and closer to the Nile Delta. This would supplement newly constructed storage
behind the original Aswan Dam in Upper Egypt. Geology was important because of the potential
for economically valuable mineral resources. Neither proved worthy of development, and the
lasting legacy of Beadnell’s fieldwork was the discovery of an unexpected richness of fossils,
notably Eocene whales (discussed below under Vertebrate Paleontology).

Vertebrate Paleontology of Wadi El-Hitan

Vertebrate fossils have received the most attention in Wadi El-Hitan because of their abundance
and exceptional preservation. Virtually all are early Priabonian late Eocene in age. Older

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Lutetian and Bartonian middle Eocene whales and other marine vertebrates are found farther
south in the lower strata of Wadi El-Rayan. Younger middle and late Priabonian late Eocene
whales and other marine vertebrates are found farther east in the escarpments north of Birket
Qarun. Thus, Wadi El-Hitan lies at the heart of an exceptional stratigraphic record of marine life
in the Eocene Tethys Sea.

Chondrichthyes: Sharks and rays are the most abundant vertebrate fossils found in Wadi El-
Hitan. Many specimens described by Case and Cappetta (1990) came from unknown localities
and uncertain stratigraphic intervals. Underwood et al. (2011) published a more comprehensive
overview of sharks and rays in Wadi El-Hitan.

Osteichthyes: Two studies have been published on the bony fishes of Wadi El-Hitan. Fierstine
and Gingerich (2009) described the rostrum of a previously known billfish, Xiphiorhynchus
aegyptiacus. El-Sayed et al. (2017) described a new genus and species of marine catfish,
Qarmoutus hitanensis.

Testudines: Two studies have been published on the turtles of Wadi El-Hitan. Wood et al. (1996)
placed the leatherback turtle Psephophorus eocaenus in a new genus Egyptemys. Cherney et al.
(2020) included a specimen identified as Cordichelys aff. C. antiqua.

Crocodylia: The only crocodilian described from Wadi El-Hitan to date is the narrow-snouted
gavialid Tomistoma kerunense (Andrews, 1905). Paratomistoma courti of Brochu and Gingerich
(2000) is a junior synonym of Tomistoma. A small broad-snouted crocodilian is also present in
the fauna.

Aves: A single bird, the early pelican Eopelicanus aegyptiacus is known from the Birket Qarun
Formation of Wadi El-Hitan (El Adli et al. 2021).

Mammalia:
Sirenia: The first sirenian described from Wadi El-Hitan was an edentulous mandible collected
by a Yale field party in 1964–1965. This was identified by Domning et al. (1982) as Protosiren

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sp. but is now recognized as a specimen of Eotheroides. Later, Domning and Gingerich (1994)
named the new species Protosiren smithae from Wadi El-Hitan, and Zalmout and Gingerich
(2012) named two new species of Eotheroides, E. clavigerum and E. sandersi. All three of these
species are based on well-preserved skulls and partial skeletons.

Cetacea: Andrews's (1906) description of the archaeocete Prozeuglodon atrox (now Dorudon
atrox) from Wadi El-Hitan was mentioned in the Introduction chapter, as was a collection of a
Basilosaurus isis skull by the University of California. Kellogg (1936) recognized that teeth in
the type of Prozeuglodon atrox are deciduous, considered P. atrox to be immature
representatives of P. isis, and synonymized the two species as Prozeuglodon isis — an
interpretation endorsed by Barnes and Mitchell (1978). However, field research in Wadi El-Hitan
in the 1980s showed that three forms are common:
(1) 15-meter-long skeletons of mature Basilosaurus isis;
(2) 5-meter-long skeletons of mature Dorudon atrox; and
(3) even smaller skeletons of immature Dorudon atrox (Gingerich et al., 1990).
Immature Dorudon are most easily recognized by their smaller size, retention of deciduous teeth,
and presence of unfused endplates on vertebrae (Uhen, 2004). (Figure 2.19) shows the great
difference in size of adult B. isis and D. atrox. Gingerich and Uhen (1996) described a third
basilosaurid from Wadi El-Hitan, Ancalecetus simonsi, that is distinctive in having
anteroposteriorly narrow scapulae; limited mobility of the shoulder joint; fusion of the humerus,
ulna, and radius at the elbow; and relatively small carpal bones in the wrist.
Gingerich (2007) referred to several large but relatively short lumbar vertebrae as a fourth
basilosaurid Masracetus markgrafi. Both of these species are basilosaurids found in the Birket
Qarun Formation. Moreover, Anter et al. (2023) recorded the basilosaurid Tutcetus rayanensis
near the Wadi El-Hitan area. Furthermore, two specimens of a new late-surviving protocetid,
Aegicetus gehennae, were found in Wadi El-Hitan in 2007. These were found in the Gehannam
Formation (Gingerich et al., 2019). Finally, a Gehannam Formation Basilosaurus isis skeleton
excavated in 2010 had stomach contents showing that Basilosaurus preyed on juvenile Dorudon
atrox (Voss et al., 2019).

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Proboscidea: Most fossil mammals found in Wadi El-Hitan are marine sea cows and cetaceans,
but there are also rare proboscideans. The first was a partially articulated lumbus, with pelves,
of a pygmy-hippo-sized semiaquatic Moeritherium lyonsi (Gingerich, 1992). The specimen was
found in the marine strata of the Birket Qarun Formation. Isolation in an offshore open-water
environment and limited articulation suggests that the Moeritherium may have been a victim of
archaeocete predation.
The teeth and occasional limb bones of Moeritherium lyonsi and the larger elephant-sized
Barytherium grave are also found in Wadi El-Hitan in the Priabonian P-2 incised valley fill
shown on the map in. The P-2 incised valley fill separates the lower and middle parts of the Qasr
El-Sagha Formation, and the proboscidean elements are reworked from the lower Qasr El-Sagha
Formation or even from the underlying Birket Qarun Formation. Consistent with reworking,
these are always isolated elements.

Scientific and Cultural Value

Wadi El-Hitan is important for three reasons. First, Wadi El-Hitan has value for the fossils it
produced and continues to produce, including some of the most complete skeletons of archaic
whales known anywhere. Whales are abundant and well preserved, but their value is enhanced
by entombment in strata with the animals that live around them. Invertebrates are abundant and
diverse, including many kinds of mollusks, sea urchins, crabs, and corals. Fish include a rich and
complex fauna of marine sharks, rays, and bony fishes. Reptiles include crocodilians and turtles.
One bird is known, and there were undoubtedly more. Mammals include primitive sea cows and
proboscideans in addition to whales. Much is known, but there is still much to learn about
biodiversity and ecological relationships in the Eocene of Wadi El-Hitan.
Second, Wadi El-Hitan has value for observing and understanding the development of complex
architectures in geological strata that produce and trap petroleum in marginal marine
environments. Many studies of ‘sea level sequence stratigraphy’ are based on the interpretation
of seismic profiles of buried strata. In Wadi El-Hitan the strata are at the surface, dissected by
wind and weather, and an investigator can study depositional sequences, literally, by walking
through them. This is a big advantage for seeing and sampling the strata, for identifying potential
petroleum sources, for integrating stratigraphic sequences with the micro- and macrofossils they
contain, and for interpreting the rise and fall of sea level.

