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First nonavian dinosaur from Lebanon: a brachiosaurid sauropod from the Lower Cretaceous of the Jezzine District


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Two sauropod teeth from an Early Cretaceous (Neocomian) fluviodeltaic sandstone near Jezzine (Southern Lebanon) are the first nonavian dinosaur remains to be reported from Lebanon. Their distinctive character places them within Brachiosauridae. The sauropod teeth from Lebanon are a significant addition to the very scanty dinosaur record from the Levant, which hitherto consisted mainly of very poorly preserved and not easily identifiable specimens. The Basal Cretaceous Sandstone of Lebanon, thus, appears to be a potentially important source of fossil vertebrate material.
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DOI 10.1007/s00114-006-0124-z
Eric Buffetaut .Dany Azar .André Nel .Kamil Ziadé .
Aftim Acra
First nonavian dinosaur from Lebanon: a brachiosaurid sauropod
from the Lower Cretaceous of the Jezzine District
Received: 17 November 2005 / Accepted: 9 April 2006
#Springer-Verlag 2006
Abstract Two sauropod teeth from an Early Cretaceous
(Neocomian) fluviodeltaic sandstone near Jezzine (South-
ern Lebanon) are the first nonavian dinosaur remains to be
reported from Lebanon. Their distinctive character places
them within Brachiosauridae. The sauropod teeth from
Lebanon are a significant addition to the very scanty
dinosaur record from the Levant, which hitherto consisted
mainly of very poorly preserved and not easily identifiable
specimens. The Basal Cretaceous Sandstone of Lebanon,
thus, appears to be a potentially important source of fossil
vertebrate material.
The published record of dinosaurs (sensu lato) from the
Levant (Lebanon, Syria, Israel, Palestine, and Jordan) is
extremely scanty, being based mostly on specimens that
cannot be determined with any accuracy, so that this area,
like the Middle East as a whole, is still largely a terra
incognita in terms of dinosaur biogeography. The record
includes theropod footprints from the Cenomanian of the
vicinity of Jerusalem, Israel (Avnimelech 1966), a fragmen-
tary theropod tibia from the Late Cretaceous of the vicinity
of Damascus, Syria (Hooijer 1968), a small fragment of a
tibia, probably from an ornithopod, from the Maastrichtian
phosphates of Roseifa, Jordan (Martill et al. 1996), and
feathers from the Early Cretaceous amber of Wadi Zerqa,
Jordan (Kaddumi 2005,pp3639). In Lebanon, birds are
among the various tetrapods known from the Cenomanian
sublithographic limestones of Nammoura (Dalla Vecchia
and Chiappe 2002), and feathers have been found in the
Early Cretaceous amber of Jezzine (Schlee 1973;Schleeand
Glöckner 1978) and Hammana, but no nonavian dinosaurs
had so far been reported. We describe, in this study, sauropod
teeth from the Lower Cretaceous of Southern Lebanon
which are the first nonavian dinosaur remains from that
country and, being identifiable to the family level,
considerably improve our knowledge of the Cretaceous
dinosaur faunas of the Middle East.
The material has been found in the Neocomian
sandstone of Southern Lebanon (Mouhafazit Loubnan El-
Janoubi), District of Jezzine (Caza Jezzine), in the locality
of Jouâr Es-Souss, and at the entrance of the villages of
Jezzine and Bkassine (Figs. 1and 2). The same outcrop
contains amber-bearing beds. This Basal Cretaceous
Sandstone (also known as Grès de Base) is considered
to have been deposited in a fluviodeltaic environment
(Maasaad 1976).
Description and comparisons
The material comprises two teeth (Fig. 3), one of which,
found in August 2005 by André Nel, is well preserved
while the other one, found in 1969 by Aftim Acra, is
incomplete. Both specimens are in the Natural History
Museum of the Lebanese University at Fanar, Lebanon.
