ArticlePDF Available

Abstract and Figures

We describe a giant titanosaurid sauropod dinosaur discovered in coastal deposits in the Upper Cretaceous Bahariya Formation of Egypt, a unit that has produced three Tyrannosaurus-sized theropods and numerous other vertebrate taxa. Paralititan stromeri is the first tetrapod reported from Bahariya since 1935. Its 1.69-meter-long humerus is longer than that of any known Cretaceous sauropod. The autochthonous scavenged skeleton was preserved in mangrove deposits, raising the possibility that titanosaurids and their predators habitually entered such environments.
Content may be subject to copyright.
29. The compilation is provided in the supplementary
material (38).
30. C. D. Gebelein, in Stromatolites, M. R. Walter, Ed.
(Elsevier, Amsterdam, 1976), pp. 499–515.
31. J. P. Grotzinger, J. F. Kasting, J. Geol. 101, 235 (1993).
32. J. Bertrand-Sarfati, M. R. Walter, Precambrian Res. 15,
353 (1981).
33. J. P. Grotzinger, D. H. Rothman, Nature 383, 423
(1996).
34. A. H. Knoll, I. J. Fairchild, K. Swett, Palaios 8, 512
(1993).
35. R. Riding, J. Geol. Soc. London 149, 979 (1992).
36. B. Laval et al., Nature 407, 626 (2000).
37. D. L. Parkhurst, D. C. Thorstenson, L. N. Plummer, U.S.
Geol. Surv. Wat.-Res. Invest. Rep. 80-96 (1990).
38. Supplementary data are available on Science Online
at www.sciencemag.org/cgi/content/full/292/5522/
1701/DC1.
39. We thank R. Riding and P. Westbroek for com-
ments on an earlier draft and the Pyramid Lake
Paiute Tribal Council and the Western Australian
Department of Conservation and Land Manage-
ment for sampling permits. Funded by the German
Research Foundation and the Studienstiftung des
deutschen Volkes (G.A.).
6 November 2000; accepted 10 April 2001
A Giant Sauropod Dinosaur
from an Upper Cretaceous
Mangrove Deposit in Egypt
Joshua B. Smith,
1
* Matthew C. Lamanna,
1
Kenneth J. Lacovara,
2
Peter Dodson,
1,3
Jennifer R. Smith,
1
Jason C. Poole,
4
Robert Giegengack,
1
Yousry Attia
5
We describe a giant titanosaurid sauropod dinosaur discovered in coastal de-
posits in the Upper Cretaceous Bahariya Formation of Egypt, a unit that has
produced three Tyrannosaurus-sized theropods and numerous other vertebrate
taxa. Paralititan stromeri is the first tetrapod reported from Bahariya since
1935. Its 1.69-meter-long humerus is longer than that of any known Cretaceous
sauropod. The autochthonous scavenged skeleton was preserved in mangrove
deposits, raising the possibility that titanosaurids and their predators habitually
entered such environments.
In the early 20th century, the Bavarian geologist
Ernst Stromer described a diverse biota from
the Upper Cretaceous [Cenomanian: 93.5 to
99.0 million years ago (Ma)] Bahariya Forma-
tion (1) of the Bahariya Oasis, Egypt (Fig. 1).
The vertebrate discoveries included fish, turtles,
plesiosaurs, squamates, crocodyliforms, and
four dinosaurs: the theropods Spinosaurus,
Carcharodontosaurus, and Bahariasaurus, and
the sauropod Aegyptosaurus (2). Tragically,
Stromer’s collections were largely destroyed
during an Allied bombing of Munich in 1944
(3). With exceptions from Morocco (46) and
Algeria (7), evidence of Late Cretaceous Afri-
can dinosaurs remains limited. An improved
understanding of Late Cretaceous African ter-
restrial vertebrates is important for the paleo-
ecology of this region and is needed to evaluate
biogeographic hypotheses pertaining to Gond-
wanan fragmentation (5, 810). Here we de-
scribe the partial skeleton of an extremely large
sauropod dinosaur, the first tetrapod reported
from Bahariya since 1935 (11). The specimen
consists largely of vertebrae, pectoral girdle,
and forelimb elements and is preserved in sed-
iments indicative of intertidal deposits. A num-
ber of morphological differences distinguish the
humerus of the specimen (Fig. 2A) from that of
Aegyptosaurus, precluding referral to that ge-
nus (12). Because of these distinctions and its
possession of several autapomorphies, we des-
ignate the new specimen as Paralititan stro-
meri, gen. et sp. nov. (13).
Two preserved caudal sacral centra of
Paralititan lack pleurocoels. The centrum of
the first caudal vertebra (Fig. 2B) is wider
than high and procoelous, and has a convex
distal articular condyle. The centrum is not
biconvex, as in the titanosaurids Alamosau-
rus (14), Neuquensaurus (14 ), and Pelle-
grinisaurus (15). Its ventral surface has
weakly developed longitudinal ridges lateral-
ly bordering a sagittal concavity. A postspi-
nal lamina is present between spinopostzyg-
apophyseal laminae on the distal surface of
the neural spine. A second proximal caudal
(Fig. 2C) is strongly procoelous and has a
well-developed distal condyle.
The scapula is concave medially. A prom-
inent dorsomedial rugosity borders the medi-
al concavity as in the titanosaurids Aeolo-
saurus, Lirainosaurus, Neuquensaurus, and
Saltasaurus (14, 16). Distal to the glenoid, a
well-developed tabular process projects from
the caudoventral margin of the scapula. The
development of this structure in Paralititan is
equaled only in a scapulocoracoid referred to
the brachiosaurid Brachiosaurus altithorax
(17).
The humerus is strongly expanded proxi-
mally and distally. Because of the modest de-
1
Department of Earth and Environmental Science,
University of Pennsylvania, 240 South 33rd Street,
Philadelphia, PA 19104 6316, USA.
2
Engineering Ge-
ology, Drexel University, 3141 Chestnut Street, Phil-
adelphia, PA 19104, USA.
3
Department of Animal
Biology, School of Veterinary Medicine, University of
Pennsylvania, 3800 Spruce Street, Philadelphia, PA
191046045, USA.
4
Academy of Natural Sciences,
1900 Ben Franklin Parkway, Philadelphia, PA 19103,
USA.