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Finally, Wadi El-Hitan has value for public education. Eocene fossils are displayed where they
were found, in a site museum with interpretive exhibits, but also in situ in the open air.
Scientists work in the Wadi El-Hitan “Valley of Whales” to improve our geoheritage — our
understanding of life and environments of the geological past. Schools and colleges bring
students to Wadi El-Hitan to give them a deep-time perspective on the natural world. Tourists
come to
Wadi El-Hitan from Egypt and many other countries to visit the World Heritage Site. Wadi El-
Hitan is an open book in the sense that fossils and the surrounding strata lie fully exposed in the
desert. UNESCO’s World Heritage designation acknowledges and endorses the scientific and
educational importance of the site. It requires that the Government of Egypt provide a corps of
guides and staff who live and work at the site, where the government added an interpretive
museum. The site deserves protection in perpetuity, and this seems assured by the UNESCO
endorsement and Egypt’s ongoing commitment.

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Figure 2.1. Panorama and general view showing some geomorphological settings of Wadi El-
Hitan. © MUVP.

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 Figure 2.2. Isolate hill to the North of Wadi El-Hitan (from Abu El-kheir, 2010).

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Figure 2.3. General view of some dunes (Ghard Gehannam) at the east of Garet Gehannam
(from Abu El-kheir, 2010).

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Figure 2.4. Rocky sandy bajada in the northeast of Wadi El Hitan (from Abu El-kheir, 2010)

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Figure 2.5. A. Location map of the Fayum Depression, Egypt. B. Geological map of Fayum and
Wadi El-Hitan. C. Zoomed portion of the Camp area in Wadi El-Hitan (compiled and modified
after El-Sayed, 2017).

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Figure 2.6. Correlation of stratigraphic units of the Cairo and Fayum areas (following
Schweinfurth, 1883; Said, 1990; Strougo and Haggag, 1984; Strougo 1985a,b; Gingerich, 1992;
and Strougo, 1992; Sieffert et al., 2006; 2008; Zalmout and Gingerich, 2012).

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 Figure 2.7. Wadi El-Hitan from 40 million years ago.

Figure 2.8. wadi El-Hitan now.

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Figure 2.9. Skeletons and reconstructions of Wadi El-Hitan archaeocetes Basilosaurus isis and
Dorudon atrox mounted in the University of Michigan Natural History Museum. A, B (upper).
B. isis, Egyptian Geological Museum specimen 42195, in left lateral view. A, B (lower). D.
atrox, composite Egyptian Geological Museum specimen 42183 with additions from University
of Michigan specimens, left lateral view. Note the retention of hind limbs in both species
(arrows). In life, the left and right pelvic bones remained attached at a midline symphysis, but
these were no longer connected to the vertebral column. Figure reproduced from Voss et al.
(2019).

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 Chapter Three
METHODOLOGY

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 METHODOLOGY

The UNESCO World Heritage site Wadi El-Hitan, located north of the Western
Desert of Egypt, has yielded an exceptional collection of Eocene vertebrates, particularly the
fossil whales from which it gets its name. A field exploration effort by the Mansoura University
Vertebrate Paleontology Center (MUVP) in collaboration with the Egyptian Environmental
Affairs Agency (EEAA) has led to the study and preparation of well-preserved remains of
Dorudon atrox at the Open-Air Museum of Wadi El-Hitan world heritage site, Fayum
Depression (Figure 3.1).

The fossil material described here is part of the activities of the Mansoura University Vertebrate
Paleontology Center (MUVP). The collection of the necessary materials for this project required
fieldwork and laboratory work. During a trip organized by MUVP, under the supervision of Prof.
Hesham Sallam, the team spent 5 days at a training camp in Wadi El-Hitan in collaboration with
EEAA. The main objective of the field trip was to learn about the stratigraphy and vertebrate
paleontology of the Wadi El-Hitan area. During the trip, the team received training on how to
excavate and prepare the skeletons of Dorudon atrox and other remains .

1- Techniques in the Field at Wadi El-Hitan

There are various methods of collecting fossils, each adapted to specific situations. Ensuring the
safe collection of specimens is essential, and the specimens collected in the field were later
prepared in the MUVP laboratory. Prof. Sallam made collecting decisions to acquire stable
specimens with the maximum amount of preserved information in mind.

No fossils were collected without good data, which typically includes GPS coordinates, digital
photographs of both the specimen and the site, as well as field notes about the stratigraphic
placement and taphonomic information.

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A- Prospecting in the Field

While prospecting (Figure 3.2), we frequently paused to observe our surroundings, noting
landmarks and familiarizing ourselves with our route. Using a loupe, we carefully examined
potential bone fragments, distinguishing them from rocks by looking for specific grain and
cancellous patterns.

Upon finding a bone fragment, we thoroughly inspected the area, often getting on our hands and
knees to avoid missing any small pieces. Tracing the source of the fragment uphill was essential,
as many fragments originated from higher locations. We carefully circled the site from various
angles without disturbing the surrounding rock.

B- Preliminary Excavation

Using an ice pick, we gently removed surface dirt to assess whether the find was part of a
complete bone or skeleton. We refrained from digging deeply at this stage to avoid damaging
the specimen. Confident in our discovery, we photographed the site and recorded GPS
coordinates, using landmarks to ensure easy relocation. Temporary markers were placed but later
removed to leave no trace (Figure 3.3-4).

C- Collaborative Assessment

After documenting our findings, we discussed the specimens with the Sallam Lab team. Given
the limitations of our expedition, we prioritized the excavation of the highest-quality fossils.

D- Detailed Documentation

We began by photographing the exposed fossils, using markers to highlight fragments, and
taking detailed notes on the location, appearance, condition, and orientation. GPS readings were
recorded, and a field number was assigned to each specimen.

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E- Surface Collection

Nearby surface fragments, likely belonging to the same specimen, were carefully collected,
wrapped in tissue paper, and placed in bags with corresponding field number slips. This ensured
that we could accurately reconstruct the specimen's context (Figure 3.5).

F- Excavation of the Specimen

With the necessary tools on hand, we outlined the fossil area, avoiding full exposure as the
surrounding matrix supported the bones. We revealed only the top and edges to understand the
fossil's extent. Digging tools such as points, chisels, and rock hammers were used alongside
consolidants such as Paraloid B-72 (ethyl methacrylate co-polymer, formerly called Acryloid)
for fragile bones.(Figure 3.6)

G- Observations of the Matrix

We carefully examined the matrix, noting any taphonomic clues, such as microfossils or plant
remains, which provided valuable insights into the site's history.(Figure 3.7)

H- Bone Consolidation

For fragile or crumbly bones, we applied a thin consolidant (Paraloid B-72 (ethyl methacrylate
co-polymer, formerly called Acryloid)) using a pipette, ensuring minimal use in the field to avoid
later removal complications.

