Received: 1 July 2021 Revised: 26 November 2021 Accepted: 29 November 2021

DOI: 10.1002/gj.4357

RESEARCH ARTICLE

First occurrence of sauropod and theropod footprints in the

pre-Campanian deposits of the Nubian Sandstone, south
Eastern Desert, Egypt

Walid G. Kassab1 | Gebely A. Abu El-Kheir2 | Mohamed K. AbdelGawad1

1
Faculty of Science, Geology Department,
Cairo University, Giza, Egypt
2
Faculty of Science, Geology Department,
New Valley University, New Valley, Egypt
Correspondence
Mohamed K. AbdelGawad, Faculty of Science,
Geology Department, Cairo University, Giza,
Egypt.
Email: mkabdelgawad@cu.edu.eg;
mkabdelgawad@sci.cu.edu.eg
Funding information
New Valley University; Cairo University
Handling Editor: I. D. Somerville
Dinosaur footprints occur in the Upper Cretaceous strata known as the Nubian Sand-
stone Formation in the south Eastern Desert, Egypt. The Nubian Sandstone unit is
subdivided into three informal members: A lower member is marked by reddish-
brown, trough cross-bedded, coarse- to medium-grained sandstone interbedded with
thin beds of varicolored mudstone, reflecting deposition in a braided fluvial system, a
middle member is characterized by reddish-white to yellow, planar cross-bedded,
medium-grained sandstone with ferruginous siltstone intercalations, interpreted as
meandering fluvial deposits, and an upper member consists of repeated coarsening-
upward cycles of rippled siltstone to fine- to medium-grained sandstone and planar
cross-bedded, yellowish-white sandstone, containing many vertebrate footprints and
invertebrate trace fossils, reflecting deposition in a coastal to deltaic environment.
Sixteen dinosaur footprints are recorded at the base of the upper member of the
Nubian Sandstone, on inclined surfaces of rippled, fine-grained sandstones. Twelve
are overlapping each other and belong to distinct individuals of dinosaurs, the other
four are distributed on a younger bedding surface. Twelve footprints are from more
than one sauropod dinosaur, while three of the isolated footprints belong to a thero-
pod dinosaur. The footprints described are semiplantigrade and digitigrade. This dis-
covery is the second record of tetrapod footprints in Egypt and the first record of
tetrapod footprints in the Eastern Desert.

KEYWORDS
dinosaur, Eastern Desert, Egypt, footprints, Late Cretaceous, Nubian Sandstone, tetrapod

1 | I N T RO DU CT I O N footstep known as a track, footprint, or ichnite (Leonardi, 1987). Three

Vertebrate trace fossils are the result of the interaction between ver-
tebrate behaviour and a medium, which includes softgrounds,
firmgrounds, hardgrounds, plants, and other animals. Trace fossils are
used in conjunction with plants, vertebrates, and invertebrate fauna,
as well as other proxies to measure changes in the geosphere, bio-
sphere, and atmosphere (Hasiotis, Platt, Hembree, & Everhart, 2007).
Tetrapod locomotion traces may contain sequences of impressions
of feet (pes) and hands (manus) and any part of the trackmaker that
was in contact with the sediment. Each impression represents a single
or more successive footprints from the same animal are defined as a
trackway (Leonardi, 1987; T. Thulborn, 1990). Assemblages of foot-
prints produced by one or more individuals from the same species, for
which it is impossible to identify whether the animal was moving
and/or its direction of movement, is defined as footprint group
(Mezga & Bajraktarevi
c, 1999). The morphology of locomotion tracks is
defined as the function of foot shape, body and tail motion, and control,
substrate consistency, and trace maker locomotion dynamics and
posture (Brand, 1996; Demathieu & Demathieu, 2002; Falkingham &
Gatesy, 2014).

