Macronaria Sereno, 1997
Definition- (Saltasaurus loricatus <- Diplodocus longus)
(Wilson, 2005; modified from Wilson and Sereno, 1998)
= Morosauria Haeckel, 1895
= "Brachiosauria" Upchurch, 1997
References- Haeckel, 1895. Systematische Phylogenie der Wirbelthiere:
(Vertebrata). 660 pp.
Sereno, 1997. The origin and evolution of dinosaurs. Annual Review of Earth
and Planetary Sciences. 25, 435-489.
Upchurch, 1997. A cladistic analysis of sauropod dinosaur phylogeny. Journal
of Vertebrate Paleontology. 17(3), 82A.
Wilson and Sereno, 1998. Early evolution and higher-level phylogeny of sauropod
dinosaurs. Society of Vertebrate Paleontology Memoir 5. Journal of Vertebrate
Paleontology. 18(2 suppl), 68 pp.
Wilson, 2005. Overview of sauropod phylogeny and evolution. in Curry Rogers
and Wilson (eds.). The Sauropods: Evolution and Paleobiology. University of
California Press, Berkeley. 15-49.
Haestasaurus Upchurch, Mannion
and Taylor, 2015
H. becklesii (Mantell vide Anonymous, 1852) Upchurch, Mannion
and Taylor, 2015
= Pelorosaurus becklesii Mantell vide Anonymous, 1852
= Morosaurus becklesii (Mantell vide Anonymous, 1852) Marsh, 1889
= Camarasaurus becklesii (Mantell vide Anonymous, 1852) McIntosh, 1990
vide Upchurch, Mannion and Taylor, 2015
Late Berriasian-Valanginian, Early Cretaceous
Hastings Beds Group, England
Holotype- (NHMUK R1870) humerus (599 mm), radius (404 mm), ulna (421 mm),
skin impression
Diagnosis- (after Upchurch et al., 2015) anteromedial corner of distal
humerus projects to form anterior entepicondylar process; two small vertical
ridges situated between lateral and medial anterodistal processes of humerus;
proximal articular surface of radius widest anteroposteriorly along lateral
margin; lateral edge of proximal radius nearly straight; anterodistal surface
of radius shallowly concave between anterolateral and anteromedial ridges; combination
of robust ulna (proximal width / proximodistal length > 0.4) and slender
radius (transverse proximal width / proximodistal length ratio < 0.3).
Comments- The holotype was discovered in 1852. Note that while the initial
description is usually cited as 'Mantell, 1852', the paper is an anonymous report
of Mantell's talk. While initially referred to Pelorosaurus based on
vague humeral resemblences, Lydekker (1888) suggested it may belong to "Cetiosaurus"
brevis (a synonym of Pelorosaurus), Titanosaurus or a new
genus. McIntosh (1990) incorrectly stated Huene (1932) referred becklesii
to Camarasaurus (while he did place becklesii in his Camarasaurinae,
he only referred to the taxon as "Gen. (?) becklesii"), but
Upchurch et al. (2015) are the first authors to explicitly use the combination
Camarasaurus becklesii. Although authors starting with McIntosh have
generally believed becklesii to be a distinct genus from Pelorosaurus,
this was only made official with the publication of Upchurch et al. (2015).
D'Emic (2012) placed the taxon in Titanosauriformes based on the "anteromedial
arm of the ulna being much longer than its anterolateral arm", but outside
Chubutisaurus+Titanosauria based on the lack of an undivided notch on the humeral
radial condyle. Mannion et al. (2013) included it as an OTU in their analysis
and recovered a position in Laurasiformes as a non-titanosaurian somphospondylan,
as a basal titanosauroid close to Phuwiangosaurus, or as a basal andesaurid
andesauroid in their analyses depending on assumptions made. Upchurch et al.
recovered Haestasaurus as a camarasaurid in a version of Harris' sauropod
analysis, and a non-camarasauromorph macronarian or basal somphospondylan in
Mannion et al.'s macronarian analysis.
References- Anonymous, 1852. On the structure of the Iguanodon
and on the fauna and flora of the Wealden Formation. Notices of the Proceedings
at the Meetings of the Members of the Royal Institution. 1, 141-146.
Lydekker, 1888. Note on a new Wealden iguanodont and other dinosaurs. Quarterly
Journal of the Geological Society of London. 44, 46-61.
Marsh, 1889. Comparison of the principal forms of the Dinosauria of Europe and
America. American Journal of Sciences, series 3. 37, 323-330.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte.
Monographien zur Geologie und Palaeontologie. 4(1), viii + 361 pp.
McIntosh, 1990. Sauropoda. In Weishampel, Dodson and Osmolska (eds.). The Dinosauria.
University of California Press. 345-401.
D'Emic, 2012. The early evolution of titanosauriform sauropod dinosaurs. Zoological
Journal of the Linnean Society. 166, 624-671.
Mannion, Upchurch, Barnes and Mateus, 2013. Osteology of the Late Jurassic Portuguese
sauropod dinosaur Lusotitan atalaiensis (Macronaria) and the evolutionary
history of basal titanosauriforms. Zoological Journal of the Linnean Society.
168(1), 98-206.
Upchurch, Mannion and Taylor, 2015. The anatomy and phylogenetic relationships
of "Pelorosaurus" becklesii (Neosauropoda, Macronaria) from
the Early Cretaceous of England. PLoS ONE. 10(6), e0125819.
Camarasauromorpha Salgado, Coria and
Calvo, 1997
Definition- (Camarasaurus supremus + Titanosaurus indicus)
(modified from Salgado et al., 1997)
Other definitions- (Camarasaurus supremus + Saltasaurus loricatus)
(modified from Upchurch et al., 2004)
References- Salgado, Coria and Calvo, 1997. Evolution of titanosaurid
sauropods. I: Phylogenetic analysis based on the postcranial evidence. Ameghiniana.
34(1), 3-32.
Upchurch, Barrett and Dodson, 2004. Sauropoda. in Weishampel, Dodson and Osmolska
(eds.). The Dinosauria (2nd edition). University of California Press, Berkeley.
259-322.
Camarasauridae Cope, 1877
= Morosauridae Marsh, 1882
= Camarasaurinae Cope, 1877 vide Nopcsa, 1928
References- Cope, 1877. On Amphicoelias, a genus of saurians from
the Dakota epoch of Colorado. Proceedings of the American Philosophical Society.
17, 242-246.
Marsh, 1882. Classification of the Dinosauria. American Journal of Science.
23, 81-86.
Nopcsa, 1928. The genera of reptiles. Palaeobiologica. 1, 163-188.
Camarasaurus
Kimmeridgian, Late Jurassic
Salt Wash Member of the Morrison Formation, Wyoming, US (Lake’s Quarry
1A; = Big Canyon Quarry)
Referred- ?(YPM 58526; formerly part of YPM 1879, the Camptonotus amplus
holotype) (juvenile) pedal ungual I (~70 mm)
Comments- The ungual YPM 58526 was originally believed to be part of
the Camptonotus amplus holotype pes, but was noted to be similar to a
sauropod's pedal ungual I by Gilmore (1909). It was assigned to Neosauropoda
and probably Camarasaurus by Galton et al. (2015), and given a new catalogue
number. While Camarasaurus specimen YPM 4633 is also known from the same
quarry, it is much too large to be the same individual. Although Lakes collected
further small elements from the quarry that might belong to this juvenile sauropod,
they are no longer preserved and are believed to have been discarded by Marsh.
References- Marsh, 1879. Principal characters of American Jurassic dinosaurs.
Part I. American Journal of Science, series 3. 16, 411-416.
Gilmore, 1909. Osteology of the Jurassic reptile Camptosaurus, with a
revision of the species of the genus, and descriptions of two new species. Proceedings
of the United States National Museum. 36, 197-332.
Galton, Carpenter and Dalman, 2015. The holotype pes of the Morrison dinosaur
Camptonotus amplus Marsh, 1879 (Upper Jurassic, western USA) - Is it
Camptosaurus, Sauropoda or Allosaurus? Neues Jahrbuch f�r
Geologie und Pal�ontologie, Abhandlungen. 275(3), 317-335.
unnamed clade
Definition- (Titanosaurus indicus <- Camarasaurus supremus)
Comments- Many taxa are referred to basal Titanosauriformes, but since
that clade is defined using Giraffatitan or Brachiosaurus, it's
likely a portion technically fall outside it even if they are closer to Titanosauriformes
than Camarasaurus. An example is Astrodon as analyzed in Wilson's
matrix by Rose (2007).
References- Rose, 2007. A new titanosauriform sauropod (Dinosauria: Saurischia)
from the Early Cretaceous of Central Texas and its phylogenetic relationships.