I- Single Bone Extraction

For small, sturdy bones, we excavated them completely, wrapped them in tissue, and placed
them in labeled bags. Medium-sized bones were additionally wrapped in aluminum foil for extra
support.(Figure 3.8)

J- Excavation and Preparation

We defined the fossil boundaries, excavating around and below it, creating a pedestal to support
the specimen. For fragile matrices, we began jacketing immediately; otherwise, we continued
excavation to ensure the fossil’s stability.(Figure 3.9)

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K- Applying the Jacket

Using tissue paper as a barrier, we molded it snugly over the fossil to prevent plaster from
sticking. For smaller fossils, we applied plaster bandages, overlapping them for strength. For
larger fossils, we used burlap strips soaked in plaster, applying multiple layers for adequate
protection.(Figure 3.10-11)

L- Finalizing the Jacket

Once the jacket hardened, we excavated beneath the pedestal and carefully flipped the fossil to
prevent matrix loss. We then capped the underside with additional plaster and burlap layers. The
jacket was labeled with the field number, identification, and orientation details.(Figure 3.12)

M- Labeling

Each specimen and plaster jacket were labeled with a field number, and detailed notes were
taken, including GPS coordinates, stratigraphic data, and orientation. Photographs and
comprehensive field notes are essential for later lab work. Specimens from Wadi El-Hitan are
then transported to the lab, where careful extraction from the matrix begins, revealing the ancient
history of these remarkable whale fossils.(Figure 3.13)

2- Laboratory Preparation at MUVP lab

A- Cleaning and Preparation

In the lab, we first unwrapped and unjacketed the specimens and then meticulously removed any
remaining matrix using precision tools, such as air scribes and fine brushes, taking care not to
damage the fossil.(Figure 3.15)

B- Consolidation, Reconstruction, and Restoration

We applied consolidants (Paraloid B-72 (ethyl methacrylate co-polymer, formerly called

Acryloid)) as needed to stabilize fragile areas, ensuring thorough impregnation and subsequent
hardening. Missing parts of the fossil can be reconstructed using various materials like plaster,
resin, or epoxy putty. This helps in creating a complete specimen for study or display. Damaged

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parts of fossils are carefully restored to their original state. This requires detailed knowledge of
the anatomy and structure of the vertebrates being worked on.

C- Computed Tomography (CT) CT Scanning

Utilizing CT scans, we analyzed the internal structure of the fossils, followed by segmentation
for detailed studies. We also performed 3D laser scanning to create digital models for further
examination. The specimen was entirely subjected to CT imaging to supplement observations of
the morphology on the external surface. The specimen was scanned helically using a Philips
Incisive CT Scanner at Al-Mawji Radiology Center in Mansoura, Egypt. The overview scans
were made with a field of view of 500 mm, a tube voltage of 140 kV, a tube current of 272 mA,
and a slice thickness of 0.8 mm. The scanning resulted in 1949 slices with a pixel matrix of 512
× 512, a field of view of 500 mm, and a 0.2 mm interval between slices. Observations on image
slices and three-dimensional (3D) visualization were done using the software package Amira
4.1.2 (Figure 3.16).

D- 3D Laser Scanning

The 3D laser scanning of fossils was performed with the handheld Micro Leo 3D scanner with a
3D point accuracy of up to 0.1 mm and a 3D resolution of up to 0.5 mm, while its 3D accuracy
over 1 m is as small as 0.03%. Its furthest range of linear field of view is 838 mm × 488 mm,
and its largest angular field of view is 38.5° × 23°. The scanned data from the Artec Leo was
transformed into the post-processing software Artec Studio 17 (Figure 3.17).

E- 2D imaging and Illustrations

Nikon D3100 and D750 cameras with 60 mm and 105 mm lenses were used to photograph the
fossil element at MUVP. Adobe Photoshop (CS 6) and Adobe Illustrator (CS 6) were used to
create and edit the photographs to generate illustrations used herein and for the resultant
manuscripts (Figure 3.18).

F- Microscopy

Each specimen was studied under high-profile stereozoom microscopes at MUVP, allowing for
detailed morphological analysis(Figure 3.19-20).

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G- Documentation

Throughout the preparation and analysis process, we documented each step with photographs
and detailed records of the fossil's condition and preparation stages(Figure 3.21).

H- Mechanical Preparation

This involves the physical removal of the matrix using tools such as air scribes, pneumatic tools,
pin vises, dental picks, and small chisels. Fine preparation was done under the sterozoom
microscopes at MUVP. In some cases, the CT was used as a guide for this mechanical
preparation.

I- Anatomical description
The anatomical nomenclature and description used throughout the thesis follow Kellogg (1936),
Uhen (2004), Geisler and Luo (1998), Luo and Gingerich (1999), and Geisler et al. (2005). Teeth
are referred to as abbreviated capital letters throughout the thesis, with incisors abbreviated as
(I), canines as (C), premolars as (P), and molars as (M). Maxillary teeth are indicated by
superscript numbers (e.g., M1 refereeing the first upper molar), and mandibular teeth by subscript
numbers (e.g., M1 refereeing the first lower molar).

J- Measurements
Measuring variable dimensions of cranial and postcranial elements has been employed to
describe the morphological variation in these primitive whales. Postcranial elements with
significant variable attributes include vertebrae length and height, and lengths of rib fragments.
These were used to show variation among the different groups of the Eocene protocetids around
the world. The skull was measured in detail for comparative purposes. Most measurements of
both cranial and postcranial elements are following the protocol of Geisler et al. (2005). A digital
caliper was used for the linear dimensions. Circumferences, arc lengths, and perimeters were all
measured by using a measuring tape. During all measuring events, digital calipers and measuring
tape have not been replaced. The frontal shield breadth has been measured based on the
assumption that the skull is essentially symmetrical; so, we measured the complete left side and
doubled that value. In measuring transverse diameters of the sixth thoracic vertebra,
measurements of the anterior face of the centrum were taken at the approximate centers of the
demifacets, and those of the posterior face were taken at the lateral margins of the demifacets.

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In teeth measurements, the anteroposterior length of the crown was taken along the base, and the
transverse diameter was measured at the base perpendicular to the long axis of the crown. The
vertical dimension of the crown was measured from the base to the tip of the apical cusp. In the
descriptions and comparisons of the new specimens’ vertebral elements, the methods of utilizing
length (L)/width (W) ratios of vertebral centra presented by Geisler et al. (2005) have been
employed. Ratios of the dimensions of the centra were determined by dividing the width (W) of
the anterior face by the anteroposterior length of the centrum. Higher values reflect a relatively
short centrum and lower values indicate more elongated ones.

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Figure 3.1. Prof. Hesham Sallam explaining the anatomical features of Dorudon atrox at the
Open-Air Museum of Wadi El-Hitan.

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 Figure 3.2. Principal prospecting for fossils at the Wadi El-Hitan.

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 Figure 3.3. Transporting tools needed for fossil collecting.

Figure 3.4. Transporting equipments to the excavation site.

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 Figure 3.5. Collecting small-sized fossils using sieves.

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 Figure 3.6. Detecting the external boundaries of the specimen.

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 Figure 3.7. Using chisels and rock hammers for the fossil excavation.

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 Figure 3.8. Single bone fragment found during prospecting.

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 Figure 3.9. A broken rib on the surface, with another still in situ.

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 Figure 3.10. Partial skeleton of Dorudon atrox specimen ready for jacketing.

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 Figure 3.11. The trenching process in preparation for jacketing.

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Figure 3.12. Preparing the remains of Dorudon atrox at the Open-Air Museum, Wadi El-Hitan.

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 Figure 3.13. MUVP team at the Dorudon atrox site at the end of the fieldwork.

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 Figure 3.14. Preparing a layer of gypsum for jacketing.

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 Figure 3.15. Cleaning and lab preparing the specimen.

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 Figure 3.16. CT scanning at Al-Mawji Radiology Center, Mansoura.

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 Figure 3.17. 3D laser scanning with the Leo Micro 3D scanner.

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 Figure 3.18. Photographing the specimen at MUVP.