5
Egyptian Geological Museum, Egyptian Geolog-
ical Survey and Mining Authority, Athar El Nabi,
Maadi, Cairo, Egypt.
*To whom correspondence should be addressed. E-
mail: smithjb@sas.upenn.edu
Table 1. Phylogenetic data matrix. The macronarian Camarasaurus is postulated as an outgroup of titanosauriformes (45, 46). Character codings are as follows:
0, hypothesized plesiomorphic states; 1 and 2, hypothesized derived states; ?, missing or uncertain data (19).
Taxa
Characters
5 10152025303540455055
Outgroup
Camarasaurus 00000 00000 00000 00000 00000 00000 00000 00000 00000 00000 00000 0
Ingroup
Brachiosaurus brancai 11001 01000 00000 00000 00000 001?1 00000 001?0 00001 01010 01000 0
Andesaurus ??111 11000 1100? ?0?00 00??0 001?? ????? ????? ????? ??111 ????? ?
Epachthosaurus ??111 ??010 11011 ?0?01 1?011 1110? 1???? ???1? ???11 111?1 ?101? 0
Opisthocoelicaudia ???12 11101 ?1111 1?002 01110 10111 11011 01111 11111 11101 11011 0
Alamosaurus ????? ????? ????? ??101 10011 11101 11?11 101?1 1111? ?1??1 1???? ?
Malawisaurus 11?12 1???1 1100? ?0101 10011 00111 1???0 ????? ????? ????1 ????? 1
Paralititan ????? ????? ????? ??011 1101? ????? ??1?? ?111? ???1? ????? ????? ?
Saltasaurus ??112 11111 11001 11111 11101 11101 1?111 11111 1???1 11?11 ?1111 1
R EPORTS
1 JUNE 2001 VOL 292 SCIENCE www.sciencemag.org1704
velopment of a proximolateral process, the
proximal end is sinusoidal in cranial view, as in
Saltasaurus and Opisthocoelicaudia (18). A
well-defined muscular depression occupies the
proximal region of the cranial surface. The
deltopectoral crest is extremely developed and
medially deflected, extending more than 53%
of the length of the element. The rectangular
radial condyle is well developed distally. A
shallow fossa, bounded by medial and lateral
ridges, occupies most of the proximocaudal
surface of the humerus. The lateral ridge devel-
ops into a tuberosity approximately 350 mm
from the proximal margin, as in Lirainosaurus
and Saltasaurus (16), whereas the medial ridge,
unknown in titanosaurids, is most developed
560 mm from the proximal end. Elongate su-
pracondylar ridges that extend over one-third of
the length of the humerus define an extensive
olecranon fossa. The distal surface of the pre-
served metacarpal is rectangular and flattened,
suggesting phalangeal reduction or absence on
this digit.
Phylogenetic analysis of titanosauriform
sauropods (19) places Paralititan within Ti-
tanosauridae (Fig. 3 and Table 1). Characters
supporting its inclusion within the clade include
strongly procoelous proximal caudal centra
with well-developed postspinal laminae, a
proximolateral process on the humerus, and
reduced manual phalanges. The position of
Paralititan within Titanosauridae permits an
estimation of its body size (Fig. 2D). The hu-
merus is 1.69 m in length, 14% longer than
the next longest known humerus from a Creta-
ceous sauropod (20). The South American ti-
tanosaurid Argentinosaurus is regarded as the
most massive terrestrial animal known and may
have approached 90 metric tons (21) and 30 m
in length (22). The humerus of Argentinosaurus
is unknown, but we estimate its length at 1.81 m
(7.5% longer than that of Paralititan), using
more complete titanosauriforms (23). There-
fore, Paralititan is probably not as large as
Argentinosaurus but represents one of the
heaviest terrestrial vertebrates yet discovered.
Fig. 1. (A) The Bahariya Oasis, 300 km south-
west of Cairo, Egypt. (B) Location of BDP 2000-
18.
Fig. 2. Anatomy, taphonomy, and estimated size of Paralititan.(A) Right humerus in cranial and
caudal views. (B) First caudal vertebra in distal and right lateral views. (C) Proximal caudal
vertebra in proximal and right lateral view. Scale bars in (A) through (C) equal 10 cm. (D) Size
comparisons between Paralititan and an African elephant. (E) Quarry map of BDP 2000-18.
Fig. 3. Summary of
phylogenetic, temporal,
and geographic relation-
ships among titano-
sauriform sauropods.
The cladogram depicts
the strict consensus of
12 most parsimonious
trees (length, 72 steps;
consistency index, 0.806;
retention index, 0.759)
resulting from a parsi-
mony analysis of eight
titanosauriforms and
56 anatomical charac-
ters (19).
R EPORTS
www.sciencemag.org SCIENCE VOL 292 1 JUNE 2001 1705
Paralititan is preserved in low-energy
paralic sediments, representing vegetated tidal
flats and tidal channels. These units alternate
vertically and are laterally variable. The tidal flat
facies is a brown friable shale, rich in rhizoliths
and plant remains, often containing leaf com-
pressions and stems of the mangrove (24) tree
fern Weichselia reticulata (25, 26 ). In some
places the facies is found, conformably, above
glauconitic nearshore marine sands, a strati-
graphic relationship that supports a mangrove
interpretation. The relationship appears to be
conformable, as evidenced by a lack of ravine-
ment surfaces between occasional rhizoliths that
extend from lagoonal muds into marine sands.
This succession of environments is known to
occur along modern low-energy coasts, where
mangroves prograde out onto the active shore
face (27, 28). The seaward migration of man-
groves into the open marine realm requires both
a low-energy littoral zone and wave-resistant
salt-tolerant plants, such as Weichselia.
The Paralititan quarry spans both tidal
channel and vegetated tidal flat facies (Fig. 2E).
In situ plant roots, throughout the bone layer,
indicate limited water depth, and fine-grained
sediments suggest low current velocities. The
closely associated elements could not have
been transported to this location as clasts. In
addition, the shallow, vegetated tidal flat would
prevent a large sauropod carcass from float-
ing to this location. Evidence therefore indi-
cates that this individual walked to this loca-
tion, over tidal flats and along tidal channels,
before its death. Furthermore, the specimen
shows indications of being scavenged by a
carnivorous dinosaur (29). Therefore at least
two species of Bahariya dinosaur traversed
paralic environments.