E. Buffetaut (*)
CNRS (UMR 5125),
16 cour du Liégat, Paris 75013, France
D. Azar
Faculty of Science II, Natural Sciences Department,
Lebanese University,
Fanar, P.O. Box 26110217, Fanar-Matn, Lebanon
A. Nel
Laboratoire dEntomologie and CNRS UMR 8569,
Muséum National dHistoire Naturelle,
45 Rue Buffon, Paris 75005, France
K. Ziadé
Amine Gemayel Street, Awkar, P.O. Box 90-674,
Jdeideh, Lebanon
A. Acra
The Lakes Apartments, 5666 East Hampton Street,
Apartment 135, Tucson, AZ 85712, USA
The best specimen (JAzar 1), shows the well preserved
crown and part of the root. It is 23 mm in height and 8 mm
mesiodistally. The crown is hardly broader than the root
and there is no significant constriction between them. The
root, which is oval in cross-section, can be distinguished
from the crown mainly because its surface is smooth,
unlike the distinctly wrinkled enamel of the crown. The
enamel wrinkles form low irregular ridges which become
fainter toward the apex (probably because of wear, as
described by Janensch (1935)inBrachiosaurus). The
crown is strongly convex labially, both mesiodistally and
apicobasally. The lingual face is markedly concave
apicobasally and convex mesiodistally. Thus, the crown
cannot be described as spoon-shaped or spatulate, as its
lingual face is neither concave mesiodistally nor signifi-
cantly broadened relatively to the root. The mesial margin
of the crown is slightly concave at the base and slightly
convex near the apex. The distal margin is convex at the
base and slightly concave closer to the apex. As a result,
when seen in labial or lingual view, the crown appears
twisted. The lingual and labial faces of the crown are
separated by blunt carinae. The mesial one is only faintly
marked, especially in the basal part. The distal carina is
more distinct, being separated from the convex labial face
by a well-marked groove, which extends from the base to
the apex. A mesial groove is also present, but much fainter
than the distal one. Close to the apex, the distal carina bears
faint, heavily worn serrations. The apex of the tooth bears a
large oval wear facet, sloping markedly in a mesiolingual
direction, which has completely removed the enamel and
exposes the underlying dentine.
The second specimen, (JAcra 1), is a fragment of a
broader tooth (14 mm mesiodistally), consisting of the
apical portion of a crown, broken above the base, and
damaged at the apex. The labial face is convex mesiodis-
tally and apicobasally, while the lingual face is convex
mesiodistally and slightly concave apicobasally. The whole
surface of the tooth is much abraded and pitted, the enamel
being largely worn away, especially on the lingual face, so
that no wrinkling is apparent. Both the mesial and distal
carinae are worn and blunt. At the level of the break, the
fragment is subcircular, with a small pulp cavity. The apex
is irregularly broken, so it is uncertain whether a wear facet
was present. On the lingual side, there is a more or less
circular and rather deep pit, probably corresponding to a
wear facet, as facets in a similar position occur, for
instance, in Brachiosaurus (Janensch 1935).
Fig. 1 The outcrop of Neocomian sandstone at Jouâr Es-Souss,
Jezzine District, Southern Lebanon, where the sauropod teeth
described in the present paper have been found
Fig. 2 Geological map of the
Jezzine area (Southern Lebanon)
showing the location of the
Neocomian sauropod locality at
Jouâr Es-Souss (adapted from
Dubertret 1950). J6 Kimmerid-
gian; J7 Tithonian; C1 Neoco-
mian; C2a Barremian (?)Early
Aptian; C2b Late Aptian;
C3 Albian; C4 Cenomanian;
QQuaternary. Scale bar
represents 1 km
The teeth from Jouâr Es-Souss are different in size and
shape, but this may be the result of different positions in the
tooth row. The better preserved slender specimen is used in
this study for comparisons. The tooth can be described as a
rather simple, somewhat flattened and twisted cylinder,
indicating that it belongs to a sauropod dinosaur, a
conclusion supported by the wrinkled surface of the
enamel (a sauropod synapomorphy: Wilson and Sereno
1998). Although the tooth cannot be described as spoon-
shaped or spatulate, it differs from the typical peg-like teeth
of diplodocoids and titanosaurs, which are straighter and
more slender, with less strongly wrinkled enamel, and the
affinities of the Lebanese sauropod must be sought among
the broad-toothedsauropods. The twistedshape of the