Geological Journal. 2021;1–12. wileyonlinelibrary.com/journal/gj © 2021 John Wiley & Sons Ltd. 1
2 KASSAB ET AL.
In this study, we document for the first time the occurrence of
tetrapod footprints from the pre-Campanian Nubian Sandstone For-
mation (Fm) in the Quseir area, in the south Eastern Desert, Egypt.
This study aims to (a) document the first occurrence of tetrapod foot-
prints in their stratigraphic context within the Nubian Sandstone For-
mation; (b) understand the relationship between the vertebrate trace
fossils and the sediment nature (grain size, substrate type), and the
depositional and ecological conditions.
Fossil footprints in the Nubian Sandstone Formation are pre-
served in originally soft, fine-grained sediment. The footprints are
recorded in situ on inclined bedding planes of rippled fine-grained
sandstone beds.
The integration of depositional analysis with fossil footprints data
in the Nubian Sandstone Formation at the Quseir area highlights the
environment of Late Cretaceous dinosaur communities. The preserva-
tion of the footprints is poor (Haubold, Hunt, Lucas, & Lockley, 1995;
Marchetti et al., 2019; Peabody, 1948), so ichnogeneric and
ichnospecific names were not assigned to the studied material. This
discovery is the second record of tetrapod footprint in Egypt after
Demathieu and Wycisk (1990). It is the first record of dinosaur foot-
prints in the Eastern Desert of Egypt.
2 | G EO LO GI C BA C KGR OU N D
In the Quseir area at Gebel Duwi (Figure 1), the Precambrian basement
rocks include ultramafics and pillowed tholeiites with ophiolitic affinities,
banded iron-formation metasediments, and serpentinite mélanges, in
addition to various plutonic rocks (Ries, Shackleton, Graham, &
Fitches, 1983; Stern, 1979; Sturchio, Sultan, & Batiza, 1983). The high
deformation is attributed to the closing of small oceanic basins and conti-
nental collision. The basement rocks are unconformably overlain by
Upper Cretaceous-Palaeogene deposits that represent the main sedi-
mentary sequence in the Quseir area. This succession was deposited in a
faulted basin, formed before the Red Sea rifting (Khalil & McClay, 2002).
The Late Cretaceous to Palaeocene sedimentary succession shows a ver-
tical change of facies, from continental to shallow marine, reflecting a
general deepening-upward trend, due to the continuous rise of sea level
during the Campanian to Palaeocene interval in North Africa. The 70 m
to nearly 200 m-thick lower part consists of massive, thick-bedded,
siliciclastic, nonfossiliferous rocks, except for the occurrence of tetrapod
footprints in the Nubian Sandstone Formation and lies unconformably
above the basement rocks. The presence of trough, tabular, planar, or
cross-bedded, medium- to coarse-grained sandstone suggests channel
deposits of low sinuosity rivers. The Nubian Sandstone Formation is
assumed to be pre-Campanian in age, based on its stratigraphic position.
The Nubian Sandstone Formation grades upward during the Cam-
panian into a 220 to 370 m-thick sequence of fossiliferous alternating
shales, sandstones, and limestones. This sequence comprises
(a) fossiliferous, variegated shales, deposited in a shallow marine envi-
ronment (Quseir Formation; Youssef, 1957), and (b) phosphate-rich
shales, deposited in shallow marine-isolated basins (Duwi Formation;
Youssef, 1957). During the Maastrichtian-Palaeocene times, the Duwi
Formation was covered by marine shale, carbonate, and green shale
that reflect increasingly deep marine facies. They represent the
Dakhla, Tarawan, and Esna formations, respectively.
2.1 | Lithostratigraphy and depositional
environments of Nubian Sandstone Formation
In Gebel Duwi (Latitude: 26
60 59.5800 N; Longitude: 34
20 22.4000 E), the
Nubian Sandstone Formation rests unconformably on the basement
complex (Figures 2 and 3a) and underlies a thick Upper Cretaceous
variegated marine shale sequence, known as the Quseir Formation.
The Nubian Sandstone Formation consists mainly of poorly to well-
sorted, fine- to coarse-grained, reddish-brown to yellowish sandstone,
with some siltstone and/or mudstone intercalations. Youssef (1957)
suggested a Santonian age for the Nubian Sandstone Formation. It is
considered by other authors to be as old as Cenomanian (Ward &
McDonald, 1979). Based on fossil evidence within the overlying
Quseir Formation, Issawi, Francis, Youssef, and Osman (2009)
assigned a pre-Campanian age for the Nubian Sandstone Formation.
The thickness of the Nubian Sandstone Formation in the Eastern
Desert is highly variable, depending on the presence or absence of
basement topographic highs. Its stratigraphic thickness reaches about
200 m in the Gebel Duwi area. Based on field observations in Gebel
Duwi, the Nubian Sandstone Formation could be subdivided into
three informal members (Figure 2). Each of these members exhibits a
unique set of properties that distinguish it from the others, including
colour, grain size, sorting, geometry, and sedimentary structures.
The lower member of the Nubian Sandstone Formation reaches
about 60 m in thickness, is composed of reddish-brown, trough cross-
bedded sandstones (Figure 3b) with some conglomerate levels
(Figure 3c), intercalated with varicolored mudstone with some roots
and palaeosol remains (Figure 3d), exhibiting an overall fining-upward
trend. The amalgamated channel pattern of this member as well as
the fining-upward cycles reflects deposition in braided fluvial channel
and flood plain environments.
The middle member (40 m thick) is reddish-white, friable,
medium- to fine-grained, tabular cross-bedded sandstone (Figure 3e),
interbedded with varicolored siltstone/mudstone layers interpreted as
meandering fluvial channels and flood plain deposits.
The upper member reaches about 100 m in thickness. The basal
portion is about 40 m thick and is made of fining-upward, black to
reddish-white, fine-grained sandstone. It is characterized by well-
developed ripple marks (Figure 3f) and vertebrate footprints and hori-
zontal invertebrate burrows. The dinosaur footprints are sup-
erimposed on the rippled surfaces. Sauropod dinosaur footprints are
present in the first level of the upper member (elevation 100 m on
Figure 2) and belong to different individuals. The theropod footprints
are found in a higher stratigraphic level, at 110 m in the succession,
and show a few isolated footprints. The rippled, fine-grained sand-
stone of the lower part of the upper member grades upward into tab-
ular cross-bedded sandstone intercalated with rippled siltstone and
upper portion is varicolored silty shale with plant remains. The middle
KASSAB ET AL. 3

