Articles

https://doi.org/10.1038/s41559-017-0455-5

New Egyptian sauropod reveals Late Cretaceous

dinosaur dispersal between Europe and Africa
Hesham M. Sallam   1*, Eric Gorscak   2,3,4, Patrick M. O’Connor   3,5, Iman A. El-Dawoudi1,
Sanaa El-Sayed1, Sara Saber1,6, Mahmoud A. Kora1, Joseph J. W. Sertich7, Erik R. Seiffert8 and
Matthew C. Lamanna   9

Prominent hypotheses advanced over the past two decades have sought to characterize the Late Cretaceous continental
vertebrate palaeobiogeography of Gondwanan landmasses, but have proved difficult to test because terrestrial vertebrates
from the final ~30 million years of the Mesozoic are extremely rare and fragmentary on continental Africa (including the then-
conjoined Arabian Peninsula but excluding the island of Madagascar). Here we describe a new titanosaurian sauropod dinosaur,
Mansourasaurus shahinae gen. et sp. nov., from the Upper Cretaceous (Campanian) Quseir Formation of the Dakhla Oasis of
the Egyptian Western Desert. Represented by an associated partial skeleton that includes cranial elements, Mansourasaurus is
the most completely preserved land-living vertebrate from the post-Cenomanian Cretaceous (~94–66 million years ago) of the
African continent. Phylogenetic analyses demonstrate that Mansourasaurus is nested within a clade of penecontemporaneous
titanosaurians from southern Europe and eastern Asia, thereby providing the first unambiguous evidence for a post-Cenomanian
Cretaceous continental vertebrate clade that inhabited both Africa and Europe. The close relationship of Mansourasaurus to
coeval Eurasian titanosaurians indicates that terrestrial vertebrate dispersal occurred between Eurasia and northern Africa
after the tectonic separation of the latter from South America ~100 million years ago. These findings counter hypotheses that
dinosaur faunas of the African mainland were completely isolated during the post-Cenomanian Cretaceous.

N
umerous palaeobiogeographic studies have proposed
hypotheses to explain the nature of the terrestrial ver-
tebrate faunas that inhabited continental Africa dur-
ing the post-Cenomanian Cretaceous (PCC; ~94–66  million
years ago)1–9. Some of these works have argued that the
African mainland was home to an endemic terrestrial verte-
brate assemblage that was isolated from other land areas on an
‘island continent’1–3,6, whereas others have postulated the exis-
tence of Late Cretaceous dispersal routes between Africa and
neighbouring landmasses that would have led to the devel-
opment of faunal commonalities among these areas4,7,8,10.
In particular, many studies have proposed the existence of biotic
connections between northern Africa and southern Europe
based largely on the presence of continental vertebrate taxa
with hypothesized Gondwanan affinities in terminal Cretaceous
deposits of the latter region4,8,10,11. Nevertheless, in the absence
of anatomically and phylogenetically informative terrestrial ver-
tebrate fossils from the PCC of continental Africa (including the
then-conjoined Arabian Peninsula but excluding Madagascar)
these hypotheses have remained essentially untestable.
Here we present a new titanosaurian sauropod dinosaur from
the Late Cretaceous (Campanian) of Egypt that is represented by the
most complete terrestrial vertebrate skeleton yet discovered from
the PCC of the African mainland. Phylogenetic analysis of this com-
paratively complete and informative taxon provides an opportunity
to test hypotheses of biotic connections between northern Africa
and southern Europe during the PCC.
Results
Systematic palaeontology.
Sauropoda Marsh, 1878
Titanosauria Bonaparte and Coria, 1993
Lithostrotia Upchurch, Barrett and Dodson, 2004
Mansourasaurus shahinae gen. et sp. nov.
Etymology. ‘Mansoura’, for Mansoura University in Mansoura,
Egypt, home institution of the research collaborative that
undertook the field and laboratory work; ‘sauros’, Greek, lizard.
‘shahinae’ honours M. Shahin for her contributions to the foun-
dation of the Mansoura University Vertebrate Paleontology
Center (MUVP).
Holotype. MUVP 200, an associated partial skeleton consisting of
cranial fragments, dentaries, cervical and dorsal vertebrae and asso-
ciated ribs, scapulocoracoid, sternal plate, humeri, radius, metacar-
pal III, metatarsals I, III and II or IV, probable partial osteoderms,
and several unidentified fragments.
Locality. North of the road between Mut and Balat, Dakhla Oasis,
Western Desert of Egypt (Fig. 1a).
Horizon. Upper member of the Upper Cretaceous (Campanian12–16)
Quseir Formation.

