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A reconstruction of Tazoudasaurus naimi (Dinosauria,

Sauropoda) from the late Early Jurassic of Morocco
a a
Karin Peyer & Ronan Allain
a
Département Histoire de la Terre, Muséum National d'Histoire Naturelle, CR2P, UMR 7207
du CNRS, CP 38, 57 rue Cuvier, F-75231, Paris Cedex 05, France

Available online: 27 Apr 2010

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the late Early Jurassic of Morocco, Historical Biology: An International Journal of Paleobiology, 22:1-3, 134-141

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 Historical Biology
Vol. 22, Nos. 1 – 3, March – June – September 2010, 134–141

A reconstruction of Tazoudasaurus naimi (Dinosauria, Sauropoda) from the late Early

Jurassic of Morocco
Karin Peyer* and Ronan Allain
Département Histoire de la Terre, Muséum National d’Histoire Naturelle, CR2P, UMR 7207 du CNRS, CP 38, 57 rue Cuvier, F-75231
Paris Cedex 05, France
(Received 20 July 2005; final version received 17 August 2006)

The basal sauropod Tazoudasaurus naimi from the late Early Jurassic of Morocco is represented by at least ten juvenile to
adult individuals. Over the past seven years, the Toundoute continental series of Ouarzazate Province, a lateral equivalent of
the Azilal and Wazzant Formations of the central High Atlas mountains (Toarcian to Early Aalenian age), produced over 600
skeletal elements pertaining to Tazoudasaurus. About a fifth of the available material has been prepared and studied. Except
for the skull, the osteology of Tazoudasaurus is completely known. Superb preservation and the abundance of osteological
data from the presently known Tazoudasaurus individuals make it possible to provide a detailed skeletal reconstruction for
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the first time.

Keywords: Dinosauria; Sauropoda; Tazoudasaurus; Early Jurassic; Morocco

Introduction Specimen examined

Compared to known Middle and Late Jurassic sauropod
remains, the Early Jurassic sauropod record is sparse. Only
a handful specimens such as Gongxianosaurus (He et al.
1998), Barapasaurus (Jain et al. 1975), Vulcanodon (Raath
1972), and now Tazoudasaurus (Allain et al. 2004) are
known from Lower Jurassic rocks. Especially in Africa,
recent fieldwork has been conducted foremost in late
Mesozoic sediments what might in part explain the poor
fossil record of Early Jurassic sauropods. In addition, strata
exposures of that age are scarce (Allain and Aquesbi
2008). Hence, the discovery of the virtually complete
Tazoudasaurus from the High Atlas Mountains of Morocco
can explain basal sauropod morphology in much greater
detail on the genus level, but also on the family level more
precisely, the Vulcanodontidae (Cooper 1984; Allain et al.
2004). Vulcanodontidae, a basal group of Gravisauria
(Allain and Aquesbi 2008), have so far only been
represented by the fragmentary specimen of Vulcanodon,
the skull elements of which are unfortunately unknown.
The High Atlas is known for its Middle Jurassic
alternating continental and marine fossil bearing facies
and the underlying continental units of the Azilal and its
lateral equivalents, the Wazzant and the Toundoute
Formation (Jenny 1988; Montenat et al. 2005). This facies
are considered Torcian to Early Aalenian in age (Jenny
1985, 1988; Allain et al. 2004) and have yielded sauropod
and theropod remains.
The holotype specimen of the basal sauropod
Tazoudasaurus (Allain et al. 2004) and referred skeletal
material belonging to several juvenile and adult
specimens (Allain and Aquesbi 2008) have been
collected at different fossil sites, all in proximity of
each other (Figures 1 and 2). The fossil sites of the
upper bone beds (Allain and Aquesbi 2008) have been
termed, from the South to the North: O, To1, To10 , Pt
haut, Pt and R (Figure 1). The To1 site yielded one
adult (holotype) and two juvenile specimens. At To10 ,
one adult, a subadult and a juvenile specimen were
recovered. The site, which consists of Pt haut and Pt, is
a downward-sloping fossil bearing site that produced an
adult specimen at Pt haut and another adult plus
remains of the theropod Berberosaurus liassicus at Pt
(Figure 2). Current fieldwork at the new sites O and R
promise further remains of Tazoudasaurus skeletons and
theropod remains, all of which are still being excavated
and prepared. And finally, the lower bone bed has only
yielded dinosaur remains at the To2 site, including one
juvenile and a subadult skeleton of Tazoudasaurus and
an enigmatic theropod. Today, a total of 10 sauropod
Tazoudasaurus specimens and at least two theropod
specimens are known (Figure 1). Known skeletal
elements of each of the known specimens of
Tazoudasaurus are shown in Figure 3 and listed in
Table 1.