Palaeontologia Electronica. 10(2), 8A.
unnamed camarasauromorph (Lydekker, 1893)
Mid Cenomanian-Turonian, Late Cretaceous
Bajo Barreal Formation, Chubut, Argentina
Material- (MLP 21; paratype of Argyrosaurus superbus) incomplete
femur (~1.63 m) (Lydekker, 1893)
Comments- MLP 21 was referred to Argyrosaurus superbus by Lydekker
(1893) based on size, but cannot be compared to the holotypic forelimb of that
taxon and was assigned merely to Titanosauriformes indet. by Mannion and Otero
(2012) in their revision of the genus. This was based on the proximolateral
femoral bulge, but this is also present in more basal taxa (e.g. Janenschia,
Tehuelchesaurus, Aragosaurus, Dongbeititan).
References- Lydekker, 1893. The dinosaurs of Patagonia. Anales del Museo
de la Plata, Seccion de Paleontologia. 2, 1-14.
Huene, 1929. Los Saurisquios y Ornitisquios del Cretaceo Argentino. Anales del
Museo de La Plata. 3, 1-196.
Bonaparte and Gasparini, 1979. Los sauropodos de los grupos Neuquen y Chubut
y sus relaciones cronologicas. Actas V Congreso Geologico Argentino, Neuquen.
2, 393-406.
Mannion and Otero, 2012. A reappraisal of the Late Cretaceous Argentinean sauropod
dinosaur Argyrosaurus superbus, with a description of a new titanosaur
genus. Journal of Vertebrate Paleontology. 32(3), 614-638.
unnamed camarasauromorph (Huene, 1929)
Early Santonian, Late Cretaceous
Bajo de la Carpa Formation of the Rio Colorado Subgroup, Neuquen, Argentina
Material- (MLP 27) incomplete femur (~2.12 m)
Comments- Huene (1929) referred this to Argyrosaurus, but it cannot
be compared to the holotypic forelimb of that taxon and was assigned merely
to Titanosauriformes indet. by Mannion and Otero (2012) in their revision of
the genus. This was based on the proximolateral femoral bulge, but this is also
present in more basal taxa (e.g. Janenschia, Tehuelchesaurus,
Aragosaurus, Dongbeititan).
References- Huene, 1929. Los Saurisquios y Ornitisquios del Cretaceo
Argentino. Anales del Museo de La Plata. 3, 1-196.
Bonaparte and Gasparini, 1979. Los sauropodos de los grupos Neuquen y Chubut
y sus relaciones cronologicas. Actas V Congreso Geologico Argentino, Neuquen.
2, 393-406.
Powell, 2003. Revision of South American titanosaurid dinosaurs: Palaeobiological,
palaeobiogeographical and phylogenetic aspects. Records of the Queen Victoria
Museum. 111, 173 pp.
Mannion and Otero, 2012. A reappraisal of the Late Cretaceous Argentinean sauropod
dinosaur Argyrosaurus superbus, with a description of a new titanosaur
genus. Journal of Vertebrate Paleontology. 32(3), 614-638.
Tehuelchesaurus
Aragosaurus
Galveosaurus
Titanosauriformes Salgado, Coria and
Calvo, 1997
Official Definition-
(Giraffatitan brancai + Saltasaurus loricatus)
(Silva Junior, Martinelli, Lori, Marinho, Hechenleitner and Langer,
2022; originally Wilson, 2005; Registration Number 438)
Other definitions- (Giraffatitan brancai + Chubutisaurus insignis
+ Titanosaurus indicus) (modified from Salgado et al., 1997)
(Brachiosaurus altithorax + Saltasaurus
loricatus) (Taylor, 2009; modified from Wilson and Sereno, 1998)
= Titanomorpha Sereno, 1997
References- Salgado, Coria and Calvo, 1997. Evolution of titanosaurid
sauropods. I: Phylogenetic analysis based on the postcranial evidence. Ameghiniana.
34(1), 3-32.
Sereno, 1997. The origin and evolution of dinosaurs. Annual Review of Earth
and Planetary Sciences. 25, 435-489.
Wilson and Sereno, 1998. Early evolution and higher-level phylogeny of sauropod
dinosaurs. Society of Vertebrate Paleontology Memoir 5. Journal of Vertebrate
Paleontology. 18(2 suppl), 68 pp.
Wilson, 2005. Overview of sauropod phylogeny and evolution. in Curry Rogers
and Wilson (eds.). The Sauropods: Evolution and Paleobiology. University of
California Press, Berkeley. 15-49.
Taylor, 2009. A re-evaluation of Brachiosaurus altithorax Riggs 1903
(Dinosauria, Sauropoda) and its generic separation from Giraffatitan brancai
(Janensch 1914). Journal of Vertebrate Paleontology. 29(3), 787-806.
Silva Junior, Martinelli, Lori, Marinho, Hechenleitner and Langer, 2022 (online 2021). Reassessment of Aeolosaurus maximus,
a titanosaur dinosaur from the Late Cretaceous of southeastern Brazil.
Historical Biology. 34(3), 403-411.
Astrodontidae Huene, 1932 vide Huene, 1948
Astrodontinae Huene, 1932
References- Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre
Entwicklung und Geschichte [The fossil reptile order Saurischia, their development
and history]. Monographien zur Geologie und Palaeontologie, serie 1. 4(1-2),
361 pp.
Huene, 1948. Short review of the lower tetrapods. in Du Toit (ed.). Robert Broom
Commemorative Volume. Royal Society of South Africa Special Publication. 65-106.
Astrodon
Duriatitan Barrett, Benson
and Upchurch, 2010
D. humerocristatus (Hulke, 1874) Barrett, Benson and Upchurch,
2010
= Ceteosaurus humerocristatus Hulke, 1874
= Ornithopsis humerocristatus (Hulke, 1874) Lydekker, 1888
= Pelorosaurus humerocristatus (Hulke, 1874) Lydekker, 1890
Early Kimmeridgian, Late Jurassic
Lower Kimmeridge Clay, England
Holotype- (NHMUK R44635) humerus (~1.4 m)
Diagnosis- (after Barrett et al., 2010) acute ridge at posterolateral
corner of proximal humerus, posterior to deltopectoral crest.
Comments- Given to the NHMUK in 1873. Barrett et al. (2010) considered
Duriatitan a basal titanosauriform, and Mannion et al. (2013) placed
it in Titanosauriformes based on the medially extending deltopectoral crest.
References- Hulke, 1874. Note on a very large saurian limb-bone adapted
for progression on land, from the Kimmeridge Clay of Weymouth, Dorset. Quarterly
Journal of the Geological Society. 30, 16-17.
Lydekker, 1888. Catalogue of the Fossil Reptilia and Amphibia in the British
Museum (Natural History). Part I. Containing the Orders Ornithosauria, Crocodilia,
Dinosauria, Squamata, Rhynchocephalia, and Proterosauria. British Museum (Natural
History), London. 309 pp.
Lydekker, 1890. Catalogue of the Fossil Reptilia and Amphibia in the British
Museum. Part IV. Containing the orders Anomodontia, Ecaudata, Caudata and Labyrinthodontia;
and Supplement. British Museum (Natural History), London. 295 pp.
Barrett, Benson and Upchurch, 2010. Dinosaurs of Dorset: Part II, the sauropod
dinosaurs (Saurischia, Sauropoda) with additional comments on the theropods.
Proceedings of the Dorset Natural History and Archaeological Society. 131, 113-126.
Mannion, Upchurch, Barnes and Mateus, 2013. Osteology of the Late Jurassic Portuguese
sauropod dinosaur Lusotitan atalaiensis (Macronaria) and the evolutionary
history of basal titanosauriforms. Zoological Journal of the Linnean Society.
168(1), 98-206.
"Microdontosaurus"
Zhao, 1983 (preoccupied Gilmore, 1902)
"M. dayensis" Zhao, 1985
Etymology- The genus name refers to small teeth, while the species name
references the Daye township in Qamdo.
Middle Jurassic
Middle Dapuka Group, Dabuka, Qamdo County, Tibet, China
Material- specimen seemingly including teeth
Comments- As Gilmore (1902) already named an ichthyosaur Microdontosaurus
petersonii (currently a junior synonym of Baptanodon discus), Zhao's
sauropod will need a new name if it is officially described in the future. Glut
(1997) incorrectly dates the species to 1983 and places it in the Late Cretaceous.
Relationships- Stated by Zhao (1983) to be a homalosauropodoid, in which
he includes peg-toothed taxa like diplodocoids (including mamenchisaurids in
his view) and titanosaurs. Similarly, Zhao (1985) states it is a primitive peg-toothed
sauropod. Chure and McIntosh (1989) listed it as a cetiosaurid, Lambert (1990)
listed it as a diplodocid, and Olshevsky (1991) as a melanorosaurid. Glut (1997)
merely refers it to Sauropoda incertae sedis. Most recently, Fang et al. (2006)
place it in Brachiosauridae.