1
Mansoura University Vertebrate Paleontology Center, Department of Geology, Faculty of Science, Mansoura University, Mansoura, Egypt. 2Department of
Biological Sciences, Ohio University, Athens, OH, USA. 3Ohio Center for Ecology and Evolutionary Studies, Ohio University, Athens, OH, USA. 4Integrative
Research Center, The Field Museum of Natural History, Chicago, IL, USA. 5Department of Biomedical Sciences, Ohio University Heritage College of
Osteopathic Medicine, Athens, OH, USA. 6Department of Geology, Faculty of Science, Assiut University, Assiut, Egypt. 7Department of Earth Sciences,
Denver Museum of Nature and Science, Denver, CO, USA. 8Department of Integrative Anatomical Sciences, Keck School of Medicine, University of
Southern California, Los Angeles, CA, USA. 9Section of Vertebrate Paleontology, Carnegie Museum of Natural History, Pittsburgh, PA, USA.
*e-mail: sallam@mans.edu.eg

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Articles
a b
Mediterranean Sea
North
Mansoura
Cairo
Sinai
er
Riv
Libya
Bahariya Oasis
Nile
Egypt
Re
Left dentary
d
Dakhla Oasis
Se
a
Kharga Oasis
Right dentary
200 km
Skull
fragments
Sudan
Fig. 1 | Location, quarry map and skeletal reconstruction of Mansourasaurus shahinae gen. et sp. nov. (MUVP 200). a, A map of Egypt showing the
location of the Dakhla Oasis, the type locality of Mansourasaurus (indicated by the dashed box). b, A site map showing the disposition of the skeletal
elements in situ. c, A skeletal reconstruction in right lateral view, showing the preserved elements. The colour-coding in b and c is as follows: red, skull;
purple, postcranial axial skeleton; blue, appendicular skeleton; green, osteoderm fragments.
Diagnosis. Mansourasaurus shahinae is regarded as a lithostrotian
titanosaur on the basis of the following suite of synapomorphies:
coracoid with squared anteroventral corner17,18; sternal plate
length exceeds 70% that of humerus17,19; humeral distal condyles
divided17,18; distal end of radius bevelled ~20 degrees relative to long
axis17,18,20. Mansourasaurus is diagnosed by the following autapo-
morphies: ten dentary tooth positions; dentary symphyseal region
with pronounced ventral projection (‘chin’) that comprises about
one-third of the dorsoventral depth of the anterior end of the bone;
Meckelian groove faces predominantly ventrally; anterior to middle
cervical centrum with foramen in posterior portion of lateral sur-
face; anteroposterior length of parapophysis in anterior to middle
cervical vertebra subequal to that of centrum; foramen in capi-
tulotubercular web of anterior cervical rib; distal radius four times
broader mediolaterally than anteroposteriorly.
Discussion
Osteological description. The skeleton of Mansourasaurus shahinae
is the most complete terrestrial vertebrate specimen known from the
PCC of the entire African continent (Fig. 1b,c). Although a few other
African PCC sauropod fossils have been described, the majority con-
sist of isolated, poorly informative bones21,22; the two exceptions in
this regard are the holotypic scapula and forelimb of Angolatitan from
the Turonian of Angola23 and a partial hind limb of an unidentified
titanosauriform from the Maastrichtian of Morocco24. Consisting of
parts of the cranial and postcranial skeletons, the Mansourasaurus
specimen represents a titanosaurian individual estimated at 8–10 m
in length (Fig. 1c; Table 1). Although the neurocentral sutures of the
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Metacarpal III
North Dorsal vertebra
Dorsal ribs
Metatarsal I Sternal plate
Dorsal ribs
Dorsal ribs
Coracoid Scapula Dorsal vertebra
Cervical vertebrae Metatarsal III
Humerus
Cervical vertebra
Radius
Humerus
1m
vertebrae are fused, the scapula and coracoid are not, so we estimate
that MUVP 200 was not skeletally mature at death.
Two cranium fragments and both dentaries are preserved
(Fig. 2a–f). The left dentary is nearly complete, whereas the right
preserves only the anterior end. The dentary curves medially, ren-
dering the anterior part of the lower jaw parabolic in dorsal view
(Fig. 2c), comparable to the condition in many other titanosauri-
ans25–29 but unlike the squared snout of several South American
titanosaurs30. The dentary of Mansourasaurus possesses ten alveoli,
a uniquely low number within Titanosauria (the Ampelosaurus den-
tary preserves nine alveoli but is slightly incomplete26). The symphy-
seal region exhibits an unusually well-developed ‘chin’ (Fig. 2d–f).
The symphysis is perpendicular to the long axis of the dentary, as
in many other titanosaurians27–30. The Meckelian groove occupies
the ventromedial surface of the dentary, opens mainly ventrally and
persists for most of the length of the bone (Fig. 2e).
The three preserved cervical vertebrae are opisthocoelous,
anteroposteriorly short and internally composed of camellate bone
(Fig. 2g–i; see Supplementary Information). The neural arches and
spines are tall relative to those of comparably positioned cervicals
of most other titanosaurians. The lateral surface of the centrum is
excavated by a fossa that occupies much of its length. In the anterior