part of this member is composed of fining-upward cycles of reddish
to greenish-yellow, tabular cross-bedded, fine-grained sandstone
intercalated at the base with sandy siltstone and topped by varicol-
ored silty shale. The top of the upper member consists of
coarsening-upward, repeated cycles of yellow, tabular cross-bedded,
well-sorted, and fine-grained sandstone interbedded with siltstone.
This member reflects deposition in coastal plain/shallow marine to
delta plain environments. The sandy sediments of this unit grade
upward into the variegated shales of the overlying Quseir
Formation.

F I G U R E 1 (a) Location map of the studied area; (b) Geologic map of Gebel Duwi, modified after Khalil and McClay (2002) and Khalil, Fathy,
Abdeldayem, and Ghobara (2016)
4 KASSAB ET AL.

FIGURE 2 Stratigraphic column of the Nubian Sandstone Formation in the studied area

The lower and middle members of the Nubian Sandstone Forma- changes upward to a meandering fluvial environment with a flood
tion are predominantly fluvial sediments deposited in a wide alluvial plain. The upper member consists of coastal plain and/or tide-
plain, of a dominantly braided fluvial system. The lower member dominated delta plain deposits, which grade upward into the coastal
KASSAB ET AL. 5

F I G U R E 3 Field observations of the three informal members of Nubian Sandstone Formation in Gebel Duwi, Quseir area. (a) Nubian
Sandstone Formation (NSF) rests unconformably on the basement complex (b); (b) Reddish-brown, trough cross-bedded sandstone, lower
member of NSF; (c) Reddish-brown, highly weathered, moderately hard, large-scale trough cross-bedded, pebbly to coarse sandstone with minor
conglomeratic content, lower member of NSF; (d) Varicolored mudstone with some plant roots and remains of palaeosols, lower member of
Nubian Sandstone Formation; (e) Reddish-white, friable, medium- to fine-grained, tabular cross-bedded sandstone, the middle member of Nubian
Sandstone Formation; (f) Well-developed ripple marks in the upper member of Nubian Sandstone Formation

and near-shore marine deposits of the Quseir Fm (Ward, 3 | M A T E R I A L S A N D M ET H O D S