A placement outside of Sauropoda is unlikely for a Middle Jurassic taxon (though
not unheard of- see Yunnanosaurus? youngi), suggesting Olshevsky's assignment
is incorrect. While traditionally only diplodocoids and titanosaurids were seen
as having nonspatulate teeth, they are also found in more basal brachiosaur-grade
titanosauriforms and even some basal eusauropods like Shunosaurus. This
suggests Fang et al.'s assignment to Brachiosauridae may be most accurate, especially
as Zhao is a coauthor while Chure, McIntosh and Lambert have not seen the specimen.
Since Brachiosauridae as traditionally conceived is paraphyletic, "Microdontosaurus"
is here tentatively assigned to Titanosauriformes incertae sedis.
References- Gilmore, 1902. Discovery of teeth in Baptanodon, an
ichthyosaurian from the Jurassic of Wyoming. Science. 16(414), 913-914.
Zhao, 1983. Phylogeny and evolutionary stages of Dinosauria. Acta Palaeontologica
Polonica. 28(1-2), 295-306.
Zhao, 1985. The Jurassic Reptilia. In Wang, Cheng and Wang (eds.). The Jurassic
System of China. Stratigraphy of China. 11, 286-289, 347, plates 10 and 11.
Chure and McIntosh, 1989. A Bibliography of the Dinosauria (Exclusive of the
Aves) 1677-1986. Museum of Western Colorado Paleontology Series #1. 226 pp.
Lambert, 1990. The Dinosaur Data Book. New York: Avon Books. 320 pp.
Olshevsky, 1991. A Revision of the Parainfraclass Archosauria Cope, 1869, Excluding
the Advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.
Glut, 1997. Dinosaurs - The Encyclopedia. McFarland Press, Jefferson, NC. 1076
pp.
Fang, Zhang, Lu, Han, Zhao and Li, 2006. Collision between the Indian Plate
and the paleo-Asian late and the appearance of Asian dinosaurs. Geological Bulletin
of China. 25(7), 862-873.
Ornithopsidae Lydekker, 1887
Reference- Lydekker, 1887. On certain dinosaurian vertebrae from the
Cretaceous of India and the Isle of Wight. Quarterly Journal of the Geological
Society of London. 43, 156-160.
Ornithopsis
Pleurocoelidae Marsh, 1888
Reference- Marsh, 1888. Notice of a new genus of Sauropoda and other
new dinosaurs from the Potomac Formation. American Journal of Science. 35, 89-94.
Pleurocoelus
"Poekilopleuron" schmidti
Kiprijanow, 1883
= Megalosaurus schmidti (Kiprijanow, 1883) Olshevsky, 1991
Albian-Cenomanian, Early-Late Cretaceous
Sekmenevsk Formation (= Seweri Osteolith), Kurskaya Oblast, Russia
Syntypes- (ZIN? coll.) (titanosauriform) proximal metacarpal IV (~330 mm)
(ZIN? coll.) (cf. Pervushovisaurus bannovkensis) four incomplete jugals
Diagnosis- chimaera with no diagnostic elements.
Previous diagnoses- Kiprijanow (1883) did not provide a formal diagnosis,
but differentiated P. schmidti from P. bucklandii based on its
later age and geographic separation. While the former near certainly means it
is not conspecific, stratigraphy and biogeography are not currently viewed as
valid criteria for erecting a new species.
Comments- Kiprijanow (1883) described Poekilopleuron schmidti
based on a diapophysis and four slender fragments, supposedly a humerus and ribs.
He referred it to the Seweri Osteolith, which was recognized as being of Gault
age (Albian). Molnar (1990) noted its age was difficult to determine, but stated
it appeared to be Cenomanian-Santonian. Nessov (1995) believed it was Albian-Cenomanian
instead based on his study of the area and also noted Neogene mammal fossils
were present. While no catalogue numbers or repository were given by Kiprijanow,
Storrs et al. (2000) stated his material is presumably in the Russian Academy
of Sciences in St. Petersburg.
Description- The diapophysis is large, about 132 mm wide at its end. The shaft is cylindrical
and apparently hollow, with a bone wall to diameter ratio of ~23%. The end is
asymmetrically expanded primarily in one plane, with one corner projecting to
the side and end significantly more than the other. The less projected corner
has the broken base of a large bump on one side, and a lower but more proximodistally
elongate bump on the other side, the latter bump placed closer to the midline.
Though saddle-shaped in one view, the distal surface appears irregularly bumpy
in another, and no obvious condyles exist. The end's outline is roughly shaped
like a flattened parallelogram, with one of the short sides being about twice
as long as its opposite.
There are four slender fragments illustrated (table IV, figures 1-3).
All are slightly curved with one end (posterior?) gently expanded, and
broken both anteriorly and posteriorly (minimum depth ~20-26 mm). It is
difficult to tell which posterior edges are broken and which are
natural. The element in figure 1 has a longitudinal ridge down the
center of one side with fossae (perhaps caused by crushing) on either
side of it at the proximal end. The posterior cross section (figure 1C)
is very flat except for the ridge. The element in figure 3 is flat on
both sides and has a triangular cross section anteriorly (figure 3C').
Figure 2 seemingly consists of two elements judging by their different
lengths, with 2A having a rounded surface and 2B having a
longitudinally ridged surface.
Kiprijanow devotes several figures and most of
his description to schmidti's histology, which could prove useful to
someone knowledgable on that subject.
History- Kiprijanow referred this material to Poekilopleuron (then thought to
be a crocodile) because he thought the form of the elements was identical. However
this is obviously not so as even if there is a general resemblence to Poekilopleuron's
distal humerus in anterior view, side view shows at the very least schmidti
lacks Poekilopleuron's deltopectoral crest. Further differences from
theropod humeri are mentioned below. Rozhdestvensky (1973) reidentified the
diapophysis as a distal tibia, referring it to Megalosaurus sp. (he included
Poekilopleuron in that genus as well). This may be the origin of Olshevsky's
(1991) combination Megalosaurus schmidti (listed as a junior synonym
of P. schmidti), as it is not possible to sink a named species into "sp.",
regardless of how undiagnostic it is. It would be quite coincidental if schmidti
were still referrable to the same kind of animal, after the one somewhat diagnostic
fragment's position was reinterpreted. Yet schmidti differs from both
Poekilopleuron and Megalosaurus in lacking a shelf for the astragalar
ascending process and sharply tapered lateral edge in distal view. Efimov and
Chkhikvadze (1987) in a review of Soviet crocodilians, believed schmidti
to be dinosaurian. Molnar stated it "could belong to almost any large Late
Cretaceous theropod" and referred it to Theropoda indet., but no known
theropod has a similar distal tibia, leading me to doubt its identity (see below).
Molnar later (pers. comm. to Olshevsky, 1991) stated it may be ceratosaurid,
but Ceratosaurus differs in the same major ways as Megalosaurus
and Poekilopleuron.
Nessov was doubtful of its theropod identity, based on a "slit-like
pit" on the more broken side, "separating the region of contact with
other bones from the side of the diaphysis." I'm unsure which feature
Nessov was referring to, though the bone does have many cracks and
small pits, the latter largely from wear showing the internal texture.
Nessov thought the supposed rib fragments were not obviously
dinosaurian and probably belonged to marine reptiles (he discovered
ichthyosaur remains at the site). Arkhangelsky and Averianov (2003)
stated "the bones identified by Kiprianoff as ribs (1883, pl. 4) are
rather reminiscent of the ichthyosaur jugal." Holtz et al. (2004)
listed it as Tetanurae indet. without comment. Carrano et al. (2012)
described the main element as a humerus and tentatively confirmed its
theropod nature, while stating it could not be referred to Poekilopleuron.
Poekilopleuron schmidti is generally listed as Theropoda indet. in modern
references.
Metacarpal- The diapophysis was identified by Kiprijanow as a distal humerus and Rozhdestvensky
as a distal tibia. However, it differs from both of these elements. Dinosaurian
humeri have more symmetrical distal outlines as opposed to schmidti's
one side being much deeper. Both condyles in theropods are usually projected
anteriorly and rounded in side view, whereas schmidti has a prominent
and elongate projection on only one side and tapers towards the end in side
view. Most non-coelurosaur theropod tibiae have a deep shelf projecting diagonally
across the anterior end for reception of the astragalar ascending process, which
is lacking in schmidti. Coelurosaurs and a few other taxa lack this shelf,
but always have a much shallower distal end which is strongly tapered on one
side. This distal outline is even true of basal taxa like ceratosaurs and megalosauroids
(though not the much earlier coelophysoids, which differ more strongly in having
even deeper astragalar shelves), while coelurosaurs exaggerate it further. Another
point of difference is that theropod tibial shafts are wider compared to the
width of their distal transverse flare. Other dinosaur tibiae differ as well,
with ornithopods having a very strongly projected lateral flange for instance.
While schmidti bears a general resemblence to a proximal tibia, it lacks
all the detailed features one would expect- posterior intercondylar groove,
fibular crest, lateral condyle. Similarly, proximal fibulae are narrower, and
ulnae more triangular in proximal view. An exhaustive comparison of dinosaur
limb elements indicates it is near certainly a metapodial. The closest resemblence
among theropods is to the proximal end of metatarsal IV of large carnosaurs
such as Mapusaurus (mislabeled II in its description) and Chilantaisaurus.