to middle cervical vertebra (Fig. 2h), the posterior region of the lat-
eral surface exhibits a foramen anterior to the cotyle that is unique
to Mansourasaurus among titanosaurians. In at least the two ante-
rior-most cervicals, the parapophysis is more than half the length
of the centrum, and in the anterior to middle cervical vertebra, it
extends nearly the entire centrum length, an autapomorphy of
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whereas the other vertebra preserves the centrum and part of the
Table 1 | Measurements (mm) of Mansourasaurus shahinae gen.
neural arch (Fig. 2k). The opisthocoelous anterior dorsal centrum
et sp. nov. (MUVP 200)
lacks a ventral keel and has a shallow, subcircular cotyle. The antero-
Dentary posteriorly elongate lateral fossa (pleurocoel) is not subdivided by
Anteroposterior length (along straight line) 182L internal laminae. The other dorsal vertebra preserves the anterior
and posterior centrodiapophyseal laminae and a weakly developed
Dorsoventral height, symphysis 40.8L, 36.0R
lamina dividing the centrodiapophyseal fossa.
Dorsoventral height, ramus 24.9L, 29.0R The right scapulocoracoid is nearly complete (Fig. 2l). On the
Anterior cervical vertebra basis of their preservation, the dorsal margins of the coracoid and
Anteroposterior length, centrum (with condyle) 147 the acromial process of the scapula would have been flush, or at
Dorsoventral height, centrum, posterior 69
least at the same height. The depth of the acromial region is more
than 275% that of the anterior scapular blade (Table 1), contrasting
Dorsoventral height, total 235 with the low acromion exhibited by some other titanosaurians31,32.
Middle to posterior cervical vertebra The blade is D-shaped in cross-section with a medial concav-
Anteroposterior length, centrum (with condyle) 200+​ ity that is demarcated ventrally by a ridge that is also present in
Dorsoventral height, centrum, posterior 127 Ampelosaurus26 and Lirainosaurus32. Mansourasaurus also has a
dorsomedial tubercle on the anterior scapular blade that occurs in
Dorsoventral height, total 330
Lirainosaurus32. The scapula contributes more to the glenoid than
Anterior dorsal vertebra does the coracoid, unlike in Lirainosaurus (coracoid contributes
Anteroposterior length, centrum (with condyle) 148 more than scapula32) or Opisthocoelicaudia (subequal contribution31).
Dorsoventral height, centrum, anterior 104 The span of the coracoid is greater dorsoventrally than anteropos-
teriorly, as in Ampelosaurus, but in Opisthocoelicaudia this condi-
Transverse width, centrum, anterior 85
tion is reversed. The infraglenoid lip is not as well developed as
Scapulocoracoid R
in Opisthocoelicaudia31. The coracoid foramen is patent along the
Anteroposterior length, total 730 scapulocoracoid suture, as in a few other titanosaurs32,33. The right
Dorsoventral height, scapula, glenoid–acromion 428 sternal plate (Fig. 2m) is crescentic and similar to those of other
Dorsoventral height, scapula, anterior end of blade 128 titanosaurians and closely related titanosauriforms17–19. Its length
is roughly 76% that of the humerus, a ratio comparable to other
Dorsoventral height, scapula, posterior end of blade 90
lithostrotians17.
Anteroposterior length, coracoid 215 The proximolateral corner of the humerus (Fig. 2n) is squared,
Dorsoventral height, coracoid 340 as in most titanosauriforms17. The deltopectoral crest is slightly
Sternal plateR mediolaterally expanded at its distal extreme, projects anteriorly
and persists until approximately midshaft (Table 1). The ulnar and
Anteroposterior length 470
radial condyles are moderately developed anteroposteriorly and dis-
Mediolateral width, anteroposterior midline 262 tinct from one another. The anterior portion of the radial condyle
HumerusL is not divided, as in other titanosaurians17. The proximal end of the
Proximodistal length 620 nearly complete left radius is mediolaterally expanded but damaged
Proximodistal length, deltopectoral crest 275
(Fig. 2o). The distal end is about four times wider mediolaterally
than anteroposteriorly (Fig. 2p; Table 1); we consider this autapo-
Mediolateral width, midshaft 120 morphic of Mansourasaurus since most other titanosaurians do not
Mediolateral width, distal 75 exceed a ratio of approximately 2.0. The distal margin is distolater-
RadiusL ally bevelled at 20 degrees to the horizontal, as in saltasaurids17,18.
Proximodistal length 380+​ The distal end of left metacarpal III is mediolaterally expanded and
bevelled slightly distolaterally (Fig. 2q). Of the hind limb, only the
Mediolateral width, midshaft 47
right metatarsals I (Fig. 2r) and III (Fig. 2s) and a metatarsal II or
Mediolateral width, distal 124 IV (probably also right) are preserved. The mediolateral axis of the
Anteroposterior breadth, distal 30 distal end of metatarsal I is bevelled. The proximal end of the stout
Metacarpal IIIL metatarsal III is subovoid in outline, similar to that of Notocolossus34.
Several flattened, elongate fragments with irregular outlines and
Mediolateral width, distal 54
interwoven bone texture probably represent pieces of osteoderms.
Metatarsal I R