It is interesting that this extremely produc-
tive biota, containing some of the largest known
terrestrial vertebrates (30), occurs during a time
with extremely low thermal gradients from pole
to pole and high global sea levels (31, 32).
Mid-Cretaceous ocean-atmosphere systems are
of particular interest in paleoclimate research,
representing extreme “hothouse” conditions for
post-Pangaean Earth history. The apparently
high productivity of this Cenomanian environ-
ment may reflect a biotic response to some
aspect of this condition.
References and Notes
1. W. Dominik, Berl. Geowiss. Abh. Reihe A 62, 1 (1985).
2. E. Stromer, Abh. Bayer. Akad. Wiss. Math. Nat. Abt.
N. F. 33, 1 (1936).
3. Material from only 2 of Stromer’s 15 Bahariya tetra-
pods survived.
4. R. Lavocat, Comptes rendus 19e Congre´s ge´ologique
international (Alg.) 1952, Academie des Sciences de
Paris 15, 65 (1954).
5. P. C. Sereno et al., Science 272, 986 (1996).
6. D. A. Russell, Bull. Mus. Natl. Hist. Nat. Paris 4th Ser.
18, 349 (1996).
7. P. Taquet, D. A. Russell, C. R. Acad. Sci. Paris 327, 347
(1998).
8. S. D. Sampson et al., Science 280, 1048 (1998).
9. S. D. Sampson, M. T. Carrano, C. A. Forster, Nature
409, 504 (2001).
10. D. W. Krause, G. V. R. Prasad, W. v. Koenigswald, A.
Sahni, F. E. Grine, Nature 390, 504 (1997).
11. Other new Bahariya vertebrates include remains of
chondrichthyans, osteichthyans, turtles, plesiosaurs,
squamates, crocodyliforms, and dinosaurs. Theropod
remains represent Spinosaurus, cf. Carcharodonto-
saurus, and two indeterminate forms. Sauropod re-
mains include an indeterminate partial skeleton and a
possible rebbachisaurid scapula.
12. Although the holotype of Aegyptosaurus (1912VIII61)
was destroyed, and comparison with Paralititan is thus
difficult, several characters distinguish them. Specimen
1912VIII61 was substantially smaller (humerus length
59% that of Paralititan), may have had pleurocoelous
proximal caudals (33), possessed a weakly medially
convex scapula with no dorsomedial prominence, had a
humerus with a weak proximomedial expansion and
more medially positioned deltopectoral crest restricted
to the proximal third of the element, and lacked the
autapomorphies of Paralititan. Stromer (33) tentatively
referred several isolated elements to Aegyptosaurus,
including an indeterminate vertebra (1912VIII66), two
possible caudal cervicals (1912VIII67), and an isolated
procoelous caudal (1912VIII65). Because two Bahariya
Formation titanosaurids are now recognized, this ma-
terial referred to Aegyptosaurus must be considered
Titanosauria incertae sedis.
13. Etymology: Paralititan stromeri (paralititan
stromeri): paralos (Greek), near the sea (paralic refers
to tidal environments); Titan (Greek), an offspring of
Uranus and Gaea, symbolic of brute strength and large
size (effectively, “tidal giant”); stromeri, in honor of
Ernst Stromer. Holotype: CGM 81119 (Egyptian Geo-
logical Museum, Cairo). Locality: BDP 2000-18, near
Gebel Fagga (28°2010.7⬘⬘N latitude, 28°5904.7⬘⬘E
longitude). Material: Two fused caudal sacral vertebrae
(probably 5 and 6), first caudal vertebra; proximal cau-
dal vertebra; dorsal and sacral ribs; incomplete scapulae;
complete right and incomplete left humeri; distal meta-
carpal; and several additional elements. Diagnosis: Very
large titanosaurid characterized by proximal caudal cen-
tra wider than tall; prominent tabular process on cau-
doventral margin of scapula distal to the proximal ex-
pansion; and humerus with medial ridge on the proxi-
mocaudal face and rectangular radial condyle. Paraliti-
tan shares the following characters with other
titanosaurids: lack of pleurocoels on caudal sacrals;
proximal caudal vertebrae with strongly concave prox-
imal articular surface, well-developed distal articular
condyle, ventral excavation, and postspinal lamina; hu-
merus with proximolateral process and strong supra-
condylar ridges; and reduction or absence of manual
phalanges. Referred material: Stromer (33) described a
very large cranial dorsal vertebra (1912VIII64). Speci-
men 1912VIII64 was opisthocoelous, pleurocoelous,
and caudally wider than tall, as in Epachthosaurus and
Pellegrinisaurus (15), and may pertain to Paralititan.
14. L. Salgado, R. A. Coria, J. O. Calvo, Ameghiniana 34,3
(1997).
15. L. Salgado, Ameghiniana 33, 355 (1996).
16. J. L. Sanz, J. E. Powell, J. Le Loeuff, R. Martinez, X.
Pereda-Suberbiola, in Geology and Palaeontology of the
Upper Cretaceous Vertebrate-Bearing Beds of the Lan˜o
Quarry (Basque-Cantabrian Region, Iberian Peninsula),
H. Astibia, J. C. Corral, X. Murelaga, X. Orue-Etxebarria,
X. Pereda-Suberbiola, Eds. (Estudios del Museo de Cien-
cias Naturales de Alava, Alava, Portugal, 1999), pp.
235–255.
17. B. D. Curtice, K. L. Stadtman, L. J. Curtice, in The
Continental Jurassic, M. Morales, Ed. (Museum of
Northern Arizona, Flagstaff, AZ, 1996), pp. 87–95.
18. P. Upchurch, Zool. J. Linn. Soc. 124, 43 (1998).
19. The character list is available as Appendix 1 of the
supplementary information (34).
20. Humerus lengths of large Cretaceous sauropods are
as follows: Jobaria, 1360 mm (35); Chubutisaurus,
1450 mm (36); Aegyptosaurus, 1000 mm (33); Argy-
rosaurus, 1435 mm (estimate) (37); Titanosaurus?
colberti, 1480 mm (38).
21. G. S. Paul, in Dinofest International, D. L. Wolberg, E.
Stump, G. D. Rosenberg, Eds. (Academy of Natural
Sciences, Philadelphia, PA, 1997), pp. 129–154.
22. C.-M. Jianu, D. B. Weishampel, Geol. Mijnbouw 78,
335 (1999).