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 Figure 3.19. Fine preparation using an air scribe pen under the stereozoom microscope at
MUVP.

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 Figure 3.20. Specimen preparation under the stereozoom microscope at MUVP.

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 Figure 3.21. Investigating the specimen through measurements and descriptions at MUVP.

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 Chapter Four
SYSTEMATIC PALEONTOLOGY

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 SYSTEMATIC PALEONTOLOGY

Order CETACEA Brisson, 1762

Suborder ARCHAEOCETI Flower, 1883

Archaoceti are early, primitive cetaceans that retain generalized mammalian skulls with nares on
the rostrum, cheek teeth with complex crowns and multiple roots, and periotics attached to
surrounding bones. Skeletons range from semiaquatic to fully aquatic, with fore- and hindlimbs
variably modified for swimming. The elbow joint is generally mobile. Early semiaquatic forms
have skeletons with double-pulley astragali and other features indicating evolutionary derivation
from early Artiodactyla (Gingerich et al., 2001; 2003; 2019).

Family BASILOSAURIDAE Cope, 1868

Basilosauridae are generally later, more derived, and fully aquatic archaeocetes with only two
molar teeth in the maxilla. Basilosaurids generally lack evidence of a distinct sacrum in the
vertebral column and have innominates of reduced size and altered proportions. It is useful to
separate Basilosaurus with extreme vertebral elongation from more typical basilosaurids. The
former are sometimes classified in Basilosaurinae and the latter in Dorudontinae. However, the
relative size of carpal bones and other differences cut across this simple dichotomy, and a better
understanding of basilosaurid morphology in more taxa will be required before a reliable
subdivision is possible (Uhen, 2004; 2010; Anter et al., 2023).

Subfamily DORUDONTINAE Miller, 1924

Genus: DORUDON Gibbes, 1845.

Species: Dorudon atrox (Andrews, 1906).

Type locality: Wadi EI-Hitan World Heritage Site. The type’s stratigraphic position of the
holotype is the Birket Qarun Formation.

Age and distribution: Late Bartonian (late middle Eocene) and early Priabonian (early late
Eocene) of the Gehannam and Birket Qarun formations, northern Fayum, Egypt.

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Diagnosis: Dorudon atrox (Figure 4.1)can be distinguished from all other archaeocetes by its
unique conformation of cranial bones in the posterior narial region. In Dorudon atrox, the
posterior nasals are separated by a long, thin process of the frontals. In addition, the medial edges
of the posterior maxillae contact the lateral edges of the nasals. Both species of Basilosaurus
have small narial processes of the frontals and the medial maxillae touch the nasals only.
Dorudon atrox is distinctive as a medium-sized Fayum archaeocete with blocky posterior
thoracic, lumbar, and anterior caudal vertebrae. Posterior thoracic and anterior lumbar vertebrae
have centra that are approximately equal in length to their height and width. Dorudon resembles
Basilosaurus and differs from Saghacetus and Stromerius in having a larger number of thoracic
vertebrae (17 in Dorudon).

Skull

Premaxilla: The premaxilla (PMx in Figure 4.3) forms the anterior portion of the rostrum
dorsally and ventrally, as well as the anterior, lateral, and part of the dorsal margins of the
external bony nares. Laterally, it articulates exclusively with the maxilla. Anterior to the narial
fossa, the premaxillae are facing each other along the midline. In the dorsal view, the posterior
portion (narial process) of the premaxilla articulates medially with the nasal. In the mesorostral
region, the premaxilla is partially covered by the vomer.

The naso-premaxillary (Na-PMx) sutures are straight and diverge posterolaterally. Laterally, the
narial process contacts the maxilla forming the posterior portion of the external maxillo-
premaxillary (mx-pmx) suture. The alveoli are roughly oriented ventrolaterally, with the alveolus
for I1 more anteriorly oriented than those for I2 and I3. Moreover, both I1-I2 and I2-I3 diastemata
bear embrasure pits, which receive the crown of the posterior lower incisors (i2-i3). Thereby,
when the jaws are closed the lower incisors are anterior to their corresponding upper incisors. A
third embrasure pit is located posterior to I3 and receives the crown of the lower canine. The
three embrasure pits cited above are located lateral to the dental row, open laterally, and are
visible in lateral and ventral view. The depth and size of the embrasure pits increase posteriorly.

Maxilla: The maxilla (Mx in Figure 4.4) contributes significantly to the posterolateral wall of
the rostrum. Medially, the maxilla articulates with both premaxilla and nasal. Ventrally, it mainly
participates with the palatine in the posterior half of the palate. Posteriorly, the maxilla articulates

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principally with the frontal, but it also contacts both the lacrimal and the jugal. It contributes to
the anterior wall of the orbit. As a consequence of the torsion of the rostrum, the right maxilla is
slightly transversely wider than the left maxilla, anterior to and at the level of the external bony
nares

In the dorsal view, the maxilla contacts the premaxilla medially. The dorsal portion of the mx-
pmx suture has already been described above. The mx-pmx suture meets posteriorly the
nasomaxillary (na-mx) suture, which is parallel to the midline for almost its entire length. Indeed.
Here, the maxilla forms a narrow, but conspicuous jugular process. The alveolar process lies on
the lateral portion of the maxillary palatal surface. It bears the alveoli for six teeth (C1-M1). On
the one hand, the two anterior alveoli (for C1 and P1) are aligned with the incisor alveoli. On the
other hand, the four posterior alveoli (for P2-M1) are aligned in a row, with the right and left rows
diverging posteriorly

Only the anterior maxillary alveoli are separated by three diastemata (respectively located
between C1 and P1, P1 and P2, and P2 and P3). As for the premaxilla, the anterior diastemata are
longer on the right side of the skull, a condition that results from the asymmetry of the rostrum.
Each diastema is excavated by a conical embrasure pit, receiving the crown of P 1, P2, and P3,
respectively. The embrasure pit housing P2 (in the P1-P2 diastema) is the deepest, being followed
in depth by the pit for p1 (in the C1-P1 diastema).

Vomer: The vomer (v in Figure 4.4) covers most of the ventral surface of the presphenoid, as
it does in other basilosaurids, but little of it is exposed. It contacts the medial border of the medial
lamina of the pterygoid.

Nasal: The Nasal (na in Figure 4.3) The nasal is a thin and paired bone, that forms the median
portion of the facial region and participates in the roof of the nasal cavity. Laterally, the nasal
articulates with the premaxilla, maxilla, and frontal. Medially, both nasals meet at midline for
almost their entire length and are only separated posteriorly by the anterior (or nasal) process of
the frontal.

Jugal: The Jugal (j in Figure 4.4) The right jugal is the only preserved from both. The jugal is
a paired bone, which contributes exclusively to the zygomatic arch. As in all other basilosaurids,

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it is straight, long, and transversely compressed on most of its length. Anteriorly, the jugal
contacts the maxilla and lacrimal. Posteriorly, it does not articulate with the squamosal

Lacrimal: The lacrimal is one of the smallest bones of the skull. It is paired and located at the
posterolateral corner of the rostrum. Dorsally, the lacrimal articulates with the frontal; anteriorly
and ventrally, it articulates with the maxilla; and posteroventrally, it contacts the jugal. The
lacrimal also contributes to the orbit wall and the lacrimal and infraorbital canals. The external
surface of the lacrimal is mainly exposed in lateral view but is also visible in dorsal view.