Proximodistal length 41 Phylogenetic and palaeobiogeographic analyses. We assessed

Mediolateral width, midshaft
Metatarsal IIIR
Proximodistal length
Mediolateral width, midshaft
+​: element incomplete, measurement as preserved; L: left; R: right.
Mansourasaurus. The capitulotubercular web of the anterior cervi-
cal rib is pierced by a foramen that represents another autapomorphy
of Mansourasaurus (Fig. 2g). Two incomplete dorsal vertebrae are
preserved (Fig. 2j,k). The more anterior dorsal vertebra consists of
the centrum and the posterior centrodiapophyseal lamina (Fig. 2j),
14 the phylogenetic affinities of Mansourasaurus using the data
set and analytical protocols presented in a previous study9 (see
Methods and Supplementary Information). Mansourasaurus was
96 recovered within a clade of penecontemporaneous (Campanian–
39 Maastrichtian) saltasaurid titanosaurians that are otherwise
known only from Europe and Asia (Fig. 3). Within this clade,
Mansourasaurus is the sister taxon of the European Lohuecotitan,
with a clade consisting of the European Lirainosaurus and the
central Asian Nemegtosaurus and Opisthocoelicaudia as the sister
taxon of Lohuecotitan +​  Mansourasaurus. A clade comprised of
the European Ampelosaurus and Paludititan is the sister taxon of
the clade containing all of these other Afro-Eurasian titanosaurs.
Although clade posterior probabilities vary widely across the
topology—probably due to missing data and the lack of overlap