*Corresponding author. Email: karin_peyer@yahoo.fr

ISSN 0891-2963 print/ISSN 1029-2381 online
q 2010 Taylor & Francis
DOI: 10.1080/08912960903562317
http://www.informaworld.com
 Historical Biology 135

Figure 1. Fossil site map indicating the sites To1, To10 , To2, Pt (Pt haut, Pt, Pt bas) and the two new fossil locality sites R and O.

Reconstruction
The small sauropod Tazoudasaurus, pertaining to the
family vulcanodontidae, measured 9.5– 10 m as an adult.
A fully grown individual probably weighed around 8
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metric tonnes (calculated after Anderson et al. 1985). The

Figure 2. Fossil site map of the Pt region indicating Pt haut, Pt
and Pt bas.
juvenile specimen 5 from To1 with a femoral length of
290 mm would have weighed around 140 kg.
The skull elements have been described in detail by
Allain et al. (2004). A fairly complete lower jaw and a few
disarticulated elements of the skull and braincase have
been identified from a single animal (To1-275). Additional
information for the reconstruction of the skull of
Tazoudasaurus has been obtained from the articulated
skulls of Shunosaurus (Zhang 1988) and Abrosaurus
(Ouyang 1989). The latter two and Tazoudasaurus have
strikingly similar lower jaws in possessing a mandibular
fenestra that is bordered by the dentary, surangular and
angular, respectively. Also, the overall dimension and

Figure 3. Skeletal elements in grey indicate presently known/prepared bones for each specimen. A, specimen 1, adult (holotype); B,
specimen 6, juvenile; C, specimen 5, juvenile; D, specimen 2, adult; E, specimen 3, subadult; F, specimen 7, juvenile; G, specimen 8,
adult; H, specimen 4, adult; I, specimen 9, juvenile and J, specimen 10, subadult.
 136 K. Peyer and R. Allain

Table 1. Specimen list indicating all presently known skeletal elements for each specimen of T. naimi, locality, side of body and
measurements in mm. Measurements indicate always maximum length for long bones and centrum length for vertebrae.

Locality Specimen number Side Measurement

Specimen 1, holotype adult To1
Mandible To1-275 Left
Tooth To1-20
Postorbital Left
Quadrate Right
Astragalus Right
Proximal chevron To1-187
Middle chevron To1-217
Distal chevron To1-182
Pubis To1-103 Right 610
Ungual phalanx I To1-94
Ungual phalanx III To1-114
Ungual phalanx of IV To1-14
MtII To1-265 Left?
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MtI To1-22 Left

MtIV To1-13 Right , 250
Pedal phalanx To1-121
Pedal phalanx To1-137
Pedal phalanx To1-158
Pedal phalanx To1-67
Femur To1-04 Left , 1150
Fibula To1-42 Right 730
Fibula To1-43 Left 730
Axis To1-239 , 121
Ant. cervical To1-354 250
Mid cervical To1-64 260
Dorsal vertebrae To1-69 85
Dorsal vertebrae To1-38 A – C 122, 120, 120
Post. dorsal vertebrae To1-156 101
Ant. caudal vertebrae To1-100 78
Ant. mid caudal vertebrae To1-303 A – C 115, 107, 110
Mid caudal vertebra To1-288 88
Post. mid caudal vertebra To1-88 141
Distal caudal vertebra To1-317
Distal caudal vertebra To1-357
Specimen 2, adult To10
Ulna To1-375 Right 720
Ischium To1-378 Right ?
Ilium To1-373 Left 795
Scapula To1-518 Left 1160
Specimen 3, subadult To10
Fibula To1-377 Left 530
Specimen 4, adult Pt
Humerus Pt1 Left 1030
Ulna Pt24 Left 730
Radius Pt24 Left ?
MtV Pt25 Right
Specimen 5, juvenile To1
Femur To1-256 Right 290
Humerus To1-93 Left 185
Humerus To1-48 Right 185
Astragalus To1-135 Right
Specimen 6, juvenile To1
Tibia To1-76 Left 250
Femur To1-105 Right 490
Specimen 7, juvenile To10
Ischium To1-379 Right 285
Ulna To1-374 Right 220
 Historical Biology 137

Table 1 – continued

Locality Specimen number Side Measurement

Specimen 8, adult Pt haut

Tibia To1-380, To1-382 Right, left 710
Femur To1-381 Right 1230
5 dorsal vertebrae To1-514
Specimen 9, juvenile To2
Manus To2-112 Left
Specimen 10, subadult To2
Coracoid To2-109 Right