crown indicates that the tooth is from the posterior part of
the jaw, as more anterior teeth have more symmetrical
crowns. The narrow and convex lingual face of the crown
distinguishes this tooth from the posterior teeth of most
broad-toothed sauropods, such as camarasaurids and
euhelopodids, in which the posterior tooth crowns are
broad and have a concave lingual face. The tooth from
Lebanon is strongly reminiscent of the posterior teeth of
brachiosaurids. In Brachiosaurus brancai from the Upper
Jurassic of Tanzania, the posterior teeth of the upper jaw
show the same twisted shape and their crown is not spatulate
(Janensch 1935, pl.XII, Fig. 1). Their enamel shows a similar
wrinkling and when worn, they bear an oblique apical wear
facet (JanenschsTerminal-Usurfläche), as on the Leba-
nese tooth. The main difference is that on the tooth from
Jezzine the distal groove on the labial face appears to be
much more distinct than in B. brancai. The dentition of other
brachiosaurids is poorly known. Isolated teeth from the Early
Cretaceous Arundel Formation of Maryland (USA), for
which the name Astrodon johnsoni must be used (Carpenter
and Tidwell 2005), were described by Marsh (1888,p.90)as
having crowns which are mainly compressed cones, and
not spoon-shaped. Some of them closely resemble the tooth
from Jezzine (Lull 1911, pl.XIV, Fig. 8). Astrodon is usually
placed among the Brachiosauridae (McIntosh 1990;
Weishampel and Young 1996; Naish and Martill 2001),
although Carpenter and Tidwell (2005) referred it to a family
incertae sedis of the Titanosauriformes (a group which
includes brachiosaurids).
From these comparisons, the slender tooth from Jezzine,
probably from the posterior part of a left upper tooth row, can
be referred to the Brachiosauridae, although no identification
at a lower taxonomic level is possible. The broader tooth, to
judge from its apparently more symmetrical crown, is
probably from a more anterior location in the tooth row, and
both teeth may well belong to the same taxon.
The teeth from Jezzine allow identification to the family
level and are, thus, more informative than the previously
reported dinosaur material from the Middle East. However,
they do not provide much new information about the exact
age of the Basal Cretaceous Sandstone of Lebanon, as
brachiosaurids are known from both the Late Jurassic and
the Early Cretaceous. All that can be said is that the
occurrence of a brachiosaurid is in accordance with the
Early Cretaceous (Neocomian) age of the Basal Cretaceous
Sandstone of Lebanon.
The ascertained geographical distribution of brachio-
saurids encompasses North America, Europe, and Africa
Fig. 3 Sauropod teeth from the Neocomian of Jouâr Es-Souss,
Southern Lebanon in the collection of the Natural History Museum of
the Lebanese University, compared with brachosaurid teeth. JAzar1
in mesial (a), lingual (b), distal (c), and labial (d) views. JAcra1 in
mesial (e), lingual (f), labial (g), and distal (h) views. IThe five
posteriormost teeth of the right upper tooth row of Brachiosaurus
brancai from the Late Jurassic of Tendaguru (Tanzania) in labial
(above) and lingual (below) views, after Janensch (1935, pl.XII,
Fig. 1). JIsolated tooth of Astrodon johnsoni from the Early
Cretaceous of Maryland (USA) in (from left to right) labial, mesial,
lingual, and distal views, after Lull (1911, pl.XIV, Fig. 8). All scale
bars represent 10 mm
(Upchurch et al. 2004); in addition, an isolated tooth from
the Lower Cretaceous of Korea has been referred to the
Brachiosauridae (Lim et al. 2001). In the Early Cretaceous,
the region of present-day Lebanon was in the northeastern
part of Gondwana, on the Afro-Arabian plate. There is no
well-ascertained record of brachiosaurids in the Early
Cretaceous of Gondwana. The brachiosaurid Brachio-
saurusnougaredi (Lapparent 1960) from Algeria, con-
sidered as Early Cretaceous by Upchurch et al. (2004) and
Weishampel et al. (2004), in fact comes from beds which
contain Late Jurassic plants (Boureau and Caillon 1958;
Lapparent 1960). Early Cretaceous sauropods so far
reported from Africa (see Weishampel et al. 2004) are
either indeterminate or belong to other groups, including
primitive eusauropods and rebacchisaurids from Niger
(Sereno et al. 1999) and titanosaurs from Malawi (Jacobs et
al. 1996), to which the teeth from Lebanon cannot be
referred. By contrast, brachiosaurids are well represented in
the Early Cretaceous of North America (Upchurch et al.