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Articles Nature Ecology & Evolution

a c d e al10 al1 f

al1 mg
ns
b ch ch
ch
al10
sprl
bt? ns
prz
poz
pcdl
g h i prdl j
lf
ns cprl
cpol
prz ct
ct
pcdl

ct ct
pp
for for
cr cr pp
acp
pp
cr

k l
scb
cof
dr sc

al

ct
cd
cor

sct

vp igl
gl
o
m n
ca dpc

10 cm

r s

rc
uc

Fig. 2 | Skeletal anatomy of Mansourasaurus shahinae gen. et sp. nov. (MUVP 200). a, Skull fragment (frontal or parietal?) in ?dorsal view. b, Braincase
fragment in ?posterior view. c, Dentaries in dorsal view. d–f, Left dentary in lateral (d), medial (e) and anterior (f) views. g, Anterior cervical vertebra
in right lateral view. h, Anterior to middle cervical vertebra in right lateral view. i, Middle to posterior cervical vertebra in right lateral view. j, Anterior
dorsal vertebra in ventral view. k, Dorsal vertebra (with superimposed dorsal rib) in left anterolateral view. l, Right scapulocoracoid in lateral view
(with reconstructed dorsal margins of acromial region and coracoid indicated by dashed line). m, Right sternal plate in ventral view. n, Left humerus in
anterior view. o,p, Left radius in anterior (o) and distal (p) views. q, Distal left metacarpal III in anterior view. r, Right metatarsal I in lateral view. s, Right
metatarsal III in dorsal view. acp, acromial process; al, accessory lamina; al1, alveolus 1; al10, alveolus 10; bt?, basal tuber?; ca, coracoid articulation;
cd, condyle; ch, ‘chin’ of dentary; cof, coracoid foramen; cor, coracoid; cpol, centropostzygapophyseal lamina; cprl, centroprezygapophyseal lamina;
cr, cervical rib; ct, cotyle; dpc, deltopectoral crest; dr, dorsal rib; for, foramen; gl, glenoid; igl, infraglenoid lip; lf, lateral fossa (=​pleurocoel); mg, Meckelian
groove; ns, neural spine; pcdl, posterior centrodiapophyseal lamina; poz, postzygapophysis; pp, parapophysis; prdl, prezygodiapophyseal lamina;
prz, prezygapophysis; rc, radial condyle; sc, scapula; scb, scapular blade; sct, supracoracoideus tuberosity; sprl, spinoprezygapophyseal lamina;
uc, ulnar condyle; vp, ventral process.

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Nature Ecology & Evolution Articles
a

E A

Afro-Eurasia
F

b Saltasaurus
0.96
Neuquensaurus
0.11 Paralititan
0.05 Diamantinasaurus

0.12 Maxakalisaurus
Futalognkosaurus
0.31 Epachthosaurus
0.12
Mendozasaurus
0.16
0.45 Atsinganosaurus
Pellegrinisaurus
0.19
Alamosaurus
0.47 0.56
Baurutitan
0.56
Dreadnoughtus
0.46 Paludititan
0.15 Ampelosaurus
Africa–South Lohuecotitan
America 0.21 0.15
Mansourasaurus
separation
0.02 Lirainosaurus

0.32 Opisthocoelicaudia
0.36 Nemegtosaurus

125.0 120.0 115.0 110.0 105.0 100.0 95.0 90.0 85.0 80.0 75.0 70.0 65.0 Ma
‘Middle’ Cretaceous Post-Cenomanian Cretaceous