Mcdonald, & Mansour, 1979), of Campanian age (Abd El-Razik,
1967; El-Dawoody, 1970; El-Tarabili, 1966; Youssef, 1957). As a
matter of fact, the occurrence of marine and freshwater gastro-
pods and bivalves from the Quseir Formation and plant remains
and fossil wood (Issawi, Francis, El Hinnawy, Mehanna, & El
Deftar, 1971; Seward, 1935) indicates a marginal marine to estua-
rine depositional environment. The Quseir Formation marks an
upward transition from a continental environment to a shallow
marine environment.
Sixteen footprints have been found on the upper surface of tectoni-
cally inclined beds of rippled, fine-grained sandstone. Twelve of them
are overlapping each other and belong to distinct tetrapod individuals,
the other four found on an upper bedding plane, are randomly distrib-
uted, these are theropod footprints (Figure 2). The footprints are num-
bered according to their position from the right position to the left
position and then the second row from the right to the left as shown
in Figure 4.
6 KASSAB ET AL.

F I G U R E 4 (a) The Twelve overlapping footprints with scale 120 cm; (b) An illustration of the twelve overlapping footprints with scale 120 cm
with colour coding may attributed to the same individuals; (c–e) Close-up views for selected footprints

Footprints were photographed, measured, and their geographic
positions were recorded (Latitude: 26
60 59.5800 N; Longitude:
34
20 22.4000 E). The measured parameters are: The overall length and
width of the footprint, length and width of each digit (if possible), the
interdigital angle between the two adjacent digits (if possible), the
depth of penetration of the claws (anterior portion of the footprint),
and of posterior margin of the footprint- and the total divarication.
The parameters were measured in centimetres (cm) (Table 1).
The equations used to determine the body length, the hip height
(leg length), the weight and mass of the body acting on one foot, and
the average mass of the body on one foot were based on
Alexander (1976), R. A. Thulborn (1984, 1989), T. Thulborn (1990),
R. A. Thulborn and Wade (1979, 1984), Henderson (2003), Mezga and
Bajraktarevi
c (1999), Day, Norman, Gale, Upchurch, and Powell (2004),
Lee and Huh (2002), Parkes (1993), Weems (2006) and Martin (2006).
Body length for biped dinosaurs = footprint length (FL)  10.
Hip height (H) (leg length) = footprint length (FL)  4
(Alexander, 1976; Day et al., 2004; Henderson, 2003; Lee &
Huh, 2002; Mezga & Bajraktarevi
c, 1999; R. A. Thulborn (1984,
1989), T. Thulborn (1990); R. A. Thulborn & Wade, 1979, 1984);
while Parkes (1993) used fraction methods for determination of
the hip height as if the theropod is small or large size. Herein, we
used the hip height which is equal to 4 times the footprint length
for all the recorded footprints.
Foot width (FW)/foot length (FL) ratio; estimated by
T. Thulborn (1990) see also Parkes (1993).
Foot area (FA) = foot length (FL)  foot width (FW), based on
Parkes (1993).
Body mass for biped dinosaurs = (4.73H)3, based on Weems (2006).
Foot stress for biped dinosaurs (Sf) = (F/2)/FA, where the
F = body mass  gravitational force (9.8 m/s2), and for correction;
multiplying the result by 0.8 as the footprint is more oval than rectan-
gular based on Martin (2006).
The results of the body length, hip height, FW/FL, FA, body mass,
and foot stress for the footprints are shown in Table 2.
4 | F O O T P RI NT ' S I D E N T I F I C A T I O N A N D
DE SC R I PT ION
The footprints are recorded in the upper member of the Nubian Sand-
stone Formation. Based on Leonardi (1987), the described footprints
are semiplantigrade (footprints lack a heel impression) and digitigrade
(footprints contain impressions of the entire digit length) footprints.
TABLE 1 Measurements (cm) of the sixteen footprints found in the Nubian Sandstone Formation

Third digit Second digit Fourth digit Interdigital angle between Depth

Second and Third and (Front) Claws and (Rear) End of the
KASSAB ET AL.