2004), with taxa such as Cedarosaurus,Sauroposeidon,
and Astrodon, and are also present in Europe, with material
from the Wealden of England that is mostly fragmentary,
although a fairly complete skeleton still awaits description
(Naish and Martill 2001). However, the record of Early
Cretaceous sauropods from Gondwana is still too sparse to
conclude that brachiosaurids were not present there, and it
would be premature to suggest that the teeth from Lebanon
indicate Euramerican rather than Gondwanan affinities.
The brachiosaurid teeth from the Early Cretaceous of
Jezzine are the first nonavian dinosaur remains from
Lebanon. The only vertebrates hitherto reported from the
Basal Cretaceous Sandstone were small fishes from a
dysodilic level (Janensch 1924) and feathers (Schlee 1973;
Schlee and Glöckner 1978) and a lizard (Arnold et al.
2002) preserved in amber. The Lebanese brachiosaur is an
important addition to the record of dinosaurs from the
Levant, and more generally from the Middle East, which is
still very scarce (see above). We fully concur with the
remark of Martill et al. (1996, p.153), to the effect that the
apparent scarcity (of dinosaurs in the Middle East) is
because so few vertebrate paleontologists have examined
the Mesozoic in this part of the world. The Basal
Cretaceous Sandstone of Lebanon is one of the formations
which are likely to provide more information about the
elusive dinosaurs of the Middle East, a vast area which was
part of the northern margin of Gondwana and for which
continental biogeographical data for the Cretaceous are still
extremely incomplete.
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... 9: 220381 17 [207][208][209]. Early Cretaceous ('Neocomian') strata in Lebanon have produced teeth that appear to be referrable to Brachiosauridae [64,87]. All of these records represent 'holdovers' from the Late Jurassic, based on comparisons with the sauropod fauna of the Tendaguru Formation of Tanzania [210,211]. ...
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... In northern and central Jordan the upper boundary is marked by the presence of pelagic chalk deposits of the Belqa Group (Wadi Umm Ghudran Formation), while to the south this facies is gradually replaced by chert, phosphatic, quartz-arenitic and dolomitic rocks. The Ajlun Group has been variously considered Albian-Early Cenomanian in age (Wetzel and Morton 1959;Bender 1974), or the top being late Turonian in age (Wetzel and Morton, 1959;Basha 1978). Six formations are recognized in this group (Fig. 1B). ...
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... Paleobotanists have also identified a new species of Weichselia, a Mesozoic fern, in Early Cretaceous strata at Makhtesh Ramon, in southern Israel (Silantieva and Krassilov, 2006). Discoveries of terrestrial fauna include a new species of snake and a brachiosaur in Cretaceous deposits in Lebanon, a salamander in similar strata in Israel, and mammal remains that are too numerous to cite (Buffetaut et al., 2005;Caldwell, 2006;Nevo and Estes, 1969). ...
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Eastern Anatolia, southern Iraq, and Jerusalem have been proposed as the regions that once contained the Garden of Eden. Several creationists have argued that it is impossible to locate the region of Eden due to the dramatic changes to the surface of the earth during the Noachian Deluge. However, a close analysis of relevant Biblical passages and the archaeology, geography, geology, paleontology, and paleobotany of Anatolia, the Near East, and North Africa suggest that the region of Eden was located in Southeastern Anatolia. This region provides source waters for four rivers, following Precambrian rift valleys or faults, which are connected to the Biblical locales of Asshur, Havilah, and Cush. The rivers traversed a landmass that originated in the Precambrian not covered by the alleged expanse of the Tethys Ocean. Furthermore, southeastern Anatolia is associated with the ancient kingdom of the House of Eden. Although the Bible states that the Flood was a global catastrophic event, the confluence of biblical texts with geographical, geological, and other data provides a compelling indication that its effects did not eradicate all evidence of Eden’s original location.
... The diversity of the species placed in Minyohelea suggests diversification of these biting midges in the Cretaceous, with probable specialisation on different hosts (7 species known) in the area of Lebanese amber formation. The fossil record of Lebanese early Cretaceous terrestrial vertebrates is scarce, but some remains of lizards (Arnold et al., 2002), pterosaurs (Kellner et al., 2019), dinosaurs (Buffetaut et al., 2006) and birds (Dalla Vecchia and Chiappe, 2003) were reported, making the possibility of relationships of haematophagous biting midges and these vertebrates plausible. ...