23. See Web fig. 1 (34).
24. Mangroves are vegetated paralic environments, de-
fined by Thanikaimoni (39) as intertidal tropical
forests. Mangroves first developed in the Carbon-
iferous (360 to 286 Ma), when salt-tolerant gym-
nosperms and pteridophytes adapted to the habi-
tat (40, 41). Modern mangroves include members
of 27 plant genera (42) and first developed along
the Late Cretaceous Tethys Seaway (43).
25. Paradoxopteris stromeri has been synonymized with
Weichselia reticulata (44).
26. Shinaq and Bandel (45) identified an Early Cretaceous
Weichselia mangrove in Jordan.
27. R. A. Davis, A. C. Hine, E. A. Shinn, in Quaternary Coasts
of the United States: Marine and Lacustrine Systems,J.F.
Wehmiller, C. H. Fletcher III, Eds. (Society for Sedimen-
tary Geology (SEPM), Tulsa, OK, 1992), pp. 193–212.
28. Additional evidence of low-energy environments in-
cludes fine-grained sediments, low-velocity bed-
forms, the absence of ravinement surfaces, in situ
bivalves, and horizontal burrows. All nonfossil clasts
in the Bahariya system are medium sand size or
smaller.
29. A cf. Carcharodontosaurus tooth (crown height
65 mm), recovered from BDP 2000-18, is larger
than clasts that could have been transported to
this location.
30. Paralititan coexisted with three Tyrannosaurus-sized
carnivores (Carcharodontosaurus, Bahariasaurus, and
Spinosaurus) and with other large vertebrates (such
as the 3.5-m coelacanth Mawsonia and the 10-m
crocodyliform Stomatosuchus).
31. B. U. Haq, J. Hadenbol, P. R. Vail, Science 235, 1156
(1987).
32. M. L. Fassell, T. J. Bralower, in Evolution of the Cre-
taceous Ocean Climate System, E. Barrera, C. C. John-
son, Eds. (Geological Society of America, Boulder, CO,
1999), vol. 332, pp. 121–142.
33. E. Stromer, Abh. Bayer. Akad. Wiss. Math. Nat. Abt.
N. F. 10, 1 (1932).
34. Supplementary Web material is available on Science
Online at www.sciencemag.org/cgi/content/full/292/
5522/1704/DC1
35. P. C. Sereno et al., Science 286, 1342 (1999).
36. L. Salgado, Ameghiniana 30, 265 (1993).
37. R. Lydekker, Ann. Mus. La Plata Sect. Paleontol. 2,1
(1893).
38. S. L. Jain, S. Bandyopadhyay, J. Vert. Paleontol. 17,
114 (1997).
39. G. Thanikaimoni, in Mangrove Palynology (Travaux
de la Section scientifique et technique, UNDP/
UNESCO Regional Project on Training and Research
on Mangrove Ecosystems, French Institute, Pondi-
cherry, India, 1987) pp. 1–100.
40. A. A. Cridland, Palaeontology 7, 186 (1964).
41. A. Raymond, T. L. Phillips, in Biology and Ecology of
Mangroves, H. Teas, Ed., vol. 8 of Tasks for Vegetative
Science (Junk, the Hague, 1983), pp. 19–30.
42. N. C. Duke, in Tropical Mangrove Ecosystems,A.I.
Robertson, D. M. Alongi, Eds. (American Geophysical
Union, Washington, DC, 1992), pp. 63–100.
43. A. M. Ellison, E. J. Farnsworth, R. E. Merkt, Global Ecol.
Biogeogr. 8, 95 (1999).
44. E. Boureau, A. F. de Lapparent, C. R. Somm. Seances
Soc. Geol. Fr. 7-8, 107 (1951).
45. R. Shinaq, K. Bandel, Freib. Forschungsh. C 474,39
(1998).
46. J. A. Wilson, P. C. Sereno, Soc. Vertebr. Paleontol.
Mem. 5, 1 (1998).
47. Supported by Cosmos Studios, MPH Entertainment,
the University of Pennsylvania, the Andrew K. Mellon
Foundation, E. de Hellebranth, and the Delaware
Valley Paleontological Society. We thank K. Soleiman
and the Egyptian Geological Survey and Mining Au-
thority, Drexel University, T. Holtz, H. Sues, J. Harris,
K. Curry-Rogers, J. Lacovara, and R. Martı´nez. We also
thank A. Tumarkin, Y. Abdelrazik, M. Said Abdel-
Ghani, P. Kane-Vanni, and J. Caton for assistance in
the field.
9 March 2001; accepted 30 April 2001
R EPORTS
1 JUNE 2001 VOL 292 SCIENCE www.sciencemag.org1706
... Nevertheless, despite the rich and ever-increasing middle and Late Cretaceous non-avian dinosaur record of Egypt (e.g. [5][6][7][8]), only highly fragmentary evidence of Abelisauridae has as yet come to light from this nation and northeastern Africa in general. At present, the only unquestioned abelisaurid fossil from Egypt is an isolated tooth from an exposure of the uppermost Cretaceous (Campanian-Maastrichtian) Duwi Formation near Idfu in the southern Nile Valley region [9,10]. ...
... Erectopus sauvagei by Stromer [5,13] and may have affinities with Ceratosauria and clades therein. Intriguingly, several of these forms-namely Spinosaurus, Carcharodontosaurus and Bahariasaurus/Deltadromeus-attained exceptionally large body sizes among non-avian theropod dinosaurs, comparable to that of Tyrannosaurus rex [6,20]. Though it undoubtedly pertains to a substantially smaller-bodied animal, the new abelisaurid vertebra (MUVP 477) confirms the presence of a fourth medium-sized to large (approximately 6 m in total body length) theropod taxon in the Bahariya Formation palaeoecosystem ( figure 5). ...
... If so, then these distinctions may be directly linked to palaeoenvironmental differences between these units. In particular, the predominant dinosaur-bearing horizons of the Bahariya Formation appear to represent a comparatively low-energy paralic habitat dominated by the mangrove tree fern Weichselia reticulata [6,68] (figure 5), whereas those of the Kem Kem Group ( particularly the Gara Sbaa Formation) were apparently deposited in a substantially higher-energy fluvial system in which the complete disarticulation and dissociation of skeletal elements prior to and during entombment was common [20]. Further taphonomic studies of continental vertebrates from the Bahariya Formation and the Kem Kem Group are needed to evaluate these hypotheses. ...