The primary differences are schmidti would have a flat anterior edge
and a narrower shaft. Another similar element is metatarsal III in hadrosauriforms,
but these differ in being broader proximally and having a more pronounced anteroproximal
projection. More importantly, the anterior surface would be oddly angled in
schmidti and it lacks the anteroposterior expansion distally which is
a consequence of hadrosauriforms' short metatarsi. This makes it an unlikely
identification. The most similar element I could find among dinosaurs is the
proximal end of metacarpal IV of titanosauriforms. cf. Laplatasaurus'
metacarpal IV is nearly identical in proximal view, while Epachthosaurus'
is a close match in lateral view. The size is also well within the range of
titanosauriforms, while only giant theropods are of comparable size. One point
of contention is that schmidti's element is illustrated and described
as being hollow, but it's possible this is taphonomic as the end is broken and
extensively fractured, perhaps leading to the loss of a spongy interior as often
happens in recent mammal bones. Unfortunately, if we accept the sauropod identification
there is a high degree of variation ontogenetically/intraspecifically, so the
element is Titanosauriformes indet.. Most other sauropods had stouter metacarpals
and were extinct by the Mid Cretaceous in any case.
Jugals- Kiprijanow identified the slender fragments as ribs without specifying which part
of the vertebral column they derived from. Molnar merely said they may be fragments
of ribs, while Nessov thought they probably belonged to marine reptiles. There
is no evidence they belong to the same animal as the metapodial, as Kiprijanow
only says they "were found all together in the Seweri Osteolith" and
were fossilized identically. Unfortunately, dinosaurian rib morphology has been
largely ignored by most authors so comparison is difficult. Despite this, comparison
to theropod, sauropod and ornithopod dorsal ribs shows at least two of the elements
differ in having the proximal broad area too proximodistally extensive compared
to shaft width. In this they are more similar to theropod cervical ribs, such
as those of Tyrannosaurus. Assuming they are theropod cervical ribs,
they are not from abelisaurs since their shafts lack a flange. The concave and
possibly crushed areas on the one specimen might suggest pneumaticity as is
common in neotheropods, however the cross section shows it is solid. They are
also roughly similar to sauropod cervical ribs, though in medial view sauropods
have a strut extending perpendicular to the shaft from the tuberculum which
is missing in schmidti.
Another possibility for at least some of the elements is that they are
the medial ends of medial theropod gastralia (other dinosaurs of the
time lacked gastralia), which share the slightly expanded end and
transition from plate-like medial portion to rod-like lateral portion.
Arkhangelsky and Averianov's suggestion these are ichthyosaur jugals
seems extremely plausible. Co-occuring ichthyosaur material was
described by Kiprijanow as Ichthyosaurus campylodon,
long a wastebasket for Cretaceous Eurasian ichthyosaurs but restudied
by Fischer (2016) and restricted to Cenomanian English material as Pervushovisaurus campylodon. Pervushovisaurus itself is based on the Cenomanian Russian type species P. bannovkensis, contemporaneous with schmidti. While no jugals are reported for P. bannovkensis or P. campylodon, "Platypterygius" australis is a sister taxon of Pervushovisaurus in Fischer et al. (2016) and has a jugal identical to the schmidti "ribs" (Kear, 2005: Fig. 7E-F). Stratigraphy suggests the schmidti jugals are plausibly referrable to Pervushovisaurus bannovkensis, but it is also likely jugals are indeterminate within a subgroup of Platypteryginae.
References- Kiprijanow, 1883. Studien �ber die Fossilen Reptilien
Russlands. IV. Theil. Ordnung Crocodilina Oppel. Indeterminirte Fossile Reptilien. M�moires de l'Acad�mie Imp�riale des
Sciences de Saint-P�tersbourg. VIIe s�rie 31(7), 1-29.
Rozhdestvensky, 1973. [The study of Cretaceous reptiles in Russia]. Paleontologischeskii
Zhurnal. 1973, 90-99.
Efimov and Chkhikvadze, 1987. The review of finds of fossil Crocodiles of the
USSR. Procedings of the Academy of Sciences of the Georgian SSR. 13(3), 200-207.
Molnar, 1990. Problematic Theropoda: "Carnosaurs". In Weishampel,
Dodson and Osmolska (eds.). The Dinosauria. University of California
Press. 306-317.
Olshevsky, 1991. A revision of the parainfraclass Archosauria Cope, 1869, excluding
the advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.
Nessov, 1995. Dinosaurs of northern Eurasia: New data about assemblages, ecology,
and paleobiogeography. Institute for Scientific Research on the Earth's Crust,
St. Petersburg State University. 1-156.
Storrs, Arkhangelskii and Efimov, 2000. Mesozoic marine reptiles of Russia and
other former Soviet republics. In Benton, Shishkin, Unwin and Kurochkin (eds.).
The Age of Dinosaurs in Russia and Mongolia. Cambridge University Press. 187-210.
Arkhangelsky and Averianov, 2003. On the find of a primitive
hadrosauroid dinosaur (Ornithischia, Hadrosauroidea) in the Cretaceous
of the Belgorod region. Paleontological Journal. 37(1), 58-61.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria Second Edition. University of California Press. 71-110.
Kear, 2005. Cranial morphology of Platypterygius longmani
Wade, 1990 (Reptilia: Ichthyosauria) from the Lower Cretaceous of
Australia. Zoological Journal of the Linnean Society. 145, 583-622.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Fischer. 2016. Taxonomy of Platypterygius campylodon and the diversity of the last ichthyosaurs. PeerJ. 4:e2604.
Fischer, Bardet, Benson, Arkhangelsky and Friedman, 2016. Extinction of
fish-shaped marine reptiles associated with reduced evolutionary rates
and global environmental volatility. Nature Communications. 7:10825.
unnamed titanosauriform (Huene, 1929)
Late Cretaceous?
Neuquen?, Argentina
Material- (MACN 5017) humerus (210 mm)
Comments- MACN 5017 was referred to Argyrosaurus superbus by Huene
(1929), but Mannion and Otero (2012) noted it lacked the medial ridge diagnostic
for that taxon so referred it to Titanosauria indet.. The latter was based on
the medially expanded deltopectoral crest and prominent supracondylar fossa,
but these features are widespread in titanosauriforms in Mannion et al.'s macronarian
matrix, so the humerus is only identified to that level here.
References- Huene, 1929. Los Saurisquios y Ornitisquios del Cretaceo
Argentino. Anales del Museo de La Plata. 3, 1-196.
Mannion and Otero, 2012. A reappraisal of the Late Cretaceous Argentinean sauropod
dinosaur Argyrosaurus superbus, with a description of a new titanosaur
genus. Journal of Vertebrate Paleontology. 32(3), 614-638.
unnamed possible titanosauriform (Woodruff, 2012)
Aptian-Albian, Early Cretaceous
Cloverly Formation, Montana, US
Material- (MOR CL-121-1^B) pedal ungual I
Reference- Woodruff, 2012. A new titanosauriform from the Early Cretaceous
Cloverly Formation of Montana. Cretaceous Research. 36, 58-66.
Brachiosauridae Riggs, 1904
Definition- (Brachiosaurus altithorax <- Saltasaurus loricatus)
(Taylor, 2009; modified from Wilson and Sereno, 1998)
Other definitions- (Giraffatitan brancai <- Saltasaurus
loricatus) (Wilson, 2005)
References- Riggs, 1904. Structure and relationships of opisthocoelian
dinosaurs, part II: The Brachiosauridae. Field Columbian Museum Geological Series.
2(6), 229-247.
Wilson and Sereno, 1998. Early evolution and higher-level phylogeny of sauropod
dinosaurs. Society of Vertebrate Paleontology Memoir 5. Journal of Vertebrate
Paleontology. 18(2 suppl), 68 pp.
Wilson, 2005. Overview of sauropod phylogeny and evolution. in Curry Rogers
and Wilson (eds.). The Sauropods: Evolution and Paleobiology. University of
California Press, Berkeley. 15-49.
Taylor, 2009. A re-evaluation of Brachiosaurus altithorax Riggs 1903
(Dinosauria, Sauropoda) and its generic separation from Giraffatitan brancai
(Janensch 1914). Journal of Vertebrate Paleontology. 29(3), 787-806.