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... This sedimentary sequence is becoming internationally known thanks to a series of palaeontological works based on amber insect inclusions (Azar, 2007 and references therein) including up to 180 different species of insects. Buffetaut et al. (2016) recently discovered in laterally equivalent deposits located near the village of Jezzine (south Lebanon) two dinosaur teeth attributed to sauropods. ...
A rich microfossil assemblage composed of six charophyte species and fifteen ostracod taxa has been recovered from eight stratigraphic sections raised in Lower Cretaceous rocks of the Abeih Formation/upper part of the Grès de base unit in Mount Lebanon (Central-South Lebanon). Four depositional environments have been distinguished within this unit based on sedimentological and microfacies analyses: fluvial, coastal freshwater lakes, estuary (intertidal and subtidal) and shallow marine. The charophyte assemblage is composed of Sphaerochara asema, aff. Mesochara harrisii, Atopochara trivolvis var. trivolvis, Ascidiella reticulata, Clavator ampullaceus, and Munieria martinclosasi extracted from costal freshwater lakes facies. The associated ostracod fauna is mainly represented by Cypridea libanensis, C. heliopolisensis, C. aaleyensis recovered along with accessory carapaces of C. baidarensis, C. quadrangulata, Cypridea sp. 1, Cypridea sp. 2, and Timiriasevia cardiformis. Part of this freshwater microfossil assemblage (A. trivolvis var. trivolvis, A. reticulata, C. ampullaceus, Cypridea libanensis, C. heliopolisensis, and C. aaleyensis) has been found in facies related to brackish water characterized by marine ostracods such as Fastigatocythere aff. naftaliensis, Fastigatocythere sp., Metacytheropteron parnesi, Majungaella (Progonocythere?) sp. Pontocyprella sp., Asciocythere sp., and Damonella sp. Shallow marine elements are also abundant including dasycladalean thalli, foraminifera and echinoid remains. This study indicates that the Abeih Formation/upper part of the Grès de base unit is late Barremian/early Aptian in age. The microfossil assemblage is dominated by endemic to subcosmopolitan taxa characteristic of the Levantine Island at the southeastern margin of the Tethys during the Early Cretaceous.
... En plus, on trouve de l'argile, des nodules de fer (Bellos, 2008), du lignite, des laminites, de l'ambre extrêmement riche en fossiles et du basalte (Massaad, 1976 ;Azar et al., 2010). Dans ces niveaux se trouvent des plantes fossiles, des dents de poissons, des gastéropodes, des fragments d'ostracodes et des os de dinosaures (Heybrock et Dubertret, 1945 ;Buffetaut et al., 2006). Selon Blankenhorn ( (Tixier, 1971(Tixier, -1972. ...
Les dysodiles sont des roches sédimentaires formées de feuillets millimétriques riches en matière organique et en fossiles. Au Liban, malgré leur intérêt pétrolier et paléontologique, les dysodiles ont été très rarement étudié, gardant leur environnement de dépôt largement inconnu. Ce travail doctoral vise une approche interdisciplinaire sédimentologique, paléontologique et géochimique, afin de reconstruire le paléoenvironnement de dépôt de 5 sites de dysodiles du Barrémien inférieur du Liban. Une diversité de fossiles est exceptionnellement préservée à l’intérieur des dysodiles : des poissons, des tortues d’eau douce, des insectes, des gastéropodes, des ostracodes, des débris de végétaux et des coprolithes. Leur étude a permis de reconstruire un lac temporaire peu profond, influencé par le volcanisme, entouré d’une savane tropicale typique du Crétacé inférieur, et abritant des précurseurs de la faune continentale actuelle. Les ostracodes montrent des différences taxonomiques et taphonomiques entre les sites, et les palynomorphes sont dominés par les grains de pollen au Nord du Liban, contrairement aux autres sites où figure une diversité de spores de fougères. Les analyses optiques, géochimiques et moléculaires indiquent l’origine algaire et bactérienne de la matière organique (dominance de matière organique amorphe fluorescente, kérogène de type I, HI moyen = 659 mgHC/gTOC, et OI moyen = 30 mgCO2/gTOC, n-alcanes à chaine courte et hopanes), et prouvent que les dysodiles sont d’excellentes roches-mères potentielles (TOC entre 3 et 25 %). Toutefois à cause de leur extension limitée, l’intérêt pétrolier des dysodiles est supplanté par leur intérêt paléontologique.