Article
Full-text available
Numerous non-avian theropod dinosaur fossils have been reported from the Upper Cretaceous (Cenomanian) Bahariya Formation, Bahariya Oasis, Western Desert of Egypt, but unambiguous materials of Abelisauridae have yet to be documented. Here we report Mansoura University Vertebrate Paleontology Center (MUVP) specimen 477, an isolated, well-preserved tenth cervical vertebra of a medium-sized abelisaurid from the Bahariya Formation. The new vertebra shows affinities with those of other Upper Cretaceous abelisaurids from Madagascar and South America, such as Majungasaurus crenatissimus , Carnotaurus sastrei , Viavenator exxoni and a generically indeterminate Patagonian specimen (Museo Padre Molina specimen 99). Phylogenetic analysis recovers the Bahariya form within Abelisauridae, either in a polytomy of all included abelisaurids (strict consensus tree) or as an early branching member of the otherwise South American clade Brachyrostra (50% majority rule consensus tree). MUVP 477, therefore, represents the first confirmed abelisaurid fossil from the Bahariya Formation and the oldest definitive record of the clade from Egypt and northeastern Africa more generally. The new vertebra demonstrates the wide geographical distribution of Abelisauridae across North Africa during the middle Cretaceous and augments the already extraordinarily diverse large-bodied theropod assemblage of the Bahariya Formation, a record that also includes representatives of Spinosauridae, Carcharodontosauridae and Bahariasauridae.
... The Cenomanian vertebrate fossils of Bahariya Oasis are collected from the association facies 1 of Catuneanu et al. (2006) of the base of Bahariya Formation at Gebel El Dist, north Bahariya Oasis (Stromer, 1931;Smith et al. 2001). ...
... 7D, 7E and 7F). Smith et al. (2001) discovered partial skeleton of the large Sauropod dinosaur Paralatitine stroemeri from the tidal flat and tidal channels facies of Bahariya Formation of Bahariya Oasis. ...
Preprint
Full-text available
The detailed stratigraphical field relations and taphonomic conditions and assessment of the Upper Cretaceous vertebrate bearing horizons in Western Desert of Egypt were investigated in the three sites; Campanian Quseir Formation, Kharga Oases, Maastrichtian Ammonite Hill member of Dakhla Shale, Abu Minqar and Lower Cenomanian Bahariya Formation in Gebel El Dist, Bahariya Oases. It was found that the most important taphonomic factors controlling the preservation state of vertebrate fossils in the selected sites are rate of transportation and sedimentation as well as bio-erosion and diagenetic events of the bearing sediments. A comparative study of these factors was achieved between the investigated vertebrates bearing sites. The study indicated that the vertebrate fossils in Bahariya and Kharga Oases are more preserved and less abraded than these in Abu Minqararea. Generally, the study vertebrate fossils that yielded in sandstones have better preserved state than other lithological types. Furthermore, the deposition in low rate of transportation and high rate of sedimentation give the vertebrate carcasses good opportunity for the rapid burying with articulated and semi articulated elements, consequently better preservation state. Due to the importance of the vertebrate bearing low topographic sites in the Western Desert, its highly recommended to carry out serious and decisive mitigation for the protection against negative impacts that caused by human activities (e.g. illegal land reclamation and ecotourism) which leading to groundwater rise in these lowlands.
... The Cenomanian vertebrate fossils of Bahariya Oasis are collected from the association facies 1 of Catuneanu et al. (2006) of the base of Bahariya Formation at Gebel El Dist, north Bahariya Oasis (Stromer, 1931;Smith et al. 2001). ...
... 7D, 7E and 7F). Smith et al. (2001) discovered partial skeleton of the large Sauropod dinosaur Paralatitine stroemeri from the tidal flat and tidal channels facies of Bahariya Formation of Bahariya Oasis. ...
Article
Full-text available
The detailed stratigraphical field relations and taphonomic conditions and assessment of the Upper Cretaceous vertebrate bearing horizons in Western Desert of Egypt were investigated in the three sites; Campanian Quseir Formation, Kharga Oases, Maastrichtian Ammonite Hill member of Dakhla Shale, Abu Minqar and Lower Cenomanian Bahariya Formation in Gebel El Dist, Bahariya Oases. It was found that the most important taphonomic factors controlling the preservation state of vertebrate fossils in the selected sites are rate of transportation and sedimentation as well as bio-erosion and diagenetic events of the bearing sediments. A comparative study of these factors was achieved between the investigated vertebrates bearing sites. The study indicated that the vertebrate fossils in Bahariya and Kharga Oases are more preserved and less abraded than these in Abu Minqararea. Generally, the study vertebrate fossils that yielded in sandstones have better preserved state than other lithological types. Furthermore, the deposition in low rate of transportation and high rate of sedimentation give the vertebrate carcasses good opportunity for the rapid burying with articulated and semi articulated elements, consequently better preservation state. Due to the importance of the vertebrate bearing low topographic sites in the Western Desert, its highly recommended to carry out serious and decisive mitigation for the protection against negative impacts that caused by human activities (e.g. illegal land reclamation and ecotourism) which leading to groundwater rise in these lowlands.
... "Godzilla" (1954) leaves that mind-boggling conundrum unresolved, the single most vital question not even posed by its otherwise wondering actors and actresses [8]. "Turd" may seem vulgar, but even today, especially at numerous places where raw sewage streams off the landscape, all too many of those fragrant and disgusting capsule-like putrefying objects are visibly afloat in Rio de Janeiro's allegedly beloved Guanabara Bay! Considering that Cretaceous dinosaurs fed on ancient shoreline mangroves [9], that survival stratagem for enrapturing Guanabara Bay-emerging 'Guanazilla' does not arise to real-world intellectual impact. ...