Dinosaur Footprint no FL FW L W L W L W third digits fourth digits Total divarication beginning of digits footprint
Sauropod 1 80 45 80 10a 5 3
45b
2 38 18 38 4a 2 0
18b
3 55 30 55 6a 3 2
30b
Theropod 4 40 22 40 6a 25.1 35.9 59.6 2 1.5
or ornithopod? 22b
Sauropod 5 45 25 45 5a 42.2 Not 3
25b recognized
6 80 60 80 10a 5 3
60b
7 50 28 50 7a 50.3 3 2
28b
8 60 33 60 9a 2 4
33b
9 50 33 50 9 50 5 Third digit 2 Third digit 3
Second digit 1.5 Second digit 2
10 95 65 95 9a 46.5 4a 1
47c 2b
65b
11 50 30 50 24a 3 2
30b
12 60 42 60 28a 5 3
42b
13 55 43 55 20a 2 cm 1 cm
43b
Theropod 14 38 31a 38 11 38 7 38 7 19.2 24.7 43.3 All three digits are 2 2.5
19b
15 40 32 40 6a 26 9a 40 8a 40.7 40.3 83.8 Third digit 1.5 1
10b 10b 12.5b Second, fourth digits 2
16 50 40 50 10a 35 5a 47 6a 16.9 8.1 1 1
16c 10b 10b
20b

Abbreviations: FL, foot length; FW, foot width.

a
Measurement at the front of the footprint.
b
Measurement at the rear of the footprint.
c
Measurement at the middle of the footprint.
7
8 KASSAB ET AL.

T A B L E 2 Footprint length (cm), hip height (cm), body length (cm), footprint width (cm), foot width (FW)/foot length (FL), foot area (FA) (cm2),
body mass (kg), foot stress (N/m2), and foot stress after correction (N/m2) for the sixteen footprints

Footprint Body FW/ Body Foot Sf after

Dinosaur no FL H length FW FL FA Mass stress (Sf) correction
Sauropod 1 80 320 45 0.5 3,600
2 38 152 18 0.4 684
3 55 220 30 0.5 1,650
Theropod or 4 40 160 400 22 0.5 880 433.5 241.4 193.1
ornithopod?
Sauropod 5 45 180 25 0.5 1,125
6 80 320 60 0.7 4,800
7 50 200 28 0.5 1,400
8 60 240 33 0.5 1980
9 50 200 33 0.6 1,650
10 95 380 65 0.6 6,175
11 50 200 30 0.6 1,500
12 60 240 42 0.7 2,520
13 55 220 43 0.7 2,365
Theropod 14 38 152 380 31a 0.8 1,178a 182.9 76.6 61.3
19b 0.5 722b 124.5 99.6
15 40 160 400 32 0.8 1,280 433.5 165.9 132.7
16 50 200 500 40 0.8 2000 846.6 207.4 165.9
a
Measurement at the front of the footprint.
b
Measurement at the rear of the footprint.