... Paleobotanists have also identified a new species of Weichselia, a Mesozoic fern, in Early Cretaceous strata at Makhtesh Ramon, in Southern Israel (Silantieva and Krassilov, 2006). Discoveries of terrestrial fauna include a new species of snake and a brachiosaur in Cretaceous deposits in Lebanon, a salamander in similar strata in Israel, and mammal remains that are too numerous to cite (Caldwell, 2006;Buffetaut et al., 2005;Nevo and Estes, 1969). ...
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Archaeologist Joel Klenck describes the location of the region, which once contained the Garden of Eden. Included in this manuscript are reviews of the debate, analyses of proposed locations for Eden in Turkey, Iraq and Israel, and a discussion of the claim that the region cannot be located due to the effects of the Noachian Deluge. In addition, Biblical references elucidate the classification of groups of plants and animals at creation, the association of certain taxonomic groups, and infer the presence of at least three types of gardens. Correlations are proposed between these Biblical gardens and paleontological assemblages from alleged geological area. The creation of plant and animal kings and their placement in gardens throughout the earth, provides a potential explanatory model for paleontological assemblages. Here, plant and animal kinds increasingly overlap their alleged temporal boundaries and yet certain floral and faunal groups are commonly found associated with each other in geographical locales.
... In addition, the position of Charentes teeth in the tooth row is unknown. During the Early Cretaceous, brachiosaurids are recorded in North America (Tidwell et al., 1999(Tidwell et al., , 2001Chure et al., 2010;D'Emic et al., 2016) Nicosia et al. (2007) possibly Afro-Arabia (Buffetaut et al., 2006). Sonorasaurus thompsoni is the last known brachiosaurid occurrence from the late Albian/early Cenomanian (middle Cretaceous) (D'Emic et al., 2016), and Brachiosauridae had become globally extinct by the earliest Late Cretaceous . ...
Algora is a recently discovered vertebrate fossil site located in the Cenomanian strata of the Arenas de Utrillas Formation (Guadalajara, Central Spain). This fossil site is rich in turtle remains, but other clades of reptiles are also identified, including plesiosaurs, crocodyliforms and dinosaurs. The only remains of dinosaurs from Algora so far studied corresponded to theropod teeth. However, several sauropod remains are here presented and analyzed, revealing the presence of several shared features with members of Titanosauriformes, some common found in derived members belonging to Lithostrotia clade. This sauropod fossil assemblage corresponds to the richest sauropod fossil accumulation found in the Cenomanian of Europe. In addition, one of the first studies of the pneumatic features in dorsal ribs using CT scan is also provided here. The performed cladistic analysis places the Algora sauropod within Lithostrotia, in a more derived position than Malawisaurus (supported, for example, by the presence of convex posterior articulation up to the posterior section of the tail), in a clade which also includes Rapetosaurus (Maastrichtian, Madagascar) and the Italian indeterminate titanosaur MSNM V7157 (late Aptian–early Albian). The Algora titanosaur and MSNM V7157 are recognized as closely related. This analysis shows the presence of two different lineages (Algora titanosaur + MSNM V7157 and Normanniasaurus, from the Albian of France) in the Early and middle Cretaceous of Europe, and at least one with a Gondwanan origin. The close relationship of the Algora titanosaur and MSNM V7157 suggests a pre-late Aptian faunal contact between the Adriatic Plate and Laurasia, in accordance with several previous studies, which proposed faunal exchanges between Laurasia and Gondwana through the Adriatic plate during the Early Cretaceous.
... After the Jurassic, Early Cretaceous brachiosaurid remains are known of the USA, of southern Africa (McPhee et al., 2016), and possibly from Afro-Arabia too (Buffetaut et al., 2006), while Padillasarusu from South America has been considered a somphospondylan (Mannion et al., 2017). The described taxa came only from Early Cretaceous of the USA: Abydosaurus, Cedarosaurus, Sonorasaruus and Venenosaurus Mannion et al., 2017). ...