Notes: , , estimated length; ?, bone incomplete.

form of the lower jaw bones resemble each other in all margin of the orbit. Posteromedially, it overlaps the
specimens. In Shunosaurus and Tazoudasaurus, the ventral frontal, anteromedially the nasal, and its descending
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margin of the dentary is slightly concave but straight in
Abrosaurus. The overall length of the skull was
reconstructed measuring 32 cm (Figure 4).
Description
Skull
The frontal and parietal are fused. The frontals are longer
than wide and contribute to the supratemporal fossa. The
parietals are considerably wider posteriorly than anteriorly
and the distance separating the supratemporal fenestrae is
very broad. This indicates that at least the back of the skull
of Tazoudasaurus was fairly wide, as in Atlasaurus
(Monbaron et al. 1999).
A recently discovered right prefrontal is rectangular
and smooth in dorsal view. It forms the anterodorsal
process contacts the lacrimal and or the maxilla ventrally
(Figure 4). Medially, the prefrontal is very thin and
plate-like.
The ventral process of the postorbital is longer than the
dorsal one. The reconstruction of the skull suggests that
the ventral process must have been quite long to reach the
very short postorbital process of the jugal (Figure 4). The
postorbital contacts and probably overlapped the anterior
process of the squamosal laterally.
The left jugal is well preserved, missing only a small
fragment of its anterior ramus, the part forming the
posterior edge of the antorbital fenestra (Figure 5). The
jugal forms the anteroventral border of the lower temporal
fenestra, the ventral border of the orbit, and reaches
forward to the antorbital fenestra. In Tazoudasaurus, the

Figure 4. Tazoudasaurus naimi (T. naimi). Reconstruction of the skull with known skull elements of Tazoudasaurus. Other available not
visible elements include: splenial, prearticular and surangular. Unknown skull elements reconstructed after Abrosaurus (Ouang 1989) and
Shunosaurus (Zhang 1988). Abbreviations: a, angular; ant f., antorbital fenestra; d, dentary; f, frontal; j, jugal; l, lacrimal; lt. f, lower
temporal fenestra; mx, maxilla; n, nasal; na, naris; o, orbit; pa, parietal; pmx, premaxilla; po, postorbital; prf, prefrontal; q, quadrate; qj,
quadratojugal; s, squamosal and sur, surangular.
 138 K. Peyer and R. Allain
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Figure 5. T. naimi. Left jugal in A, medial and B, lateral view. Right quadratojugal in C, lateral and B, medial view. Abbreviations: ant. f,
antorbital fenestra, posterior margin; c. l, contact for lacrimal; c. mx, contact surface for maxilla; c. po, contact for postorbital and c. qj,
contact surface for quadratojugal. Scale bar 5 cm.
jugals’ participation of the orbit is very limited. The
posteroventral process is somewhat flared and was
covered by the quadratojugal laterally. Just anterior and
ventral to the quadratojugal contact, a short finished
surface of roughly 1 cm length indicates the lower border
of the skull. The maxilla covered the jugal anteroventrally
and did not reach the quadratojugal. Anterior to the
ascending process of the jugal is a well-marked articular
surface for the contact with the lacrimal. The ascending
process itself is short; posterodorsally oriented and
contacted the postorbital. A small foramen is visible on
the lateral side just below the contact with the lacrimal. A
foramen of similar size opens also on the medial side in
roughly the same spot.
The newly added quadratojugal is beautifully pre-
served, uncompressed and lacks only the anterior tip of its
rostral ramus (Figure 5). It is inversely L-shaped and in
lateral view, the anterior process projects posteriorly
beyond the dorsal process as in Nemegtosaurus
(Wilson 2005). This posteriorly projecting process was
wrapped around the quadrate posteriorly and also poster-
oventrally. Its dorsal process is high and plate-like,
compared to the thin and slender dorsal process in
Shunosaurus (Chatterjee and Zheng 2002). The rostral
ramus is tongue-shaped with a slightly excavated dorsal
margin to mark the ventral border of the lower temporal
fenestra, and a straight ventral margin. It is not clear if the
anterior process contacted the maxilla.
The quadrate lacks a quadrate fossa, a feature observed
in prosauropods. In posterior view, its shaft is bent slightly
medially.
Lower jaw
The lower jaw is slender and is the deepest at or slightly
behind the mandibular fenestra. In addition to the currently
known left surangular (see Allain and Aquesbi 2008), a
somewhat smaller left surangular is now fully prepared.
This newly added element is better preserved and displays
good contacts for the angular ventrally and the dentary
anteriorly. It forms the outer wall of the adductor fossa.
The right and left mandibles of the lower jaw formed
more of a V-shaped structure in dorsal view rather than an
arch as seen in later sauropods (Figure 6). Eighteen teeth
with conical denticles on the mesial and distal margins of
the crowns are present.
Axial skeleton
The axis and seven postaxial cervicals are known. Based
on comparisons with prosauropods and other sauropods
for which the number of cervicals are known
(prosauropod Plateosaurus Huene 1926, basal Eusauro-
pod Shunosaurus Zhang 1988), the neck has been
reconstructed with 12 strongly opisthocoelous elements.
They are slightly elongated and lack true pleurocoels.
The spool-shaped cervical centra suggest that the head
was held at or below the height of the shoulder in