Fig. 3 | Phylogenetic, temporal and palaeobiogeographic context of Mansourasaurus shahinae gen et sp. nov. and other saltasaurid titanosaurian
sauropod dinosaurs. a, The palaeobiogeographical model proposed for a subset of post-Cenomanian saltasaurids. Note the division into Pan-American
and Afro-Eurasian clades, proposed to have arisen as a consequence of the final tectonic separation of Africa and South America ~100 million years ago
(Ma; see Supplementary Information). Palaeogeographic map © 2012 Colorado Plateau Geosystems Inc. (CPGI) of ref. 52 and reproduced with permission
from CPGI. b, Results of the tip-dated Bayesian phylogenetic and BioGeoBEARS palaeobiogeographic analyses. The decimals represent the posterior
probability for each node; estimated ancestral areas (colour-coded pie charts at nodes) denote the likeliest palaeogeographic range(s) at each node based
on the DEC+​j and DIVALIKE+​j models; the light purple bars represent the 95% highest posterior density (HPD) for the timing of the divergence date at
each node. The colour-coded bars for each terminal taxon indicate the palaeogeographic range and the 95% HPD of the recorded stratigraphic range
of that taxon; the vertical line within each bar indicates the mean tip age within the 95% HPD of the stratigraphic range. The hatched area indicates the
approximate timing of the final separation of Africa and South America. The geologic timescale is after ref. 53. A, Asia (magenta); E, Europe (teal); F, Africa
(orange); N, North America (green); S, South America (red); U, Australia (lavender).

of preserved skeletal elements of each taxon—the placement of
Mansourasaurus with Eurasian titanosaurians is consistent across all
phylogenetic models (see Supplementary Information). The Afro-
Eurasian clade as a whole (that is, Ampelosaurus, Lirainosaurus,
Lohuecotitan, Mansourasaurus, Nemegtosaurus, Opisthocoelicaudia
and Paludititan) is supported by a unique combination of syn-
apomorphies of the scapulocoracoid (scapular blade unexpanded,
scapular blade perpendicular to coracoid articulation, posterior
margin of acromial process concave, coracoid foramen at margin of
scapula–coracoid articulation). The Afro-Eurasian clade is, in turn,
sister to a Pan-American clade comprised of the South American
titanosaurs Baurutitan, Dreadnoughtus and Pellegrinisaurus and
the North American Alamosaurus. The Afro-Eurasian and Pan-
American clades are estimated to have diverged from one another