A new dinosaur sauropod from the Golmayo Formation (upper Hauterivian-lower Barremian) in Soria province (Spain) is described as Soriatitan golmayensis gen. et sp. nov.. The new material consists of one tooth, three dorsal vertebrae with ribs, a partial sacrum, five caudal centra, two caudal vertebral spines, one chevron, a humerus, an ulna, a radius, two partial ilia, two ischia, a fragment of pubis, and a partial femur. Cladistic analysis identified the material as belonging to Brachiosaruidae within Titanosauriformes. Sauropod material from the Golmayo Formation shares a diagnostic feature with Abydosaurus, Cedarosaurus, Tastavinsaurus and Venenosaurus including anteriorly deflected anterior–middle caudal neural spines and with Cedarosaurus a proximodistally straight lateral margin between the proximal head and the shaft of the humerus. Eight characters are potential autapomorphies in the sauropod from Spain, including the presence of a large rectangular ventral ridge below the preacetabular process of the ilium, the same length of the pubic and ischiadic blades in the ischium, and two lateral ridges in the lateral area of the deltopectoral crest of the humerus. The presence of Early Cretaceous brachiosarids in both, North America and Europe, give support to the hypothesis of a connection between the tectonic plates of these continents at some point during the Early Cretaceous.
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Brachiosauridae is a lineage of titanosauriform sauropods that includes some of the most iconic non-avian dinosaurs. Undisputed brachiosaurid fossils are known from the Late Jurassic through the Early Cretaceous of North America, Africa, and Europe, but proposed occurrences outside this range have proven controversial. Despite occasional suggestions that brachiosaurids dispersed into Asia, to date no fossils have provided convincing evidence for a pan-Laurasian distribution for the clade, and the failure to discover brachiosaurid fossils in the well-sampled sauropod-bearing horizons of the Early Cretaceous of Asia has been taken to evidence their genuine absence from the continent. Here we report on an isolated sauropod maxilla from the middle Cretaceous (Albian–Cenomanian) Longjing Formation of the Yanji basin of northeast China. Although the specimen preserves limited morphological information, it exhibits axially twisted dentition, a shared derived trait otherwise known only in brachiosaurids. Referral of the specimen to the Brachiosauridae receives support from phylogenetic analysis under both equal and implied weights parsimony, providing the most convincing evidence to date that brachiosaurids dispersed into Asia at some point in their evolutionary history. Inclusion in our phylogenetic analyses of an isolated sauropod dentary from the same site, for which an association with the maxilla is possible but uncertain, does not substantively alter these results. We consider several paleobiogeographic scenarios that could account for the occurrence of a middle Cretaceous Asian brachiosaurid, including dispersal from either North America or Europe during the Early Cretaceous. The identification of a brachiosaurid in the Longshan fauna, and the paleobiogeographic histories that could account for its presence there, are hypotheses that can be tested with continued study and excavation of fossils from the Longjing Formation.
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Sauropod material from the Arundel Formation (Aptian-Albian boundary) of Maryland has been variously referred to Astrodon johnsoni Leidy 1865 or to Pleurocoelus nanus Marsh 1888. Most of the specimens are juvenile as demonstrated by the small size of the bones, the lack of neurocentra 1 fusion, absence of an olecranon, and underdevelopment of muscle scars. Contrary to some recent statements, the Arundel sauropod is diagnostic. Only a single sauropod taxon is present in the Arundel Formation, to which the name Astrodon johnsoni must be used under the Principle of the First Reviser of the International Code of Zoological Nomenclature .
Africa became progressively isolated during the middle portion of the Cretaceous Period as it rifted apart from other Gondwana continents and the southern Atlantic Ocean was completed. Within this geological context, which controls the occurrence and distribution of Cretaceous fossil localities over most of Africa, there is considerable variety in composition and occurrence of fossils. In the Koum Basin of Cameroon in western Africa, abundant footprints of five morphotypes, but scrappy skeletal remains and teeth of the ornithopod Ouranosaurus and other taxa are found. In the Karonga area of northern Malawi, in southeastern Africa, more complete skeletal remains are known, including the sauropod Malawisaurus, which shows affinity to South American titanosaurids, and an undescribed sauropod taxon. These may demonstrate a range of morphological diversity in sub-Saharan Cretaceous sauropods comparable to that found in South America.