Padillasaurus Carballido,
Pol, Parra Ruge, Padilla Bernal, Paramo-Fonseca and Etayo-Serna, 2015
P. leivaensis Carballido, Pol, Parra Ruge, Padilla Bernal, Paramo-Fonseca
and Etayo-Serna, 2015
Early Barremian, Early Cretaceous
Paja Formation, Columbia
Holotype- (JACVM 0001) (adult) partial posterior dorsal vertebra (220 mm),
partial last dorsal vertebra, partial sacrum (?,?,?,?, 168 mm), incomplete first
caudal vertebra (115 mm), incomplete second caudal vertebra (135 mm), incomplete
third caudal vertebra (110 mm), incomplete fourth caudal vertebra (110 mm),
incomplete fifth caudal vertebra (113 mm), incomplete sixth caudal vertebra
(115 mm), incomplete seventh caudal vertebra (120 mm), incomplete eighth caudal
vertebra (125 mm)
Diagnosis- (after Carballido et al., 2015) first and second caudal vertebrae
with high and dorsally directed prezygodiapophyseal laminae that converge with
centroprezygapophyseal laminae and form lateroventral ventral margins of prezygapophyseal
processes; proximal caudal vertebrae with weakly laterally expanded transverse
processes; first caudal vertebrae with divided transverse process, the dorsal
section of which is posterodorsally directed in first two caudal vertebrae.
Comments- Carballido et al. (2015) recovered this taxon as a brachiosaurid
in their phylogenetic analysis.
Reference- Carballido, Pol, Parra Ruge, Padilla Bernal, Paramo-Fonseca
and Etayo-Serna, 2015. A new Early Cretaceous brachiosaurid (Dinosauria, Neosauropoda)
from northwestern Gondwana (Villa de Leiva, Colombia). Journal of Vertebrate
Paleontology. 35(5), e980505.
Rugocaudia Woodruff, 2012
R. cooneyi Woodruff, 2012
Late Albian, Early Cretaceous
Cloverly Formation, Montana, US
Holotype- (MOR 334) tooth (22x7.6x? mm), rib fragments, four proximal caudal
centra (112 mm), six partial to complete mid caudal centra (126 mm), ten fragmentary
to complete distal caudal centra (54-92 mm), caudal neural arch, partial chevron,
distal metacarpal, proximal femur, ?girdle and limb fragments
Diagnosis- (after Woodruff, 2012) paired, large depression on anterior
condyle margin near neural arch of proximal caudal vertebrae; mid caudal centra
square in transverse cross section; distalmost caudal vertebrae exhibit convex-upward
bend in lateral view.
Other diagnoses- D'Emic and Foreman (2012) noted two of Woodruff's diagnostic
characters (caudal condyle and cotyle margins greatly expressed with rugose
texture; neurovascular foramina expressed on condyle and cotyle margins with
greater frequency distally along caudal series) are due to damage and pathology.
They also stated that the other diagnostic characters listed are either not
present or are widespread in sauropods, thus considering Rugocaudia to
be a nomen dubium. However, the only other supposed autapomorphy mentioned by
them is the subequal length and height of the mid caudals, which are seen in
many taxa. While it is plausible the other characters listed above are also
invalid, they are retained here pending detailed study.
Comments- The material was discovered in 1985 and described as a new
taxon of titanosauriform by Woodruff (2012) without a phylogenetic analysis.
D'Emic and Foreman (2012) criticized the study, noting that some described morphologies
were not present in the material (e.g. the supposed second caudal is reversed,
so that no centra are actually procoelous). They stated Rugocaudia was
a brachiosaurid based on its lateral central fossae, which is confirmed when
the taxon is added to Mannion et al.'s macronarian matrix. Mannion et al. (2013)
placed Rugocaudia in Titanosauria based on procoelous caudals (despite
citing D'Emic and Foreman's paper), though agreed it was undiagnostic.
References- Woodruff, 2012. A new titanosauriform from the Early Cretaceous
Cloverly Formation of Montana. Cretaceous Research. 36, 58-66.
D'Emic and Foreman, 2012. The beginning of the sauropod dinosaur hiatus in North
America: Insights from the Lower Cretaceous Cloverly Formation of Wyoming. Journal
of Vertebrate Paleontology. 32(4), 883-902.
Mannion, Upchurch, Barnes and Mateus, 2013. Osteology of the Late Jurassic Portuguese
sauropod dinosaur Lusotitan atalaiensis (Macronaria) and the evolutionary
history of basal titanosauriforms. Zoological Journal of the Linnean Society.
168(1), 98-206.
Venenosaurus
Europasaurus
Brachiosaurinae Riggs, 1904 vide Janensch, 1929
Reference- Janensch, 1929. Material und Formegehalt der Sauropoden in
der Ausbeute der Tendaguru-Expedition, 1909-1912 [Material and figured content
of sauropods in the yield of the Tendaguru Expedition, 1909-1912]. Palaeontographica.
Supplement VII (1), 2(1), 3-34.
Brachiosaurus
Abydosaurus Chure, Britt,
Whitlock and Wilson, 2010
A. mcintoshi Chure, Britt, Whitlock and Wilson, 2010
Mid-Late Albian, Early Cretaceous
Mussentuchit Member of the Cedar Mountain Formation, Utah, US
Holotype- (DINO 16488) skull (~500 mm), sclerotic plates, mandibles, hyoid,
atlas, partial axis, partial third cervical vertebra (230 mm), partial fourth
cervical vertebra (320 mm)
Paratypes- (DINO 17848) partial skull, partial mandibles
(DINO 17849) incomplete skull
(DINO 39728) skull roof, braincase
(DINO coll.) sacrum, proximal caudal series, scapula, partial humerus (~1.6
m), metacarpus, partial pelvis
Diagnosis- (after Chure et al., 2010) nasal lateral process directed
anteriorly and lacking posterior hook; maxilla narial process with dorsal articulation
for nasal; nasals overlap asymmetrically on midline; external naris smaller
than orbit; upper crown apices shifted distally; upper tooth shafts twist through
an arc of nearly 45 degrees; tooth wear present only mesially.
Comments- Chure et al. (2010) added Abydosaurus to a version of
Wilson's sauropod analysis and recovered it as a brachiosaurid. This has been
supported by D'Emic (2012) who found it as a derived brachiosaurid closest to
Venenosaurus and Cedarosaurus, and Mannion et al. (2013) who recovered
it as a derived brachiosaurid close to Cedarosaurus and sometimes Venenosaurus,
Giraffatitan and Sonorasaurus.
References- Chure, Britt, Whitlock and Wilson, 2010. First complete sauropod
dinosaur skull from the Cretaceous of the Americas and the evolution of sauropod
dentition. Naturwissenschaften. 97(4), 379-391.
D'Emic, 2012. The early evolution of titanosauriform sauropod dinosaurs. Zoological
Journal of the Linnean Society. 166, 624-671.
Mannion, Upchurch, Barnes and Mateus, 2013. Osteology of the Late Jurassic Portuguese
sauropod dinosaur Lusotitan atalaiensis (Macronaria) and the evolutionary
history of basal titanosauriforms. Zoological Journal of the Linnean Society.
168(1), 98-206.
Giraffatitan
Cedarosaurus Tidwell, Carpenter
and Brooks, 1999
C. weiskopfae Tidwell, Carpenter and Brooks, 1999
Barremian, Early Cretaceous
Yellowcat Member of the Cedar Mountain Formation, Utah, US
Holotype- (DMNH 39045) (adult) partial to incomplete fourth-twelfth dorsal
vertebrae (4th 260, 12th 100 mm), several incomplete dorsal ribs, first caudal
vertebra, second caudal vertebra (92 mm), third caudal vertebra (95 mm), fourth
caudal vertebra (113 mm), fifth caudal vertebra (95 mm), sixth caudal vertebra
(97 mm), seventh caudal vertebra (105 mm), eighth caudal vertebra (112 mm),
ninth caudal vertebra (107 mm), partial tenth caudal vertebra, eleventh caudal
vertebra (113 mm), twelfth caudal vertebra (102 mm), thirteenth caudal vertebra
(100 mm), fourteenth caudal vertebra (98 mm), fifteenth caudal vertebra (101
mm), sixteenth caudal vertebra (100 mm), seventeenth caudal vertebra (105 mm),
eighteenth caudal vertebra (112 mm), nineteenth caudal vertebra (100 mm), twentieth
caudal vertebra (117 mm), twenty-first caudal vertebra (115 mm), twenty-second
caudal vertebra (112 mm), twenty-third caudal vertebra (110 mm), twenty-fourth
caudal vertebra (111 mm), twenty-fifth caudal vertebra (105 mm), several chevrons
(all but one partial), proximal scapulae, coracoids, incomplete sternal plates,
humerus (1.38 m), radius (812 mm), incomplete ulna, incomplete metacarpal IV
(428 mm), incomplete metacarpal, pubes (one partial), proximal ischia, femora
(one proximal, one incomplete; 1.395 m), tibia (~884 mm), metatarsal I (152
mm), pedal ungual I, metatarsal II (201 mm), phalanx II-1, pedal ungual II,
pedal ungual III, metatarsal V, gastroliths
Early Albian, Early Cretaceous
Paluxy Formation of the Trinity Group, Texas, US
Referred- (FMNH PR 977) distal tibia, distal fibula, astragalus, metatarsal
I (121 mm), phalanx I-1 (71 mm), incomplete pedal ungual I (137 mm), incomplete
metatarsal II (200 mm), phalanx II-1 (63 mm), pedal ungual II (114 mm), metatarsal
III (165 mm), phalanx III-1 (50 mm), phalanx III-2, pedal ungual III (112 mm),
proximal metatarsal IV, phalanx IV-1 (62 mm), phalanx IV-2, partial pedal ungual
IV (80 mm), metatarsal V (215 mm), phalanx V-1 (Langston, 1974; described by
Gallup, 1975)
Diagnosis- (after Tidwell et al., 1999) proximal caudal vertebrae with
deeply concave anterior faces; proximal caudal vertebrae lack well developed
hyposphenes; proximal caudal vertebrae with flat to concave posterior faces;
mid caudal vertebrae with sharp ridge extending axially along sides of neural
arch; deltopectoral crest placed closer to midshaft than Brachiosaurus
or Pleurocoelus; humerofemoral ratio (.98) higher than Pleurocoelus
(also in Brachiosaurus); humerus more robust than Brachiosaurus;
radius slender with two ridges beginning at mid shaft, one of which extends
along the distal posterior side, the other curves along
the lateral side to terminate in a prominent rugosity near distal end unlike
Brachiosaurus; distal radius subrectangular and flat; ulna with prominent
posterior condyle unlike Brachiosaurus (also in Pelorosaurus);
distinct ulnar groove separating distal condyle from lateral wall unlike Pleurocoelus.