Frontal: The Frontal (fr in Figure 4.3) The frontal is a paired bone. It is the main element of
the supraorbital shield and forms most of the roof and medial wall of the orbit. Both frontals
meet at midline and the interfrontal suture is easily distinguishable in the supraorbital region.
Dorsally, the frontal articulates posteriorly with the parietal and anteriorly with the nasal and the
maxilla. In the orbit, it contacts the lacrimal, maxilla, palatine, and orbitosphenoid.
Posteroventrally, the frontal articulates with the alisphenoid. In the dorsal view, the frontals
extend anteromedially, forming a triangular anterior process that separates the posterior ends of
the nasals. This anterior process is anteroposteriorly longer than wide.

Palatine: The palatines (pal in Figure 4.4) are paired elements forming the posterior portion of
the palate, extending from the maxillae in front to the pterygoids posteriorly. The two palatines
join at the midline and their ventral surface forms a keel that narrows posteriorly. This is part of
the posterionnost extension of the hard palate. The palatines rise dorsally and laterally along a
surface that is folded inward to form a concave surface for muscle attachment. Internal surfaces
of the palatines form the ventral and lateral walls of the narial passage.

Pterygoid: The remaining portion of the palate is formed by pterygoids. These too are paired
elements. They articulate anteriorly with the posterior parts of the palatines. The pterygoids meet
along the midline ventrally to form ventral and lateral walls of the posteriormost extension of
the narial passages. The pterygoids have several extensions and surfaces that project in different
directions

Orbitosphenoid: The orbitosphenoid (or in Figure 4.4) is a paired bone. It is a small bone
exposed on the medial wall of the orbit. In this region, the orbitosphenoid covers part of the
frontal and participates in the medial wall of the optic infundibulum. Anterolaterally, the

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orbitosphenoid forms the optic foramen and part of the path for the optic nerve and the
ophthalmic artery. Posteromedially, the orbitosphenoid participates in the medial wall of the
sphenorbital fissure. At this level, it contacts the palatine anteroventrally and basiphenoid
posteromedially.

Basisphenoid: The basisphenoid generally has little ventral exposure on the basicranium
because it is completely or almost completely covered by vomer and pterygoids. The
basisphenoid-basioccipital suture is straight and runs across the basicranium perpendicular to the
midline.

Alisphenoid: The alisphenoid (al in Figure 4.4) forms the ventral portion of the lateral wall of
the mesocranium. It is discussed here as an extension of the basisphenoid. The alisphenoid covers
the course of the ophthalmic and maxillary divisions of the trigeminal nerve (V, and V2), beneath
which is the presphenoid. A small piece of the posteroventral alisphenoid is present on each side
where it contacts the palatine anteroventrally, the pterygoid posteroventrally, and the squamosal
posteriorly.

Parietal: The parietals (pr in Figure 4.3) make up the lateral sides of the posterior portion of
the skull. Only the posterior parts of each parietal are preserved here, where they are sutured to
the squamosal. The parietal-squamosal suture runs diagonally dorsally and caudally across the
lateral wall of the braincase to a point where it turns sharply posteriorly and intersects the nuchal
crest (ncr)

Squamosal: The squamosal (sq in Figure 4.3) forms the posterolateral wall of the side of the
cranium. The squamosal articulates anteriorly along its dorsal edge with the parietal and on its
anteroventral edge meets the alisphenoid as described above. There is also a contact with the
pterygoid, enclosing the foramen ovale that carries the mandibular branch of the trigeminal nerve
(V3) into the temporal fossa. On the ventral side, the squamosal has a complex relationship with
the periotic. The squamosal projects laterally from the side of the skull and anteriorly to form
the zygomatic process of the squamosal. Once this zygomatic process turns anteriorly it becomes
laterally compressed and parallels the dorsal surface of the jugal. The two bones remained
separate along their entire parallel course

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Basioccipital: The basioccipital forms the central base of the posterior basicranium. This
articulates anteriorly with the basisphenoid and posteriorly with exoccipitals. The basioccipital
and exoccipitals are generally co-ossified in older juvenile and adult basilosaurids, but these are
readily distinguished in young individuals. The caudalmost projection of the basioccipital
extends between the occipital condyles and forms the ventral floor of the foramen magnum (frn).
This area is also known as the intercondyloid notch. Internally, the basioccipital forms the
posterior floor of the braincase.

Supraoccipital: The supraoccipital is a singular midline bone that forms the posterior portion
of the braincase and the posterior surface of the skull. Most of the supraoccipital in the specimen
has been weathered away, but the ventral-most portion remains. The supraoccipital articulates
ventrolaterally with the exoccipitals, and the suture between these elements runs diagonally from
the foramen magnum toward the ventral borders of each nuchal crest. The ventralmost projection
of the supraoccipital forms the dorsal border of the foramen magnum. The posterior surface of
the supraoccipital is concave, with the lateral edges flaring out and back to form prominent
nuchal crests. In addition, the ventral extension of the supraoccipital flares outward from the
center of the bone to the edge of the foramen magnum.

Exoccipital: The exoccipitals (eoc in Figure 4.3) are paired bones that form the ventral part of
the posterior surface of the cranium. The exoccipitals articulate dorsally with the supraoccipital
along the diagonal sutures mentioned above. The anterior surface of the exoccipitals articulates
medially with the parietals and laterally with the squamosals.

The occipital condyles (oc) arise from the medial edges of the exoccipitals. They extend ventrally
to the suture with the basioccipital, forming the intercondylar notch mentioned above. The
occipital condyles articulate with the vertebral column. Lateral and ventral to the occipital
condyles, two processes of the exoccipital form the borders of the jugular notch (jn), which lies
between them.

Ectotympanic: The tympanic (ectotympanic) auditory bulla of Dorudon atrox (Figure 4.6) is
a large ovate structure like that of other advanced archaeocetes (Kellogg, 1936), including
Zygorhiza kochii (Lancaster, 1990). It is composed of unusually dense compact bone. The right
bulla is well preserved in Dorudon atrox. It has a convex ventral surface and a deeply excavated

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dorsal surface (tympanic cavity). The tympanic cavity is an anteroposteriorly elongated space
opening into the middle ear above. The tympanic cavity is widest anteriorly and narrows
posteriorly. There is a shallow depression on the posteroventral part of the external surface of
the bulla that separates the involucrum on the medial side from the outer lip. The latter bears a
raised circular eminence curving smoothly into the sigmoid process (sp). The sigmoid process
curves posteriorly where it rises as a distinct process and normally would project dorsally above
the rest of the bulla, however, most of the sigmoid process is broken and missing. The posterior
edge of the sigmoid process normally curls backward, enclosing a space between itself and the
body of the auditory bulla. Much of the malleus is present, attached to the gracile or gonial
process. There is a distinct sulcus for the chorda tympani anterior to the gracile process.

Mandible
The left and right mandibles in Doruodon atrox are each composed of a single bone, the dentary
(Figures 4.7-8). In: Doruodon atrox the left and right dentary are complete from the alveolus for
I1 to the mandibular condyle (cd). The mandibular symphysis (sym) between these extends
posteriorly to a point below the posterior edges of the alveoli for P2. This symphysis is unfused.
Rami of the left and right dentaries parallel each other along the symphysis and then diverge
posteriorly as far as the end of the molar series. Posterior to the molar series the rami are again
more parallel. Much of the medial surface of the dentary is gently concave, while the lateral
surface is slightly convex. The lateral surface bears a series of small mental foramina (mf). At
the anterior end of the dentary these are close to the ventral border of the ramus, but successive
foramina are higher on the ramus so that the last, just below the anterior root of M1, is very near
the dorsal border of the ramus. There are some seven or eight mental foramina in total.