Article
Full-text available
Symbolic transformation of "Guanazilla" (a melding of an indifferent and ignorant public with NWO groupthink from professional politicians' mindsets) from villain to magnificent virtuousness is a transition that is positively well-worth seeking. It is posited here that Guanabara Bay can again become a naturally functional coastal marine ecosystem! The purely fanciful model "Guanazilla" is used as a mentated or metaphorical approach to address the mega-problem of potential environmental harm to the upper-bay's precious mangrove stands. This metaphor is applied with a purposeful yet playful attitude and a curious perspective. This unique perspective, which provides a forward-looking expression for both the mega-mangrove problem and its potential remediation, is offered for consideration without apology. Because Guanabara Bay is a Brazilian cultural keystone, it is prudent to avoid the development of an unwanted and degraded "Guanazilla." This avoidance of potential eco-catastrophe is possible via implementation of environmental training that will ensure the iconic mangrove's proper long-term maintenance, protection, preservation, and enhancement. Resumo: A transformação simbólica de "Guanazilla" (uma fusão de um público indiferente e ignorante com o pensamento de grupo da NWO da mentalidade de políticos profissionais), de vilania para virtuosidade magnífica, é uma transição que vale a pena procurar. Postula-se aqui que a Baía de Guanabara pode voltar a se tornar um ecossistema marinho costeiro naturalmente funcional! O modelo puramente fantasioso "Guanazilla" é usado como uma abordagem mental ou metafórica para lidar com o megaproblema de possíveis danos ambientais aos preciosos manguezais da baía superior. Essa metáfora é aplicada com uma atitude propositiva, porém lúdica, e uma perspectiva curiosa. Essa perspectiva única, que fornece uma expressão voltada para o futuro, tanto para o problema dos megamanguezais quanto para sua possível remediação, é oferecida para consideração sem desculpas. Como a Baía de Guanabara é uma pedra angular da cultura brasileira, é prudente evitar o desenvolvimento de uma "Guanazilla" indesejada e degradada. Essa prevenção de uma potencial catástrofe ecológica é possível por meio da implementação de treinamento ambiental adequado que garantirá a manutenção, proteção, preservação e enriquecimento do manguezal icônico no longo prazo. Palavras-chave: Macro-engenharia, Baía de Guanabara, Rio de Janeiro, Godzilla film, resíduos plásticos, limpeza da água do mar, preservação dos manguezais.
... Previous records of Moroccan plesiosaurs come from Late Cretaceous marine settings, including the Turonian Akrabou Formation Buchy et al., 2005;Angst and Bardet, 2016) and the Maastrichtian phosphates (Vincent et al., 2011;Vincent et al., 2013). The fossils reported here are the first freshwater occurrence of plesiosaurs from Morocco, although plesiosaurs occur in a paralic setting (Smith et al., 2001) in the coeval Bahariya Formation of Egypt (Stromer, 1935). Benton et al. (2000) report a plesiosaur vertebra from the ?Albian Chenini Formation of Tunisia, also a fluvial system. ...
Article
Plesiosaurs were a long-lived and widespread group of marine reptiles, with a worldwide distribution and a temporal range from the Late Triassic to the Late Cretaceous. Most occur in marine deposits, but some occur in low-salinity, brackish to freshwater environments. We report plesiosaurs from the freshwater fluvial deposits of the mid-Cretaceous (?Albian-Cenomanian) Kem Kem Group of Morocco. Remains include numerous shed teeth, vertebrae, and a humerus. The humerus represents a young juvenile; vertebrae likely belong to sub-adults. Teeth show heavy wear, similar to teeth of co-occurring spinosaurids. While coeval plesiosaurs from the Bahariya Formation of Egypt are members of Polycotylidae, the Kem Kem fossils show features of Leptocleididae, small-bodied plesiosaurs that were widely distributed in nearshore and non-marine settings in the Early Cretaceous. These fossils are the first freshwater plesiosaurs from Morocco, and are among the youngest representatives of Leptocleididae. The Kem Kem leptocleidids could have been infrequent visitors from the sea, freshwater-tolerant, or even freshwater-adapted, as in modern river dolphins. The abundance of shed teeth in the Kem Kem Group supports the hypothesis that they had some degree of freshwater tolerance. Furthermore, leptocleidids occur almost exclusively in shallow nearshore, brackish, or freshwater environments, suggesting adaptation to shallow, low-salinity environments. Other plesiosaur groups and other Mesozoic marine reptiles, including teleosaurids and mosasaurids, also occur in freshwater settings, suggesting plesiosaurs and other marine reptiles frequently exploited non-marine environments.
... Rebbachisaurids and somphospondylans coexisted during the early Albian in Tunisia [230][231][232] and the late Albian-early Cenomanian in Morocco [5, 213,227,[233][234][235][236][237]. Titanosaurs were evidently established in western Africa (Mali) before the Cenomanian [238], and the only sauropods present in Cenomanian deposits in Egypt (Aegyptosaurus, Paralititan) are titanosaurs [239,240]. Finally, the Namba Member of the Galula Formation in Tanzania has produced the titanosaurs Shingopana and Rukwatitan [241,242]; however, the age of this stratigraphic unit is certain, with palaeomagnetic analysis indicating either a Cenomanian-Santonian or Campanian age [243]. ...
Article
Full-text available
The Upper Cretaceous Winton Formation of Queensland, Australia, has produced several partial sauropod skeletons, but cranial remains—including teeth—remain rare. Herein, we present the first description of sauropod teeth from this formation, based on specimens from three separate sites. An isolated tooth and a dentary fragment from the Diamantinasaurus matildae type locality are considered to be referable to that titanosaurian taxon. A single tooth from the D. matildae referred specimen site is similarly regarded as being part of that individual. Seventeen teeth from a new site that are morphologically uniform, and similar to the teeth from the two Diamantinasaurus sites, are assigned to Diamantinasauria. All sauropod teeth recovered from the Winton Formation to date are compressed-cone-chisel-shaped, have low slenderness index values (2.00–2.88), are lingually curved at their apices, mesiodistally convex on their lingual surfaces, and lack prominent carinae and denticles. They are markedly different from the chisel-like teeth of derived titanosaurs, more closely resembling the teeth of early branching members of the titanosauriform radiation. This provides further support for a ‘basal’ titanosaurian position for Diamantinasauria. Scanning electron microscope microwear analysis of the wear facets of several teeth reveals more scratches than pits, implying that diamantinasaurians were mid-height (1–10 m) feeders. With a view to assessing the spatio-temporal distribution of sauropod tooth morphotypes before and after deposition of the Winton Formation, we provide a comprehensive continent-by-continent review of the early titanosauriform global record (Early to early Late Cretaceous). This indicates that throughout the Early–early Late Cretaceous, sauropod faunas transitioned from being quite diverse at higher phylogenetic levels and encompassing a range of tooth morphologies at the start of the Berriasian, to faunas comprising solely titanosaurs with limited dental variability by the end-Turonian. Furthermore, this review highlights the different ways in which this transition unfolded on each continent, including the earliest records of titanosaurs with narrow-crowned teeth on each continent.