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at approximately or shortly after the time South America and
Africa separated (~100 million years ago35,36). Palaeobiogeographic
analyses suggest that Mansourasaurus arrived in Africa via a dis-
persal event from Europe during the latest Cretaceous (Fig. 3b; see
Supplementary Information).
Implications for African PCC continental vertebrate faunas. The
discovery of Mansourasaurus and the results of the analyses pre-
sented herein hold significant palaeobiogeographic implications.
Most importantly, the palaeobiogeographic affinities of terrestrial
vertebrates that inhabited continental Africa during the PCC have
historically been poorly characterized due to sparse sampling and
a highly incomplete fossil record16,37,38. Although PCC terrestrial
vertebrate fossils have previously been recovered from the African
mainland and the then-contiguous Arabian Peninsula, almost all
are fragmentary and not diagnostic to low taxonomic levels, and
therefore have allowed for only coarse palaeobiogeographic infer-
ences16,39–41. Despite this dearth of evidence, numerous authors have
proposed hypotheses regarding the nature of African terrestrial
vertebrate faunas during the PCC, ranging from assemblages domi-
nated by relictual lineages that were geographically isolated on an
‘island continent’1–3 to those with close affinities to coeval taxa from
southern Europe4,8,10,42. Until now, these scenarios had been based
mostly on extremely limited and/or ambiguous evidence. Although
the PCC non-marine vertebrate record of Europe is much more
extensive than that of the African mainland, the European taxa that
have previously been postulated as indicators of continental verte-
brate dispersal between the two landmasses are problematic in this
regard. Some (for example, the gar Atractosteus africanus, mawson-
iid coelacanths and bothremydine turtles) were probably tolerant of
salt water8,43,44, whereas others (for example, foxemydine turtles, the
crocodyliform Doratodon and abelisaurid theropods) are of ambig-
uous or even contradictory palaeobiogeographic affinities within
Gondwana (for example, the hypothesized closest relatives of the
French abelisaurid Arcovenator are from India and Madagascar45).
Even European PCC titanosaurians, traditionally considered
Gondwanan in origin10,42,46, have, until now, been of unclear phy-
logenetic and palaeobiogeographic affinities8,47. Phylogenetic and
palaeobiogeographic analyses of Mansourasaurus demonstrate its
sister taxon relationship to the near-coeval European titanosaurian
Lohuecotitan (as well as to other Eurasian forms, to the exclusion
of taxa from elsewhere in Gondwana), thereby offering the most
robust support to date for non-marine vertebrate dispersal between
Africa and Europe during the late Mesozoic. The new Egyptian
titanosaur therefore emphasizes the need for fossils that are diag-
nostic and phylogenetically informative at low taxonomic levels to
enable the adequate assessment of palaeobiogeographic hypotheses.
Methods
Phylogenetic and palaeobiogeographic analyses. We used the data set
and analytical protocols presented in a previous study9 (see Supplementary
Information) to investigate the phylogenetic and palaeobiogeographic affinities
of Mansourasaurus. The data set consists of 52 taxa and 503 morphological
characters (including autapomorphies), making it one of the most comprehensive
phylogenetic data sets yet assembled for titanosaurian sauropods. We analysed
the data using recently developed Bayesian tip-dating phylogenetic methods and
model-based palaeobiogeographic analyses9,48–50. Stratigraphic ranges for each
taxon were obtained from the technical literature (see Supplementary Information)
and sampled under a uniform prior distribution. Characters were assumed to
be unordered and rates of character evolution were variable with rates drawn
from a gamma distribution. We used the birth–death skyline serial-sampling tree
model51—which assumes that birth and death rates may vary through time—and
a lognormal relaxed clock model. The model ran for 20 million generations
with sampling occurring every 1,000 generations followed by a 25% burn-in to
discard the initial climbing phase. An alternative model was tested with similar
assumptions except characters evolving under equal rates; however, the variable
rates model was strongly favoured over the equal rates model and used for the
palaeobiogeographic analysis. BioGeoBEARS was used for the palaeobiogeographic
analysis and tested three models (DEC, DIVALIKE and BAYAREALIKE) with
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Nature Ecology & Evolution
alternative models with the additional +​j parameter to facilitate long-distance
dispersal events alongside the assumptions of each model49.
Life Science Reporting Summary. Further information on experimental design is
available in the Life Sciences Reporting Summary.
Data availability. Data have been deposited in ZooBank under Life Science
Identifier urn:lsid:zoobank.org:act:81FD8987-8020-4C57-AABF-13DE9A9BB819.
The authors declare that all other data supporting the findings of this study are
available within the paper and its Supplementary Information.
Received: 26 June 2017; Accepted: 15 December 2017;
Published: xx xx xxxx
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Acknowledgements
We thank A. Othman and A. Habib of the Faculty of Science at Mansoura University
for logistical support, and M. El-Amir and F. Ibrahim (MUVP) for their critical roles in
the field and laboratory work. A. McAfee skilfully executed the skeletal reconstruction
in Fig. 1c and contributed greatly to Fig. 2, the other components of Fig. 1, and
Supplementary Figs. 1–18. D. and R. Blakey kindly provided permission to reproduce the
palaeogeographic map in Fig. 3a. We thank M. D’Emic and V. Díez Díaz for discussions,
and V. Díez Díaz for providing unpublished photographs of the dentary of Ampelosaurus.
Funding was provided by grants from Mansoura University, the Jurassic Foundation,
the Leakey Foundation, the National Geographic Society/Waitt Foundation
(grant no. W88-10) and the National Science Foundation (EAR-1349825 to P.M.O.).
Author contributions
H.M.S. directed the project and supervised the collection of the fossils in the field;
H.M.S., I.A.E-D., S.E-S., S.S. and M.A.K. collected the fossils; I.A.E-D. and S.E-S.
supervised fossil preparation; I.A.E-D. curated and measured the fossils; fossils were
described by E.G., P.M.O., I.A.E-D. and M.C.L.; phylogenetic analysis was performed by
E.G.; H.M.S., E.G., P.M.O., I.A.E-D., J.J.W.S., E.R.S. and M.C.L. wrote the paper.
Competing interests
The authors declare no competing financial interests.
Additional information
Supplementary information is available for this paper at https://doi.org/10.1038/
s41559-017-0455-5.
Reprints and permissions information is available at www.nature.com/reprints.
Correspondence and requests for materials should be addressed to H.M.S.
Publisher’s note: Springer Nature remains neutral with regard to jurisdictional claims in
published maps and institutional affiliations.
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