Dentition

Upper dentition: The dental formula of Dorudon atrox is 3.1.4.1/3.1.4.2. Most of the upper
dentition was destroyed by erosion. What remains is shown in (Figure 4.1). The upper premolars
are represented by left and right P3-4. These are double-rooted with a crown that is roughly
triangular in lateral view. The deciduous first premolar has a single mesiodistally elongated root.
These teeth are both worn in life and damaged by erosion, making counts of cusps and precise
measurements impossible.

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The only upper molar preserved is left M1. This tooth is heavily worn on the lingual side. It has
two roots, with the posterior root being slightly more expanded lingually than the anterior root.

Lower dentition: Dorudon atrox preserves the alveolus of a single-rooted (I1-p1). (p2-m2) and
both of these teeth appear to have been double-rooted. The third lower premolar and the fourth
lower premolar have roughly triangular lateral views. The second lower molar is erupting yet on
both side

The only upper molar preserved is left M1.The first upper molar (M1, Figure 4.5) is much smaller
than the preceding premolars. The lateral-view profile of MI is much more rounded than that of
the premolars, which are distinctly triangular. M1 is buccolingually compressed and generally
mesiodistally symmetrical. The distal root has a lingual expansion and a midline groove on its
distal margin. The crown also has a lingual projection over this root. The distal root appears to
have developed from the fusion of two separate roots. The crown projects farther up onto the
roots on the lingual side than on the buccal side. The crown is deflected away from the division
between the roots on both the buccal and lingual sides. The central cusp of M1, the paracone, is
located directly below the division between the mesial and distal roots. There are two accessory
denticles on the mesial edge of the crown. The denticles increase in size from mesial to distal.
The first denticle is on the well-developed mesial cingulum. There are also three accessory
denticles on the distal edge of the crown. The denticles decrease in size from mesial to distal.
The last denticle joins with the well-developed distal cingulum. There is no diastema mesial or
distal to M1.

Axial skeleton (postcranial)

The axial postcranial skeleton of cetaceans has been of great interest to modem and fossil
cetologists alike since it has undergone a major transformation from the terrestrial ancestors of
cetaceans, and it is the sole body support for the musculature used for propulsion in modern
cetaceans. Most vertebrae in mammalian vertebral columns can be separated into five distinct
regions. From anterior to posterior these are: cervical, thoracic (dorsal), lumbar, sacral, and
caudal. Dorudon atrox has 7 cervicals, 17 thoracics, 20 lumbars, no sacrals, and 21 caudals.

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Cervical vertebrae: The bodies of the cervical vertebrae of Dorudon atrox are compressed
like those of modern cetaceans, but they remain relatively longer than the cervical vertebrae of
most modern cetaceans. There is no cervical fusion known in any specimens of Dorudon atrox

Atlas: The first cervical vertebra or atlas (Figure 4.9) is the largest of all of the cervical
vertebrae. It has a very thin body and forms an oval ring around the foramen magnum. The neural
arch [na] is robust and is about equal in craniocaudal length to the ventral arch. The ventral arch
(va) is dorsoventrally thicker than the neural arch and it supports both the cranial articular fovea
(cafl) and the caudal articular fovea (cdf) on its cranial and caudal sides, respectively. The ventral
surface of the ventral arch is smooth and lacks a hypapophysis. The neural canal (nc) is generally
round and is approximately the same size as the foramen magnum. There is a single broad
transverse process [t] that projects laterally and caudally from the sides of the vertebrae. The
transverse process is narrow on its ventral surface and forms a broad, convex shelf on its dorsal
surface.

The transverse process contains a small foramen that opens into a canal that runs craniocaudally,
entering the vertebra lateral to the cranial articular fovea and exiting laterally to the caudal
articular fovea. This foramen is very small when compared to the vertebrarterial foramina of the
other cervical vertebrae. It is unlikely that this foramen could have been transmitted to the
vertebral artery. It is also quite variable in size from specimen to specimen. The cranial articular
foveae are concave and kidney-shaped. The ventral edges are separated by a narrow space on the
ventral arch of the vertebra. The dorsal margins of the foveae are connected to the neural arch
by bony bridges that form the lateral vertebral foramina [Ivfl through which the vertebral arteries
pass.

The caudal articular foveae project caudally from the posterior surface of the vertebra. The dorsal
projections of the foveae are rounded when viewed from the caudal side but are very thin
craniocaudally. The two sides of the foveae are confluent with the fovea of the dens across the
ventral arch. The dens of the axis rest in the fovea of the dens. The ventral tubercle of the atlas
[vt] is a small, rounded, sub-triangular process projecting from the posterior side of the ventral
arch that projects under the body of C2, but it does not articulate with C2. When in articulation
with the axis, the neural arch of the atlas and the pedicles of the axis form the intervertebral

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foramen [ivfl through which the second spinal nerve passes. The first spinal nerve passes
between the atlas and the skull anterior to the dorsal arch of the atlas.

Cervical vertebrae 4 to 6 (C4-6; Figures 4.10): The bodies of C4-C6 are craniocaudally com-
pressed and concave on the cranial and caudal ends. The vertebral bodies decrease in length from
C4 to C5 and increase from C5 to C6. The cranial epiphyses are roughly oval but somewhat
flattened along the dorsal and ventral margins. The caudal epiphyses [cae] are more nearly oval,
especially in younger individuals. Older individuals are less oval because the epiphysis extends
onto the double hypophysis present on the anterior and posterior ends of the ventral surfaces of
the bodies. There is a ventrally projecting midline ridge on the ventral surface of each vertebral
body. The double hypophysis of C4-C6 is reduced.

The pedicles are elongated laterally and rather short craniocaudally. They are not nearly as large
as those of the axis. The pedicles become more laterally elongated from C4 to C6. Spaces
between pedicles on adjacent vertebrae form the intervertebral foramina for the passage of spinal
nerves. The pedicles on each side of the vertebrae join the laminae, which together form the
neural arches. The laminae are flat and both craniocaudally and dorsoventrally thin. The laminae
become more craniocaudally narrow from C4 to C6. The neural canals are arch-shaped. The area
of the neural canal increases from C4 to C6.

The neural spines are very short and blunt. The neural spines are low on the cranial margin of
the neural arch and higher on the posterior margin. The neural spine of C7 is a little more
substantial. It has a broader base and is a little taller than the neural spines of C4 to C6.

The prezygapophyses project cranially from the neural arch, and the articular surfaces are angled
ventrally and medially. The articular surfaces of C6 are larger than those of C4 to C5. The
prezygapophyses articulate with the postzygapophyses of the preceding vertebra. The
postzygapophyses are angled ventrally and laterally.

The transverse processes are broad and flattened craniocaudally. They are formed by the fusion
of the more dorsal diapophyses with the more ventral parapophyses. The parapophyses retain
slender processes that project ventrally and cranially from the ventral margin of the transverse
processes. These processes become more robust from C4 to C5. The parapophyses are very large
and club-shaped in C6. The processes project about as far from the ventral surface of the body

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as the body is tall. The processes are thickest along their cranial margins and slightly thinner
toward their caudal margins.