sizes (large and medium, ranging from 95 to 40 cm foot length as shown

F I G U R E 5 Footprint number 14 is isolated from the other footprints;
it might be attributed to badly preserved theropod footprint. Scale 10 cm
The dinosaur's footprints were preserved as concave epirelief,
sometimes with impressions of claws, digital pads, and sole pads (refer
to Table 1, and Figures 4, 5, 6).
4.1 | Sauropod and theropod footprints from the
lower level
Twelve footprints are found on the upper surface of a rippled fine-grained
sandstone bed and preserved in concave epirelief (Figure 4). They are
overlapping each other, exhibit various shapes (U- and V-shaped heel) and
in Table 1). All footprints are characterized by the pes length longer than
the pes width, the digit (probably digit III) is longer than wider, the digital
pad is well defined while the phalangeal pads are badly preserved. The
claw marks are relatively well preserved. Footprint number 9 presents two
digits, where digit III is slightly longer and wider than digit II; the heel is
slightly U- to V-shaped and presents an asymmetric impression. The claws
and the distal parts of the digits are more deeply impressed than the heel.
Almost all sauropod footprints exhibit oval or egg-shaped outlines without
diagnostic digit impressions (Garcia-Ramos, Aramburu, Pifiuela, &
Lires, 2000; Gatesy, Middleton, Jenkins, & Shubin, 1999; Leonardi, 1987;
Lockley, 1991; Nadon, 2001). As described, the pes-manus couples and
trackways are not recognizable, while the footprints are aligned in two dif-
ferent rows or sequences perpendicular to the slope. The footprints may
show some differential depth compared to the slope as shown in Table 1.
All twelve footprints probably belong to sauropods and footprint number
4 may belong to either theropod or ornithopod dinosaur (but the criteria
are not enough to indicate which of them) individuals. Footprint number
13 is located south of the above twelve footprints and shows almost the
same characters as the other twelve footprints.
4.2 | Theropod footprints from the upper level
Footprints number 14, 15, and 16 are distributed in the same area, in
an upper stratigraphic level, on a rippled fine-grained sandstone bed
KASSAB ET AL.
FIGURE 6 Theropod footprint number 15. Scale 10 cm
(Figures 5 and 6). They are preserved as concave epirelief. The foot-
print number 15 with foot length 40 cm tridactyl with the impression
of the digit III, significantly larger than both digits II and IV. The claw
marks are present with sharp edges, medially directed for digits II and
III and laterally directed for digit IV. All the footprints in the upper
level are characterized by the presence of three digits with claw marks
and digit III is about 20–30% longer than digit II and IV (especially in
footprint number 15) which allowed us to interpret them as theropods
footprints. A theropod track is not recognizable. Sometimes, theropod
footprints display tarsometatarsal impressions (Moreno et al., 2012),
but these footprints are badly preserved and do not show this
feature.
5 | DISCUSSION
Based on the morphological description of the footprints in the lower
level, it is interpreted as follows: (a) Based on Weems (2006) estimat-
ing the relation between the hip height (H) and the body mass, the
footprint number 4 which may belong to a small- to medium-sized
theropod dinosaur with body mass about 434 kg. (b) From the hip
height results, there are seven sauropod individuals and two theropod
individuals. The footprint numbers 1 and 6 belong to the hindlimb or
forelimb of the same sauropod individual, with a pace length equals
0.4 m and the dinosaur was moving, as well as numbers 8 and 12 and
numbers 7 and 11, with a pace length equals 1.4 m for both individ-
uals, and the dinosaurs were stopping as the two feet were almost
parallel to each other. Footprints numbers 3 and 13 belong to the
same sauropod individual, but far from each other, as it might be a
trackway for this dinosaur. The other footprints 2, 5, 9, and
10 are attributed to be other sauropod individuals. (c) Several
authors (Farlow, Pittman, & Hawthorne, 1989; Farlow, 1992; Lockley,
Farlow, & Meyer, 1994; see also Day et al., 2004) had reported that
sauropod trackways belong to two distinct categories of “narrow-
gauge” named as Parabrontopodus (Lockley et al., 1994) and “wide-
gauge” named as Brontopodus (Farlow et al., 1989). Brontopodus
9
shows that the left and right prints are widely spaced from the mid-
line, with the manus prints located closer to the midline than the pes
prints with no association of the impression of pollex claw as pres-
ented in both two individuals with footprints 8, 12, and 7, 11. Para-
brontopodus shows that both manus and pes prints are located very
close to or almost intersect the trackway midline and are accompanied
by pollex claw impressions (Lockley et al., 1994), as observed from the
individual with footprints numbers 1, 6.
The morphological description for the footprints in the upper
levels shows the following: Based on the relation between the hip
height (H) and the body mass (Weems, 2006), the footprint numbers
14 and 15 belong to a small- to medium-sized theropod dinosaur with
a body mass ranging from 183 to 434 kg while the footprint number
16 belongs to a large theropod dinosaur with body mass about
847 kg.
During the Late Cretaceous, dinosaurs are found in many basins
in Egypt, such as the Baharyia Oasis during the Cenomanian time
(Smith et al., 2001; Stromer, 1936), and the Dakhla, Kharga, and Baris
oases during the Campanian (Rauhut & Werner, 1997; Sallam
et al., 2016, 2018). The presence of dinosaurs in the Baharyia, Dakhla,
Kharga, and Baris oases throughout both the Cenomanian and Cam-
panian periods suggests the existence of dinosaurs in the Quseir
region, which is located at the same latitude and to the east of these
oases (Figure 1). As a result, it is probable that some dinosaurs lived
there and/or relocated from Baharyia Oasis to the Quseir region dur-
ing the pre-Campanian period.
The preservation of the footprints had a direct relationship with
its geological context. Therefore, the importance of the
palaeoenvironment and the associated substrate water saturation
should be considered. We record the footprints as true tracks
(Lockley, 1991) and footprints rather than undertracks. All footprints
are quite shallow, comparing with their average size, which indicates
that the sediment was cohesive (firmgrounds) substrates, with consid-
erable resistance to imprinting.
Footprints are known as biosedimentary structures, and depict
a closely relation with the environment and the nature of the
substrate. Biodeformation features and the understanding of
their origin can predict the paleoenvironmental interpretation
(Avanzini, 1998; Carvalho & Leonardi, 2020; Díaz-Martínez,

nsole-Gonella, de Valais, & Salgado, 2017; Falkingham, Bates,
Co
Margetts, & Manning, 2011; Gatesy et al., 1999; Menezes, Araújo-