... Although the soil mechanical properties and other conditions supporting sauropods are not known, it is reasonable to assume that the soil must have received ample precipitation to support growth of vegetation, and foraging ranges explored by these giants were likely small considering the challenge of locomotion over wet soils. While resolving this paradox is beyond the scope of this study, such an intrinsic constraint must have imposed certain ecological adaptations to sustain ample vegetation and sauropod mobility, such as walking on well-compacted pathways while browsing away from the path or life partially suspended in water and foraging along the margins of water bodies [such as postulated in Smith et al. (47) concerning feeding on mangroves along ancient shorelines]. Both hypothesized strategies favor animals with long necks (44), a characteristic of sauropods that supported the successful evolution of these giants. ...
Article
Full-text available
Significance Mechanization has transformed agriculture over the past century, greatly improving crop production efficiency. However, the higher capacity has resulted in increased farm vehicle weights. We show that while machinery design maintains constant surface contact stresses, an insidious and largely overlooked threat of subsoil compaction has developed. We demonstrate that modern vehicles induce high soil stresses that now exceed critical mechanical thresholds for many arable soils, inducing chronic soil compaction in root zones below tillage depths and adversely affecting soil functioning. We draw parallels between modern farm vehicles and the heaviest animals that walked Earth: sauropods. We hypothesize that these prehistoric giants may have induced subsoil compaction, thus presenting a paradox for productivity of the land that supported them.
... In anterior view, the femoral head is rounded and dorsomedially directed, rising well above the level of the greater trochanter, as in most Somphospondyli (Curry Rogers, 2009;Poropat et al., 2016;Carballido et al., 2017). Posteriorly, a wide and concave surface separates the femoral head from the greater trochanter, forming an angle of ~120° with the lateral margin of the femur (Fig. 23F), as in Bonatitan, Daxiatitan, Dongbeititan, Huabeisaurus, Paralititan Smith et al., 2001 andYunmenglong (Pang &Cheng, 2000;Smith et al., 2001;Martinelli & Forasiepi, 2004;Wang et al., 2007;You et al., 2008;Lü et al., 2013). ...
Article
Osteological knowledge of the sauropod dinosaur Ligabuesaurus leanzai is increased by the description of new postcranial elements assigned to the holotype MCF-PVPH-233. Furthermore, a newly referred specimen, MCF-PVPH-228, is recognized after a detailed revision of the abundant sauropod material collected from the Lohan Cura Formation outcrops in the Cerro de los Leones locality (southern Neuquén Basin, Patagonia, Argentina). Recent laboratory preparation and fieldwork allowed us to recognize several new morphological features of the pectoral and pelvic girdles and the cervical and caudal anatomy. Thus, a new diagnosis of Ligabuesaurus is proposed that includes new autapomorphies and a unique combination of features. A phylogenetic analysis based on this new material recovers Ligabuesaurus as a non-titanosaurian somphospondylan, more derived than Sauroposeidon. Therefore, we discuss the palaeobiogeographical implications for the diversification and distribution of South American somphospondylans, especially in the Neuquén Basin, which are closely related to the early stages of evolution of Titanosauria. In this context, Ligabuesaurus represents one of the more complete Early Cretaceous Titanosauriformes and the earliest non-titanosaurian somphospondylan of South America. Finally, the new information on Ligabuesaurus contributes not only to reconstruction of the sauropod faunal composition of south-western Gondwana, but also sheds light on the early stages and emergence of titanosaurians.
... Unlike other Ibero-Armorican titanosaurs, the Abditosaurus humerus exhibits a distally expanded deltopectoral crest, a synapomorphy of Saltasauridae 44,46 shared with Opisthocoelicaudia 41 , Saltasaurus 47 , N. australis 48 and Alamosaurus 49 . The distal condyles are clearly divided as in saltasaurids 44,46 , and the articular surface of the radial condyle faces anterodistally, as in Paralititan 34 and Saltasaurus 47 . The radius (Fig. 2l) has a mediolaterally expanded distal end that is bevelled posterolaterally more than 20° relative to the long axis of the shaft and a well-defined interosseus ridge, as in other lithostrotian titanosaurs 44 . ...
Article
Full-text available
The origin of the last sauropod dinosaur communities in Europe and their evolution during the final 15 million years of the Cretaceous have become a complex phylogenetic and palaeobiogeographic puzzle characterized by the controversy on the alleged coexistence of immigrant, Gondwana-related taxa alongside relictual and insular clades. In this context, we describe a new titanosaurian sauropod dinosaur, Abditosaurus kuehnei gen. et sp. nov., from the Late Cretaceous (Maastrichtian) Tremp Group of Catalonia (Spain). Phylogenetic analyses recover Abditosaurus separately from other European titanosaurs, within a clade of otherwise South American and African saltasaurines. The affinity of the new taxon with southern landmasses is reinforced by spatiotemporal co-occurrence with Gondwanan titanosaurian oospecies in southern Europe. The large size and the lack of osteohistological features potentially related to insular dwarfism or size reduction support the idea that Abditosaurus belongs to an immigrant lineage, unequivocally distinct from some of the island dwarfs of the European archipelago. The arrival of the Abditosaurus lineage to the Ibero–Armorican Island is hypothesized to have occurred during the earliest Maastrichtian (70.6 Ma), probably as a result of a global and regional sea-level drop that reactivated ancient dispersal routes between Africa and Europe. The arrival of large-bodied titanosaurs to the European archipelago produced dramatic changes in its insular ecosystems and important evolutionary changes in its dinosaur faunas, especially with respect to the ‘island rule’ effect.
... 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., 2016Sallam et al., , 2018. The presence of dinosaurs in the Baharyia, Dakhla, Kharga, and Baris oases throughout both the Cenomanian and Campanian 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). ...
Article
Full-text available
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 discovery is the second record of tetrapod footprints in Egypt and the first record of tetrapod footprints in the Eastern Desert.