(after D'Emic, 2013) radius with tubercle on anterior face of shaft, one-third
from the proximal end; metatarsal II with well-developed medial and lateral
tubercles at mid-shaft; pedal ungual IV present; metatarsal V only slightly
expanded proximally; metatarsal V around 1.5 times longer than metatarsal I;
phalanx V-1 present.
Other diagnoses- Tidwell et al. (1999) also stated Cedarosaurus
was unlike Astrophocaudia (SMU 61732) in having a "deltopectoral
crest placed closer to midshaft" and "a distinct groove separating
the
distal condyle from the lateral wall" of the ulna, but neither humerus
or ulna are preserved in that taxon.
Comments- The holotype was discovered in 1996. Tidwell et al. (1999)
referred Cedarosaurus to Brachiosauridae without a phylogenetic analysis.
This has been confirmed by D'Emic (2012) who recovered it as a derived brachiosaurid
closest to Venenosaurus and Abydosaurus, and Mannion et al. (2013)
who find it as a derived brachiosaurid closest to Giraffatitan, Venenosaurus
and Abydosaurus (and sometimes Sonorasaurus).
Langston (1974) mentioned a sauropod hindlimb (FMNH PR 977) that was described
in Gallup's (1975) thesis as Pleurocoelus sp.. Gallup published a brief
description in 1989, but note his "in press" Journal of Paleontology
paper listed in the bibliography never made it to print. D'Emic (2013) finally
referred this specimen to Cedarosaurus based on a few autapomorphies.
References- Langston, 1974. Nonmammalian Comanchean tetrapods. Geoscience
and Man. 8, 77-102.
Gallup, 1975. Lower Cretaceous dinosaurs and associated vertebrates from north-central
Texas in the Field Museum of Natural History. MS thesis, University of Texas
at Austin. 159 pp.
Gallup, 1989. Functional morphology of the hindfoot of the Texas sauropod Pleurocoelus
sp. indet.. Geological Society of America Special Paper. 238, 71-74.
Gallup, "1989" [unpublished]. The four-clawed Lower Cretaceous dinosaur
Pleurocoelus Marsh from the Paluxy Formation of north-central Texas.
Journal of Paleontology.
Tidwell, Carpenter and Brooks, 1999. New sauropod from the Lower Cretaceous
of Utah, USA. Oryctos. 2, 21-37.
D'Emic, 2012. The early evolution of titanosauriform sauropod dinosaurs. Zoological
Journal of the Linnean Society. 166, 624-671.
D'Emic, 2013. Revision of the sauropod dinosaurs of the Lower Cretaceous Trinity
Group, southern USA, with the description of a new genus. Journal of Systematic
Palaeontology. 11(6), 707-726.
Mannion, Upchurch, Barnes and Mateus, 2013. Osteology of the Late Jurassic Portuguese
sauropod dinosaur Lusotitan atalaiensis (Macronaria) and the evolutionary
history of basal titanosauriforms. Zoological Journal of the Linnean Society.
168(1), 98-206.
Somphospondyli Wilson and Sereno, 1998
Official Definition- (Saltasaurus loricatus <- Giraffatitan
brancai)
(Silva Junior, Martinelli, Lori, Marinho, Hechenleitner and Langer,
2022; originally Wilson, 2005; Registration Number 436)
Other definitions- (Saltasaurus loricatus <- Brachiosaurus altithorax)
(Taylor, 2009; modified from Wilson and Sereno, 1998)
References- Wilson and Sereno, 1998. Early evolution and higher-level
phylogeny of sauropod dinosaurs. Society of Vertebrate Paleontology Memoir 5.
Journal of Vertebrate Paleontology. 18(2 suppl), 68 pp.
Wilson, 2005. Overview of sauropod phylogeny and evolution. in Curry Rogers
and Wilson (eds.). The Sauropods: Evolution and Paleobiology. University of
California Press, Berkeley. 15-49.
Taylor, 2009. A re-evaluation of Brachiosaurus altithorax Riggs 1903
(Dinosauria, Sauropoda) and its generic separation from Giraffatitan brancai
(Janensch 1914). Journal of Vertebrate Paleontology. 29(3), 787-806.
Silva Junior, Martinelli, Lori, Marinho, Hechenleitner and Langer, 2022 (online 2021). Reassessment of Aeolosaurus maximus,
a titanosaur dinosaur from the Late Cretaceous of southeastern Brazil.
Historical Biology. 34(3), 403-411.
Titanosauria Haeckel, 1889
Official Definition-
(Andesaurus delgadoi + Saltasaurus loricatus) (Silva Junior, Martinelli, Lori, Marinho, Hechenleitner and Langer, 2022; originally Wilson and Upchurch,
2003; Registration Number 435)
Other definitions- (Andesaurus delgadoi + Titanosaurus indicus)
(modified from Salgado et al., 1997)
(Saltasaurus loricatus <- Brachiosaurus
altithorax, Euhelopus zdanskyi) (modified from Wilson and Sereno,
1998)
(Saltasaurus loricatus <- Euhelopus zdanskyi) (modified from
Sereno, 1998)
(Saltasaurus loricatus <- Brachiosaurus altithorax) (modified
from Upchurch et al., 2004)
= "Balochisauridae" Malkani, 2003
= Poripuchia Malkani, 2020
Definition- ("Balochisaurus malkani" + Isisaurus colberti + Gspsaurus pakistani + "Maojandino alami" + "Marisaurus jeffi" + Pakisaurus balochistani + Saraikimasoom
vitakri + Sulaimanisaurus gingerichi) (Malkani, 2021)
Other definition- (Isisaurus colberti, Gspsaurus pakistani, Pakisaurus balochistani, Saraikimasoom
vitakri within Titanosauria) (modified after Malkani, 2020)
Gspsauridae Malkani, 2021
= "Gspsauridae" Malkani, 2015
Definition- (Gspsaurus pakistani + "Marisaurus jeffi" + "Maojandino alami" + "Balochisaurus malkani" + "Nicksaurus razashahi" + Saraikimasoom vitakri) (modified from Malkani, 2021)
Gspsaurinae Malkani, 2021
= "Gspsaurinae" Malkani, 2015
Definition- (Gspsaurus pakistani + "Marisaurus jeffi" + "Maojandino alami") (modified from Malkani, 2021)
Saraikimasoominae Malkani, 2021
= "Saraikimasoominae" Malkani, 2015
Definition- ("Balochisaurus malkani" + "Nicksaurus razashahi" + Saraikimasoom vitakri) (modified from Malkani, 2021)
Pakisauridae Malkani, 2021
= "Pakisauridae" Malkani, 2003
Definition- (Isisaurus colberti + Khetranisaurus barkhani + Pakisaurus balochistani + Sulaimanisaurus gingerichi) (modified from Malkani, 2021)
Comments- Haeckel (1889) was the first to use Titanosauria, as a dinosaurian
group including "elephantine dragons" such as Iguanodon and
Atlantosaurus. This was not widely adopted, and while titanosaurs were
sometimes called "titanosaurians" through the 1900s, Bonaparte and
Coria (1993) were the first to use the same again in a systematic context.
Malkani (2020) proposed a new clade Poripuchia as "the most inclusive clade of Titanosauria containing Pakisaurus and Isisaurus pakisaurids, and Gspsaurus and Saraikimasoom
gspsaurids titanosaurs" which by definition would make it synonymous
with Titanosauria. He no doubt meant the LEAST inclusive clade
and in 2021 more clearly defined it as "Pakisaurus, Sulaimanisaurus, Isisaurus, Gspsaurus, Saraikimasoom, Balochisaurus, Marisaurus, Maojandino,
their most recent common ancestor and all of its descendants."