Ribs: The second right rib (Figure 4.11) of the new specimen has two separate articular surfaces
where they meet the vertebrae, one on the capitulum, and the other on the tuberculum. The
capitulum articulates with two adjacent vertebrae. Each capitulum articulates with the thoracic
vertebra to which it corresponds in number (capitulum of rib R2 with thoracic vertebra T2), and
with the vertebra immediately anterior. The capitular articular surfaces on the vertebrae are
divided between the two adjacent vertebrae with the more cranial portion of the articular surface
on the more cranial vertebra, and the caudal portion of the articular surface on the rib's
corresponding vertebra.

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Figure 4.1. Dorudon atrox skeletal restoration. This restoration is a composite based mainly on
UM 101215 and 101222. The innominate and pes are not known from D. atrox, but the proximal
femur, patella, and astragalus of D. atrox are similar to those of Basilosaurus isis (Gingerich et
al, 1990). The innominate here is restored from closely related Chrysocetus healyorum, and the
pes is restored from B. isis. The length of the skeleton is 4.85 meters. (after Uhen, 2004)

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Figure 4.2. Dorudon atrox skull (A) and dentary (B) restoration, in right lateral view.
Restoration is a composite, based mainly on NSFM 4451, and UM 93220, 100139, and 101222.
The dentary is shown disarticulated from the skull. The length of skull is about 95 cm (after
Uhen, 2004).

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Figure 4.3. Dorsal view of the skull of Dorudon atrox Abbreviations: eoc, external occipital
crest; fr, frontal; mx, maxilla; na, nasal; nc, nuchal crest; ncv, narial cavity; pr, parietal; pmx,
premaxilla; pspf, postorbital process of the frontal; sgc, sagittal crest; sopf, supraorbital process
of the frontal; sq, squamosal. Note the skull torsion at the rostrum level.

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Figure 4.4. Ventral view of the skull of Dorudon atrox at MUVP. Abbreviations: al, alisphenoid;
bo, basioccipital; ep, embrasure pit; fr, frontal; gf, glenoid fossa; j, jugal; mx, maxilla; or,
orbitopshenoid; paf, palatine foramen; pal, palatine; ppm, palatal process of the maxilla; pgp,
postglenoid process; pmx, premaxilla; ppe, paroccipital process of the exoccipital; pt, pterygoid;
pts, pterygoid sinus; shf, fossa for the stylohyal contact; sopf, supraorbital process of the frontal;
v, vomer; zyg, zygomatic process of the squamosal.

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 Figure 4.5. The Left first upper molar (M1) in labial (A) and lingual (B) views.

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Figure 4.6. Right tympanic bulla of Dorudon atrox (A) in dorsal (B), posteromedial (C), and
ventral (D) views.

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Figure 4.7. Lateral views of the left (A) and right (B) dentaries of Dorudon atrox. Abbreviations:
cp, coronoid process; ep, embrasure pit; mf, mental foramina.

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Figure 4.8. Medial views of the left (A) and right (B) dentaries of Dorudon atrox.
Abbreviations:cp, coronoid process; mbf, mandibular foramen; sym, mandibular symphysis.

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Figure 4.9. Atlas (C1) of Dorudon atrox in anterior (A) and posterior (B) views. Abbreviations:
aaf, anterior articular fovea; da, dorsal arch of the atlas; nc, neural canal; pas, posterior articular
surface; tp, transverse process; vf, vertebrarterial foramen; vt, ventral tubercle of the atlas.

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Figure 4.10. Cervical vertebrae of Dorudon atrox, C4 in anterior (A) and posterior (B) views;
C5 in anterior (C) and posterior (D) views; C6 in anterior (E) and posterior (F) views.
Abbreviations: di, diapophysi; nc, neural canal; pa, parapophysis; prz, prezygapophysis; psz,
postzygapophysis; vf, vertebrarterial foramen.

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 Figure 4.11. The second right rib (R2) of Dorudon atrox at MUVP .

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 SUMMARY AND CONCLUSIONS
The Fayum Depression represents the first Paleogene locality that was discovered in Africa. In
the early years, the remarkable richness and diversity of the Fayum faunal assemblages sparked
the interest of many important vertebrate paleontologists, who were then involved in the
collection and description of the Fayum fossils. These paleontologists include the British Charles
W. Andrews, the Germans Eberhard Fraas, Ernst Stromer von Reichenbach, and Max Schlosser,
as well as the American Henry F. Osborn.

The geology and vertebrate paleontology of the Eocene sequence in the study Wadi El-Hitan
area, Fayoum, Egypt attracted the attention of many scientists from the nineteenth century until
the present day. This is because the study area includes well-preserved Eocene marine
vertebrates at the UNESCO World Heritage site. The present work is a trail to integrate the
paleontological and sedimentological factors governing the distribution and paleoenvironmental
framework of the Late Bartonian- Priabonian rock units and their fossil contents in Wadi El-
Hitan. The cetaceans from the Eocene of the Fayum Depression show how morphologically
advanced and diverse their early representatives from this time were. Wadi El Hitan has provided
evidence for at least four distinct whale species, Basilosaurus isis, Dorudon atrox, Ancalecetus
simonsi, and Masracetus markgrafi.

Dorudon atrox occupies a transitional position in the evolution of cetaceans. It is one of the
earliest fully aquatic cetaceans and it is much more primitive than modern cetaceans. Despite
this conclusion about its position in cetacean history, one should not view Dorudon atrox as a
poorly adapted version of a modem cetacean. Dorudon atrox has aquatic adaptations shared with
modern cetaceans and other characteristics that are not. These others are not inferior to those
adaptations seen in modem cetaceans, just different. Dorudon atrox individuals were adequately
adapted to living in the warm shallow seas of the Eocene of Egypt. Dorudon atrox has anatomical
features that suggest it was completely aquatic. All of the differences in the auditory region
between D. atrox and terrestrial mammals are shared with modem cetaceans. D. atrox has a
large, dense auditory bulla that is loosely attached to the basicranium.

The skull and jaws of D. atrox are elongated, with the anterior teeth in line with the cheek teeth.
This is a feature seen in many piscivores, including modem odontocetes, suggesting that D. atrox
was also piscivorous. This is confined by the presence of fish bones in the remains identified as

101 | Page
stomach contents in D. atrox. The upper and lower anterior teeth interlock when the jaws are
closed and moved slightly forward. D. atrox used these anterior teeth for prey acquisition and
subsequently moved prey items to the back of the oral cavity for processing. The cheek teeth of
Dorudon atrox are buccolingually compressed, serrated blades that sheared large food items into
smaller pieces when the lower jaws were brought into occlusion with the uppers. This is very
different from feeding in odontocetes. Most odontocetes use their teeth only during prey
acquisition and then swallow prey whole. Locomotion in Dorudon atrox was basically like that
of modern cetaceans.

The vertebral column of D. atrox shows many characteristics typical of modem cetaceans
including a great increase in the number of lumbar vertebrae; relatively uniform vertebral body
size in the posterior thoracic, lumbar, and anterior caudal vertebrae; and dorsoventrally
compressed posterior caudal vertebrae. More lumbar vertebrae also indicate the presence of
greater epaxial and hypaxial muscle mass to move the tail. This is significant since modern
cetaceans have abandoned primitive mammalian limb-based locomotion in favor of a new tail-
based locomotion. Studies of other basilosaurids may show that this transition is similar to the
origin of Cetacea in complexity. The last key point in cetacean evolution is the origin of the
modern suborders of Cetacea. The fossil record of early mysticetes and odontocetes is currently
improving. The fossils now known suggest a very complex early history for these groups. More
studies of dorudontine archaeocetes are essential to the study of early mysticetes and early
odontocetes since both groups are thought to have originated from dorudontines.