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Figure 6. T. naimi. Reconstruction of the anteriorly V-shaped
lower jaw.
natural position as is the case for other sauropods
(Stevens and Parrish 1996, 1999, 2005).
The dorsal vertebral column is known from an
anterior, three and respective five consecutive mid-dorsal
and a posterior element. While the anterior dorsal
vertebrae possess a high centrum and neural spine, the
mid dorsals are much lower in total height. The posterior
dorsal vertebrae seem to be a bit higher than the mid-
dorsals but never reach the height of the anterior dorsals.
The dorsal vertebral column has been reconstructed with
13 elements. It curves high over the pectoral girdle and
then shows a saddle-like depression in the mid-dorsal
region (Figure 7). The anterior dorsal vertebrae are
anteroposteriorly shorter than in the mid region and their
neural spines are almost vertically oriented rather than
more posteriorly. All dorsals are amphicoelous and they
too are devoid of true pleurocoels.
Amphicoelous, anteroposteriorly short centra (length/
height ratio , 0.5), are characteristic of the anterior caudal
Figure 7. Skeletal reconstruction of T. naimi. The length of adult specimen of Tazoudasurus was estimated to measure , 9.5 m.
Historical Biology 139
vertebrae. The centra are getting longer in the mid- to
posterior-mid dorsals (length/height ratio 0.6 –1) until
they, reach a length twice their height in the posterior
region of the tail (Allain and Aquesbi 2008). Tall caudal
neural spines transgress from a vertical orientation in the
anterior vertebrae to very short, almost completely
posteriorly directed neural spines at the extremity of the
tail. The caudal vertebral region is known from one
anterior and numerous mid and distal caudal vertebrae.
The lack of a completely articulated tail makes it difficult
to assess with certainty, the exact number of elements in
the tail and its complete length. Preliminary calculations
suggest a tail length of 4.4– 4.9 m for an adult animal.
Pectoral girdle and limbs
Recent fieldwork has produced a right scapula (Figure 8)
from the To10 quarry of an adult specimen. Except for
its antero-dorsal border, it is complete. A corresponding
coracoid has not been collected and the scapula shows
no sign of a former fusion to the coracoid. Skeletal
element ratios suggest that the scapula belonged to an
adult individual – suggesting that scapula and coracoid
did not fuse even in adult life in Tazoudasaurus. The
scapular blade measures 116 cm in length. The scapula
in an adult specimen would have been slightly longer
than the humerus (humerus/scapula: 0.89) but shorter
than the femur (femur/scapula: 1.06). The scapula of the
related Gravisauria Vulcanodon has been calculated to be
shorter than the humerus (humerus/scapula: 1.02) and
the femur (femur/scapula: 1.40), but these bones are
broken in Vulcanodon (Cooper 1984). The distal end of
the scapula in Tazoudasaurus is only slightly expanded
and its width does not exceed the maximum width of the
shaft (slightly above mid-height). The low acromion
ridge divides the large anterior fossa from the smaller
and only partially preserved posterior fossa. The scapula
is very massive near the glenoid region and shows an
exceptionally deep articular contact with the coracoid
(Figure 8). This massiveness diminishes towards the
acromion ridge. The scapular shaft is thin and assumes
 140 K. Peyer and R. Allain
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Figure 8. T. naimi. Right scapula (To10 ) in A, lateral, C,
proximal, and D, dorsal view. Right coracoid (To2) in B, lateral
view. Coracoid scaled to fit scapula. Scale bars 10 cm.
the curvature of the torso. In order to assure an optimal
contact of the vertically oriented humerus and the
glenoid fossa, the scapulocoracoid was probably oriented
almost horizontal to the ribs. The scapula distal margin
was set sufficiently below the dorsal margins of the
neural spines to leave ample room for the cartilageonous
suprascapula of sauropods (Wilhite 2005). The presence
of such a suprascapula is indicated by the rugose distal
end of the scapula of Tazoudasaurus.
A right coracoid belonging to a juvenile or subadult
specimen of Tazoudasaurus has been described earlier
(Allain and Aquesbi 2008). It measures 34 cm dorsoventrally
compared to 47 cm (estimated length) in an adult specimen.
The front limbs of juvenile to adult Tazoudasaurus
specimens and a nearly complete articulated left manus of a
juvenile individual have been described in detail (Allain
and Aquesbi 2008). Relevant for the present reconstruction
Figure 9. T. naimi. Reassembled right manus, carpal element,
and distal radius and ulna of a juvenile specimen in anterolateral
view. Scale bar 5 cm.
is the arrangement of the metacarpals and phalanges
(Figure 9). The metacarpals have a spreading configuration
and are held off the ground at a 458. The external part of the
manus was oriented anterolaterally. The massive, highly
recurved pollex ungual is slightly turned medially. The
proximal articulation surfaces together form only a half
arch compared to the highly arched configuration (2708) in
more derived Eusauropods. The distal part of the first
phalanges and the more distal phalanges would have
contacted the substrate while the proximal part of the first
phalanges did not (Allain and Aquesbi 2008). Such an
arrangement is best described as sub-unguligrade (Carrano
1997) and marks a transition between the prosauropod
condition and the fully unguligrade semi-tubular manus of
neosauropods. A block-shaped, rectangular carpal (distal
carpals 1 þ central), has been preserved contacting
metacarpal 1 and 2 distally and the radius proximally
(Allain and Aquesbi 2008).
Pelvic gridle and limbs
The preacetabular process of the ilium is rounded
anteriorly and the iliac blade is dorsally slightly concave.
The prominent pubic peduncle is oriented ventrally, the
ischial peduncle is reduced. In general, the ilium of
Tazoudasaurus resembles that of a sauropod rather than a
prosauropod (Raath 1972). The relatively slender, almost
ventrally directed, pubis is anteroposteriorly expanded at
its distal end and measures only 0.6 of the length of
the ischium. A similar ratio is given for Vulcanodon and
the camarasaurid Haplocanthosaurus (Raath 1972). The
ischium is directed posteroventrally and is slightly curved