All but one taxon in either definition are Malkani's own Pakistani
genera which have never been included in phylogenetic analyses, so the
content of Poripuchia is particularly uncertain even given the
unsettled state of titanosaur relationships. Malkani diagnosed it
based on having all caudals procoelous (except the
biconvex first one in some taxa), which would probably fall somewhere
within Lithostrotia. Note Malkani gives a
2019a paper on Saraikimassom as the original reference for Poripuchia
in his systematic section, but that publication never mentions
it. Its earliest unofficial mention may be his 2019b paper on Pakisaurus which stated "Pakisaurus balochistani
is recognised as most derived Poripuch lithostrotian titanosaur based
on procoelous anterior, middle and posterior caudal vertebrae."
Malkani (2003) first mentioned balochisaurids and
pakisaurids, but only informally as an alternative to saltasaurids and
titanosaurids respectively. Malkani (2006) later tried to formally
erect Balochisauridae as a senior synonym of Saltasauridae and
Pakisauridae as a senior synonym of Titanosauridae, but Saltasauridae
and Titanosauridae would have priority and no justification for
replacement is given. Malkani (2006) includes "Balochisaurus",
"Marisaurus" and presumably Saltasaurus in Balochisauridae, and Pakisaurus, Khetranisaurus, Sulaimanisaurus,
Isisaurus and sometimes Brohisaurus
in Pakisauridae, but the families were invalid in 2006 as neither
eponymous genus was nomenclaturally valid (ICZN Article 11.7.1.1). By
2015 Malkani added his new taxon "Maojiandino" to balochisaurids, and
created the new family Gspsauridae divided into two monotypic
subfamilies- Gspsaurus in Gspsaurinae and Saraikimasoom
in Saraikimasoominae. These new taxa were also invalid based on
Article 11.7.1.1. Malkani's first publication that regularly follows
ICZN rules is his 2021 paper that erects Pakisauridae and Gspsauridae
with the latter basically replacing "Balochisauridae" and including
polytypic Gspsaurinae and Saraikimasoominae. This paper also provides
phylogenetic definitions for each group, but these only use Malkani's
genera and Isisaurus. As Isisaurus
has an unstable position within Lithostrotia and Malkani's taxa have
never been included in a phylogenetic analysis, or had their validity
or composition independently analyzed, Malkani's hierarchy is largely
incomparable to the consensus titanosaur phylogeny. If Malkani's ideas
are correct, the names may correspond to lithostrotian clades, but they
are conservatively listed under Titanosauria here pending further
analysis.
References- Haeckel, 1889. Nat�rliche Sch�pfungs-Geschichte.
Druck und Verlag von Georg Reimer. 832 pp.
Bonaparte and Coria, 1993. Un neuvo y gigantesco saur�podo titanosaurio
de la Formaci�n Rio Limay (Albanio-Cenomaniano) de la Provincia del Neuqu�n,
Argentina [A new and huge titanosaur sauropod from the Rio Limay Formation (Albian-Cenomanian)
of Neuqu�n Province, Argentina]. Ameghiniana. 30(3), 271-282.
Salgado, Coria and Calvo, 1997. Evolution of titanosaurid sauropods. I: Phylogenetic
analysis based on the postcranial evidence. Ameghiniana. 34(1), 3-32.
Sereno, 1998. A rationale for phylogenetic definitions, with application to
the higher-level taxonomy of Dinosauria. Neues Jahrbuch f�r Geologie und
Pal�ontologie Abhandlungen. 210, 41-83.
Wilson and Sereno, 1998. Early evolution and higher-level phylogeny of sauropod
dinosaurs. Society of Vertebrate Paleontology Memoir 5. Journal of Vertebrate
Paleontology. 18(2 suppl), 68 pp.
Malkani, 2003. Discovery of partial skull and dentary of
Titanosauria (sauropod dinosaur) from the Late Cretaceous Pab Formation of Vitakri
area, Barkhan district, Balochistan, Pakistan. Geological Bulletin University
of Peshawar. 36, 65-71.
Wilson and Upchurch, 2003. A revision of Titanosaurus Lydekker (Dinosauria
- Sauropoda), the first dinosaur genus with a 'Gondwanan' distribution. Journal
of Systematic Palaeontology. 1(3), 125-160.
Upchurch, Barrett and Dodson, 2004. Sauropoda. in Weishampel, Dodson and Osmolska
(eds.). The Dinosauria (2nd edition). University of California Press, Berkeley.
259-322.
Malkani, 2006. Biodiversity of saurischian dinosaurs from the Latest Cretaceous
park of Pakistan. Journal of Applied and Emerging Sciences. 1(3), 108-140.
Malkani, 2015. Dinosaurs, mesoeucrocodiles, pterosaurs, new
fauna and flora from Pakistan. Geological Survey of Pakistan
Information Release. 823, 32 pp.
Malkani, 2019a. Smallest titanosaur from Indo- Pakistan landmass. Open Journal of Geology. 9, 627-630.
Malkani, 2019b. Large titanosaur from Indo-Pakistan peninsula. Open Journal of Geology. 9, 635-638.
Malkani, 2020. First snout with complete teeth row of small titanosaur
in Indo-Pakistan found from the Latest Maastrichtian Vitakri Formation
of Pakistan; Associated cranial and postcranial skeletons of Saraikimasoom vitakri
(Poripuchia, stocky Titanosauria, Sauropoda) from Pakistan and referred
fossils from India. Open Journal of Geology. 10, 368-407.
Malkani,
2021. Jurassic-Cretaceous and Cretaceous-Paleogene transitions and
Mesozoic vertebrates from Pakistan. Open Journal of Geology. 11,
275-318.
Silva Junior, Martinelli, Lori, Marinho, Hechenleitner and Langer, 2022 (online 2021). Reassessment of Aeolosaurus maximus,
a titanosaur dinosaur from the Late Cretaceous of southeastern Brazil.
Historical Biology. 34(3), 403-411.
Somphospondyli/Titanosauria incertae sedis
Agustiniidae Bonaparte, 1999
Reference- Bonaparte, 1999. An armoured sauropod from the Aptian of northern
Patagonia, Argentina. in Tomida, Rich and Vickers-Rich (eds.). Proceedings of
the Second Gondwanan Dinosaur Symposium, National Science Museum Monographs.
15, 1-12.
Agustinia
Arkharavia Alifanov and
Bolotsky, 2010a
A. heterocoelica Alifanov and Bolotsky, 2010a
Late Maastrichtian, Late Cretaceous
Udurchukan Formation of the Tsagayan Group, Russia
Holotype- (AEHM 2/418) incomplete proximal caudal centrum (65 mm)
Diagnosis- (after Alifanov and Bolotsky, 2010b) proximal caudal centra
narrow in lateral view; proximal caudal articular surfaces saddle-shaped- initially
platycoelous anterior surface curved mostly because of slight anterior inclination
of upper part (concave in lateral view), while posterior part because of elongation
of lateral margins of initially platyamphicoelous surface (concave in ventral
and dorsal views).
Other diagnoses- Alifanov and Bolotsky (2010a, b) also included characters
based on the paratype vertebrae in their diagnosis- proximal caudal neural spine
very high and narrow; proximal caudal neural spines have anterior groove at
their base. However, these specimens are now known to be hadrosaurids (see below).
Comments- The holotype was discovered between 2000 and 2004, and described
by Alifanov and Bolotsky (2010a, b) as a new taxon of titanosauriform closely
related to Chubutisaurus. However, Godefroit et al. (2012) stated the
type vertebrae were likely hadrosaurid instead. Mannion et al. (2013) confirmed
paratype AEHM 2/720 is hadrosaurid based on neural spine anatomy, and this would
also hold true for the figured paratype AEHM 2/997. Another paratype (AEHM 2/420)
is said to have the same kind of neural spine, so may also be hadrosaurid, but
the final one (AEHM 2/419) is neither figured nor remarked upon, so we have
no information to judge it from. Several hadrosaurid taxa are known from the
Udurchukan Formation (Amurosaurus, Kerberosaurus, Olorotitan)
and the correlated Yuliangze Formation (Charonosaurus, Mandschurosaurus,
Sahaliyania, Wulagasaurus), but only a few of these have associated
caudals, and none have been distinguished using caudal characters. Mannion et
al. stated the holotype of Arkharavia is undiagnostic, though they did
not address Alifanov and Bolotsky's proposed autapomorphies, and referred it
to Somphospondyli based on the mildly procoelous and short proximal caudal centrum.
They correctly noted the referred tooth (AEHM 1/316) is from another locality
and cannot be compared to the holotype vertebra, but assigned it to Titanosauria
based on its cylindrical shape.