102 | Page
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 ‫الملخص العربي‬

‫تعتبر منطقة وادي الحيتان بالفيوم جمهورية مصر العربية من أهم مناطق التارث الطبيعي العالمي التي جذبت عدد كبير من العلماء لدارسة‬
‫التراكيب الجيولوجية والمحتوي الحفري بالمنطقة حيث تحتوي علي عدد كبير جدا من هياكل الحيتان المتحجرة خالل العصر االي والمتوسط‬
‫وسيني العلوي ‪ .‬نركزت منطقة الدارسة الحالية علي دارسة المحتوي الحفري بالمنطقة وكذا عوامل الترسيب التي اثرت علي توزيع السحنات‬
‫وكذلك البيئة القديمة بالمنطقة خالل العصر االيوسيني المتوسط والعلوي‪ .‬كما تركزت الدارسة علي وصف عام للتركيب التشريحي لهياكل‬
‫الحيتان المتحفر الموجودة بالمنطقة وخاصة نوعي الباسيلوصورس ايزيس والدوريودون اتروكس وكذلك درجة حفظ وتماسك تلك الهياكل‬
‫بالطبقات الحاوية لها وتوزيعها الطبقي ‪.‬يُعتبر حوت الدوردون أحد األنواع المنقرضة التي كانت تعيش في المحيطات منذ حوالي ‪ 40‬إلى ‪34‬‬
‫مليون سنة خالل فترة اإليوسين المتأخر‪ .‬هذا النوع من الحيتان يمثل مرحلة مهمة في تطور الحيتان من كائنات برية إلى مائية بالكامل‪ .‬يعتبر‬
‫حوت الدوردون أحد األسالف المباشرة للحيتان الحديثة‪ ،‬ويمثل حلقة انتقالية في سلسلة التطور ‪.‬الوصف نحيفا وطويال‪ ،‬حيث بلغ طوله حوالي‬
‫‪ 4.5‬أمتار‪ .‬كانت له جمجمة طويلة وفكوك قوية مليئة بأسنان كان حوت الدوردون يمتلك جسما ‪.‬كان لديه أطراف صغيرة‪ ،‬وهو ما يميز‬
‫أسالف الحيتان‪ ،‬إذ كانت تلك األطراف الخلفية تشير حادة‪ ،‬مما يشير إلى أنه كان صيادا ماهرا إلى بداية التحول من الحياة البرية إلى الحياة‬
‫المائية ‪.‬الجمجمة الطويلة لحوت الدوردون كانت تشبه إلى حد كبير جمجمة الحيتان الحديثة‪ ،‬إال أنها كانت تحتوي على أسنان كبيرة وحادة‪،‬‬
‫مما يدل على نظام غذائي يعتمد على صيد الفرائس الكبيرة مثل األسماك والحبار‪ .‬كانت أذنه الوسطى متطورة‪ ،‬مما يدل على قدرته على‬
‫السمع تحت الماء‪ ،‬وهي سمة مشتركة بين الحيتان الحديثة ‪.‬عاش حوت الدوردون في البحار الضحلة والمناطق الساحلية‪ ،‬حيث كان يجد‬
‫غذاءه من األسماك والحبار‪ .‬وقد تم العثور على حفرياته في مواقع متعددة حول العالم‪ ،‬بما في ذلك أمريكا الشمالية وأوروبا وأفريقيا‪ .‬توزع‬
‫حوت الدوردون على نطاق واسع يدل على قدرته على التأقلم مع بيئات مختلفة وموارد غذائية متنوعة ‪.‬حوت الدوردون يحمل أهمية كبيرة‬
‫في فهم تطور الحيتان‪ .‬تشير الدراسات إلى أن حوت الدوردون كان يمتلك خصائص متقدمة في األذن الوسطى تساعده على السمع تحت الماء‪،‬‬
‫وهي سمة مهمة للحيتان الحديثة‪ .‬كما أن األطراف الخلفية الصغيرة التي كان يمتلكها حوت الدوردون تشير إلى أن هذه الحيتان كانت في‬

‫مرحلة انتقالية من الحياة البرية إلى الحياة المائية‪ ،‬مما يعطي العلماء فهما أعمق لكيفية ً تطور الحيتان‪ ..‬الدراسات التي أجريت على حفريات‬
‫حوت الدوردون أظهرت أن هذه الحيتان كانت تمتلك تركيبة عظمية مشابهة إلى حد كبير لتركيبة الحيتان الحديثة‪ ،‬مما يدل على أن العديد من‬
‫الخصائص الحيوية للحيتان قد بدأت بالتطور في هذه الفترة المبكرة‪ .‬هذه الخصائص تشمل القدرة على السباحة بسرعة والبحث عن الفرائس‬
‫في المياه العميقة ‪.‬تكيف حوت الدوردون مع البيئة المائية كان ضروريا لبقائه واستمراره‪ .‬أظهرت الدراسات أن األطراف الخلفية لحوت‬
‫الدوردون كانت ً تبدأ في االنكماش‪ ،‬مما يشير إلى تحول تدريجي نحو الحياة المائية بالكامل‪ .‬كانت هذه األطراف تعمل بشكل رئيسي‬
‫كمجاذيف تساعده على السباحة بدال من استخدامها للمشي على اليابسة ‪.‬باإلضافة إلى ذلك‪ ،‬كانت األسنان القوية لحوت الدوردون تشير إلى‬
‫أنه كان صيادا قادرا على افتراس الفرائس الكبيرة‪ ،‬مما يدل على ً نظام غذائي غني بالبروتينات والدهون االلزمة للحفاظ على جسمه الكبير‪.‬‬
‫كما أن قدرته على السمع تحت الماء ساعدته في تحديد موقع الفرائس والتواصل مع حيتان أخرى‪ ،‬مما يعطيه ميزة كبيرة في بيئته البحرية ‪.‬‬
‫لعملية التحول من الحياة البرية إلى الحياة المائية ‪.‬حوت الدوردون يمثل جزءا هاما من تاريخ تطور الحيتان‪ ،‬حيث يعطينا فهما عميقا دراسة‬
‫حفريات حوت الدوردون توفر للعلماء معلومات قيمة حول كيفية تكيف الكائنات الحية مع البيئات المختلفة وكيفية تطور الخصائص الحيوية‬
‫التي نراها في الحيتان الحديثة ‪.‬يمثل حوت الدوردون مرحلة انتقالية حاسمة في تطور الحيتان‪ ،‬ويظهر كيف يمكن للتكيف مع البيئة أن يؤدي‬
‫إلى تغييرات كبيرة في الشكل والسلوك والوظائف الحيوية للكائنات الحية‪ .‬من خالل فهم حوت الدوردون‪ ،‬يمكننا الحصول على نظرة أعمق‬
‫على عملية التطور والتكيف التي أدت إلى ظهور الحيتان كما نعرفها اليوم‪ ،‬مما يعزز فهمنا للتنوع البيولوجي وأهمية الحفاظ على االنواع‬
‫الحية‪.‬‬

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