Article
Full-text available
Consistent with geophysical evidence for the breaking up of Pangaea, it has been hypothesized that Cretaceous vertebrates on progressively isolated landmasses exhibit generally increasing levels of provincialism, with distinctly heightened endemism occurring at the beginning of the Late Cretaceous. The Cretaceous fossil record from the southern supercontinent of Gondwana has been much too poor to test this hypothesis with regards to mammals (Fig. 1 ). Early Cretaceous mammals are known only from isolated sites in Argentina, Australia,, Cameroon, and Morocco. Apart from several occurrences in South America, knowledge of Late Cretaceous Gondwanan mammals is limited to a single site in India that previously yielded a few specimens of placental mammals,, and a site in Madagascar that previously yielded only one indeterminate tooth fragment. Here we report the occurrence of a highly specialized and distinctive group of extinct mammals, the Sudamericidae (Gondwanatheria), in the Late Cretaceous of Madagascar and India. These new records comprise the first evidence of gondwanatheres outside South America and the first indication of cosmopolitanism among Late Cretaceous Gondwanan mammals. Antarctica may have served as an important Cretaceous biogeographic link between South America and Indo-Madagascar.
Article
Full-text available
Lower Cretaceous fossils from central Niger document the succession of sauropod dinosaurs on Africa as it drifted into geographic isolation. A new broad-toothed genus of Neocomian age (∼135 million years ago) shows few of the specializations of other Cretaceous sauropods. A new small-bodied sauropod of Aptian-Albian age (∼110 million years ago), in contrast, reveals the highly modified cranial form of rebbachisaurid diplodocoids. Rates of skeletal change in sauropods and other major groups of dinosaurs are estimated quantitatively and shown to be highly variable.
Article
The Florida peninsula contains five distinct coastal sections. The east coast barrier system is wave-dominated and has a large cuspate foreland (Cape Canaveral) near its middle. The Florida Keys and reef tract represent the only coastal carbonate system in the continental United States. An open-marine mangrove coast characterizes the low-energy, tide-dominated southwest part of the State. The central Gulf barrier system displays a mixed-energy morphology in a microtidal, low-energy setting. The open-coast marsh system of the Big Bend area that is north of the barrier system is also tide dominated, and is developed on a sediment-starved carbonate platform. -from Authors
Chapter
The Florida peninsula contains five distinct coastal sections, each resulting from its own spectrum of coastal processes and sediment availability during a slowly rising, late Holocene sea level. The east coast barrier system is wave-dominated and has a large cuspate foreland (Cape Canaveral) near its middle. The Florida Keys and reef tract represent the only coastal carbonate system in the continental United States. An open-marine mangrove coast characterizes the low-energy, tide-dominated southwest part of the State. The central Gulf barrier system displays a mixed-energy morphology in a microtidal, low-energy setting. The open-coast marsh system of the Big Bend area that is north of the barrier system is also tide dominated, and is developed on a sediment-starved carbonate platform. The oldest preserved coastal Holocene section is the Florida Keys area where, at about 6 to 8 ka, sequences accumulated during the Holocene. Most of the remainder of the peninsular coast is characterized by terrigenous sequences less than 3 ka. The younger sequences accumulated almost exclusively from reworking of older strata without benefit of additional sediment supply from land.
Article
Chubutisaurus insignis Del Corro (1975) from the Aptian Gorro Frigio Formation) of Chubut Province, Argentina, is a large sized dinosaur having controversial phylogenetic relationships within the Sauropoda. Orginally it was regarded as a member of a new family (Chubutisauridae) because of the amphiplatyan caudal vertebrae. Chubutisaurus insignis was then distinguished from the well-known and widely reported Late-Cretaceous titanosaurids, characterized by procoelous caudal vertebrae. Finally, it was suggested to include Chubutisaurus insignis within the Brachiosauridae because it shares several characters with Brachiosaurus and other members of the group. The holotype of Chubutisaurus insignis is here redescribed and interpreted. Probable affinities of this fossil are discussed adding information to the faunal characteristics of the mid-Cretaceous of Gondwana. -from Author
Chapter
The morphological and paleogeographic evidence suggesting that cordaitean-dominated swamps from the early Westphalian D (= 295 m.y.b.p.) of Iowa grew in salt water swamps is reviewed. Cordaitean trees, Psaronius tree ferns, and Medullosa seed ferns contributed 95% of the peat in coal-ball samples taken from three Iowa mines. Ordination analysis shows that associations dominated by Psaronius and Medullosa succeeded an association dominated by cordaitean trees at each site. Based on comparisons with Upper Carboniferous compression flora deposits, Psaronius and Medullosa seem to have grown in freshwater or only slightly brackish water habitats. The root morphology of cordaitean trees from Iowa and the paleogeographic position of these swamps suggest that cordaitean trees which produced Nucellangium glabrum and Cardiocarpus spinatus seeds grew in salt water. Results of the ordination analysis suggest that cordaitean trees which produced Mitrospermum seeds grew in freshwater or only slightly brackish habitats. The methods used to assess these deposits as mangrove peats can be applied to any structurally preserved peat. Most other putative mangrove communities in the fossil record have been described from compression-flora deposits. The non-morphological criteria for assessing these floras as mangrove swamp deposits are reviewed.
Article
The earliest record of titanosaurids anywhere in the world was established in India in 1877. Further collections from fossiliferous pockets near Jabalpur were made by C. E Matley during 1917–1919. This material formed the basis of a number of taxa from the Indian Late Cretaceous, even though most of the bones were isolated and fragmented. New titanosaurid skeletal material (collected during 1984–1986) described here represents part of an individual in associated and mostly articulated condition, though skull, hind limb and foot bones are missing. Paucity of associated skeletal remains of titanosaurids anywhere in the world makes the present find valuable in understanding these specialized sauropods. A new taxon, Titanosaurus colberti, is erected for the reception of the new material. The genus Titanosaurus is diagnosed and three species are identified from India: T. indicus, T. blanfordi, and T. madagascanensis.
Article
New occurrences of spinosaurid dinosaurs from the Aptian of Niger and Albian of Algeria augment our knowledge of Spinosaurus maroccanus and permit the description of a new taxon: Cristatusaurus lapparenti from the Gadoufaoua locality (Niger). Spinosaurs may be separated into two distinct groups: one with longirostrine muzzles and premaxillae which are rounded in cross-section, and a second with brevirostrine muzzles possessing a dorsoposteriorly expanded crest. The spanish genus Pelecanimimus may be related to spinosaurids, rendering spinosaurs a more diversified group than previously thought.