, & Medeiros, 2019; Platt, Hasiotis, & Hirmas, 2012).
Júnior, Dal' Bo
As described here, footprints are surrounded by an observed swol-
len and compressed border, indicating the squeezing of the water-
logged sediment due to the weight of the dinosaur. As the
footprints show impact structures and distortion of the ripple
marks with huge, contorted bedding, it is suggested that the sub-
strate is more humid with low cohesiveness and low water content
(Carvalho & Leonardi, 2020).
The lithofacies analysis of the beds containing footprints is inter-
preted as having been deposited in coastal plain/shallow marine to
delta plain environments under oxic conditions, where organic matter
preservation was rare. The fine grain size of the sediments that
10
formed the substrate over which the dinosaurs crossed was a signifi-
cant factor in footprint preservation.
The footprints are assigned to both sauropod and theropod dino-
saurs in both levels. The sauropod is dominant with 85 and 15% for
theropod on the lower level. In the upper level, there are only thero-
pod dinosaurs with 100% dominance.
The domain of sauropod and theropod footprints is attributed to
an ecological zonation of the dinosaurian biota. Both dinosaur types
are distributed in the low floodplain areas, where preservation possi-
bility is higher in the fine-grained sediments.
The largest theropod is known from footprint number 9 in the
lower level and footprint number 16 in the upper level, about 5 m
long. Moreover, they are the heaviest theropod with 847 kg. The larg-
est sauropod dinosaur reflected from its hip height is known from
footprint number 10 with a length of 3.8 m and the smallest sauropod
dinosaur based on its hip height is 1.5 m long, known from footprint
number 2.
We record here the presence of Parabrontopodus (Lockley
et al., 1994) and Brontopodus (Farlow et al., 1989), which indicate dif-
ferent sauropod ichnogenera.
Although the preservation of the footprints is poor, some of them
contain morphological characters. The preservation of the footprints
is affected by the cyclic accumulation of the sediments
(T. Thulborn, 1990) and rapid burial from the overlying sediments.
The direct relationship between the quality of a footprint and the
substrate consistency at the time they are formed (Carvalho, Borghi, &
Leonardi, 2013) is considered here, as all the footprints reported are
true footprints.
6 | C O N CL U S I O N S
Sixteen footprints have been found in pre-Campanian strata
deposited in coastal plain to delta plain environments, distributed
in the present-day Quseir area, Eastern Desert, Egypt. They sug-
gest that theropod and sauropod dinosaurs flourished in the inter-
connected area between the river, deltas, and sea during the Late
Cretaceous. These footprints consist of various types of small or
medium and large footprints that may reflect the occurrence of dif-
ferent genera. During the Cretaceous, it is possible that some dino-
saurs survived and inhabited the Quseir area. This discovery
reports the first evidence of the presence of dinosaurs in the south
Eastern Desert, Egypt.
ACKNOWLEDGEMEN TS
We would like to thank Cairo University and New Valley University
for their financial support in the field trip. The authors are thankful to
Prof. Dr. Etienne Jaillard (Grenoble University, France) for his critical
reviewing the manuscript and for his valuable comments. Lorenzo
Marchetti and anonymous reviewer and the Editor-in-Chief Prof. Ian
Somerville for their pertinent comments to an earlier version of the
typescript.
KASSAB ET AL.
PE ER RE VIEW
The peer review history for this article is available at https://publons.
com/publon/10.1002/gj.4357.
DATA AVAILABILITY STAT EMEN T
The data that support the findings of this study are available from the
corresponding author upon reasonable request.
OR CID
Walid G. Kassab https://orcid.org/0000-0001-9882-9607
Gebely A. Abu El-Kheir https://orcid.org/0000-0001-5905-8790
Mohamed K. AbdelGawad https://orcid.org/0000-0002-3229-3800
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