at its distal end. The slightly shorter forelimbs of
Tazoudasaurus measure 90% of the hindlimbs.
The humerus of Vulcanodon is not known but ratio
calculations of present limb bones suggest similar proportion
in limb length in Tazoudasaurus and Vulcanodon (ulna/tibia
is 1.01 and 1.06 respectively; and tibia/femur is 0.58 in both
specimens which is typical for sauropods).
The femoral head is bulbous and dorsomedially
directed. A lesser trochanter is present on the lateral side
and the fourth trochanter is reduced. The tibia exhibits a
strongly anteroposteriorly expanded proximal end and a
transversally flattened shaft.
Four metatarsals and six pedal phalanges are known
for the holotype specimen. A reconstruction of the pes is
not possible with the available material. Future fieldwork
will hopefully add more information to the arrangement
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and construction of the pedal digits.
Summary
Lower Jurassic sauropods from Laurasia are rare and not
well documented. Until recently, vulcanodontids have
only been represented by incomplete skeletal material, e.g.
Vulcanodon. The previously published description of
Tazoudasaurus (Allain et al. 2004; Allain and Aquesbi
2008) and the present description of newly discovered and
or prepared material, provide detailed information of a
virtually complete skeleton. This is the first time that a
skeleton of a vulcanodontid sauropod is preserved in
sufficient detail that a complete skeletal reconstruction can
be made. Cranial material for sauropods in general and
basal sauropods in particular is rare and often distorted.
Tazoudasaurus with its beautifully preserved cranial
material is therefore a significant fossil and aids in
evaluating sauropod evolution. In the near future, a new
paleontological museum will be built in Toundoute,
Morocco and will house and display the small sauropod
Tazoudasaurus as one of its most remarkable fossils. The
present restoration will certainly aid in a successful and
accurate specimen display mount.
Acknowledgements
This study has been supported by the National Geographic
Society (Grants 7789-05 and 8186-07), the Ministry of Energy
and Mines of Morocco, the Agence Universitaire de la
Francophonie and the Fondation des Treilles. The Natural
History Museum of Marrakech has been especially supportive
and helpful. We thank Emilie Lang for her comments and
improvement of this article, Florent Goussard for the weight
estimation of Tazoudasaurus, P. Loubry for photographs and
P. Richir, T. Vacant, M. M’Ghari, M. Rochdy and A. Faskaoune
for the preparation of the material. Special thanks also to the
Historical Biology 141
members of the DinoAtlas Project and to Nour-Eddine Jalil and
the steering committee of NAVEP1.
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