References- Alifanov and Bolotsky, 2010a. Arkharavia heterocoelica
gen. et sp. nov., a new sauropod dinosaur from the Upper Cretaceous of far earstern
Russia. Paleontologicheskii Zhurnal. 2010(1), 76-83.
Alifanov and Bolotsky, 2010b. Arkharavia heterocoelica gen. et sp. nov.,
a new sauropod dinosaur from the Upper Cretaceous of the far East of Russia.
Paleontological Journal. 44(1), 84-91.
Godefroit, Bolotsky and Bolotsky, 2012. Osteology and relationships of Olorotitan
arharensis, a hollow-crested hadrosaurid dinosaur from the latest Cretaceous
of far eastern Russia. Acta Palaeontologica Polonica. 57(3), 527-560.
Mannion, Upchurch, Barnes and Mateus, 2013. Osteology of the Late Jurassic Portuguese
sauropod dinosaur Lusotitan atalaiensis (Macronaria) and the evolutionary
history of basal titanosauriforms. Zoological Journal of the Linnean Society.
168(1), 98-206.
Bruhathkayosaurus Yadagiri and Ayyasami, 1987
B. matleyi Yadagiri and Ayyasami, 1987
Late Maastrichtian, Late Cretaceous
Kallamedu Formation, India
Holotype- (GSI PAL/SR/20; lost) ilium (1.2 m)
Paratypes- ...?(GSI coll.; lost) incomplete caudal vertebra, radius, pubis, distal femur, tibia (2 m)
Other diagnoses- (after
Yadagiri and Ayyasami, 1987) "heavy limb bones, the tibia being the
largest; the ilium large with heavy puboischiac peduncle and
posteriorly massive."
Comments- Glut (1982) reported
"a completely preserved pelvic girdle and limb bones of a theropod"
from the Kallamedu Formation, which were later described as a new taxon
of carnosaur sensu lato (Yadagiri and Ayyasami, 1987). Only the
ilium and tibia were illustrated and photographed, with at least the
photocopies of the latter being unintelligable. The supposed
dorsal and posterior views of figures 2a and 2b only make sense as a
ventral and lateral view of a right ilium. The "heavy puboischiac
peduncle" seemingly refers to the long and distally expanded pubic
peduncle illustrated, while the "posteriorly massive" portion of the
ilium might be the postacetabular process which is illustrated with a
feature taking up the posterior margin (if convex, perhaps like the
lateral boss of therizinosaurids). "Two marked depressions are
present on dorsal portion of the ilium meant for thick muscular
attachment", typical of saurischians (M. iliofemoralis and M.
iliofibularis). The tibia is described as twisted, as in most
dinosaurs, with proximal and distal widths of 650 and 600 mm
respectively. The femur is stated to have a transverse distal
width of 750 mm, and "the forth [sic] trochanter could not be
traced." The caudal vertebra was stated to be platycoelous and
lacks a neural spine, while the radius and pubis are only mentioned
without description. Olshevsky (1994) summarized the paper's
findings, stating the "symposium [was] held in 1987 but papers
apparently not published until 1989" which would make the publication
date 1989, but this has yet to be supported by other sources.
Olshevsky (DML, 1995) stated Chatterjee "notes that Bruhathkayosaurus,
formerly the titanic theropod, is almost certainly a titanic
titanosaurid sauropod", probably based on Chatterjee's 1995
semi-popular article. Similarly, Ford (DML, 1999) stated "Sankar
Chatterjee has
told George and I that Bruhathkayosaurus is a titanosaur (he did look at the material)." These are seemingly the sources used to refer Bruhathkayosaurus
to Sauropoda in references such as Upchurch et al, (2004), Krause et
al. (2006) and Hone et al. (2016). Rumors it may be petrified
wood seem to date to Holtz (DML, 1995; "The drawings and photographs of
the fossil are horrible, and I see no evidence that these aren't
petrified wood!") and have not been supported by proposed
evidence or observation. Upon inquiry, Chatterjee wrote (pers. comm.
2000) "The material is very fragmentary, but the ilium and the femur
looks like those of titanosaurids, very massive with high iliac crest.
The only theropod occurs in this region is abelisaurids. We have
recently collected almost complete postcranial skeleton of abelisaurid
similar to Indosuchus [later described as Rahiolisaurus]. They are quite different from Bruhathkayosaurus." Unfortunately, Galton and Ayyasami (2017) note "the photographs used for Bruhathkayosaurus
by Yadagari & Ayyasami (1987) were taken while the bones were still
exposed in situ. The bones started to disintegrate in the field jackets
even before reaching the GSIH and no longer exist." Zack A. under
the pseudonym palaeozoologist wrote (online, 2011) "from email
correspondence with Dr. Ayyasami, it appears that the material was
never actually properly prepared and was left exposed to the elements
so when heavy monsoon rains struck the region the fossils were carried
away in the rains. He says that the material was definitely dinosaurian
in nature, and apparently Dr. Sankar Chatterjee also was able to
confirm its dinosaurian (and apparently, titanosaurian) nature."
Thus presenting an alternative route to an identical conclusion.
The loss of the material is unfortunate, as Yadagiri and Ayyasami's
illustrations do not match titanosaur anatomy well. The elongate
ischial peduncle and long postacetabular process are more similar to
avepods, and the preacetabuilar process is not flared much laterally
unlike neosauropods. The greatly expanded pubic peduncle is like
several titanosaurs however, and Opisthocoelicaudia has an excavated postacetabular tip that could match Bruhathkayosaurus'
illustration. The tibia is more gracile than expected (proximal
width / total length and distal width / total length 33% and 30% vs.
44% and 37% in Opisthocoelicaudia),
better matching a titanosaur fibula, although these are not
twisted. It is also much longer than expected for a titanosaur
(tibio-ilial ratio 167% compared to 71% in Opisthocoelicaudia; tibia length compared to distal femoral width 267% vs. 188% in Opisthocoelicaudia).
The platycoelous caudal vertebra is unlike the procoelous caudals of
most titanosaurs, and surely any radius referred to a carnosaur would
be too small to belong to the type individual if it were
sauropodan. Olshevsky (1994) brought up the possibility of this
being a chimaera, which is plausible as it was found disarticulated in
a 30 x 10 meter pit. In the end we are left dealing with poor
drawings and the unpublished opinion of Chatterjee, neither of which
allows for a confident conclusion.
References- Glut, 1982. The New Dinosaur Dictionary. Citadel Press. 288 pp.
Yadagiri and Ayyasami, 1987. A carnosaurian dinosaur from the Kallamedu
Formation (Maestrichtian horizon), Tamilnadu. In Sastry, Sastry,
Ramanujam, Kapoor, Jagannatha Rao, Satsangi and Mathur (eds.).
Symposium on Three Decades of Development in Palaeontology and
Stratigraphy in India. Volume 1. Precambrian to Mesozoic. Geological
Society of India Special Publication. 11(1), 523-528.
Olshevsky, 1994. Bruhathkayosaurus: Bigger than T. rex? The Dinosaur Report. Winter 1994, 12-13.
Chatterjee, 1995. The last dinosaurs of India. The Dinosaur Report. Fall 1995, 12-18.
Holtz, DML 1995. https://web.archive.org/web/20190225104444/http://dml.cmnh.org/1995Sep/msg00701.html
Olshevsky, DML 1995. https://web.archive.org/web/20201110061419/http://dml.cmnh.org/1995Nov/msg00158.html
Ford, DML 1999. https://web.archive.org/web/20200112213609/http://dml.cmnh.org/1999Mar/msg00516.html
Upchurch, Barrett and Dodson, 2004. Sauropoda. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria (2nd edition). University of California Press.
259-322.
Krause, O'Connor, Curry Rogers, Sampson, Buckley and Rogers, 2006. Late
Cretaceous terrestrial vertebrates from Madagascar: Implications for
Latin American biogeography. Annals of the Missouri Botanical Garden. 93(2), 178-208.
Zack A, online 2011. https://www.deviantart.com/comments/1/221544713/1839677900
Hone, Farke and Wedel, 2016. Ontogeny and the fossil record: What, if
anything, is an adult dinosaur. Biological Letters. 12, 20150947.
Galton and Ayyasami, 2017. Purported latest bone of a
plated dinosaur (Ornithischia: Stegosauria), a "dermal plate" from the
Maastrichtian (Upper Cretaceous) of southern India. Neues Jahrbuch f�r Geologie und Pal�ontologie Abhandlungen. 285(1), 91-96.
Clasmodosaurus
References- Huene, 1929. Los saurisquios y ornitisquios del Cretac�o
Argentino. Anales del Museo de La Plata (series 3). 3, 1-196.
"Megacervixosaurus"
Zhao, 1983
"M. tibetensis" Zhao, 1986
Etymology- The genus name refers to a large neck, while the species name
refers to Tibet where the remains were found.
Late Cretaceous
Zonggu Formation, Zonggu, Markam County, Tibet, China
Material- specimen including cervical vertebrae
Comments- Zhao (pers. comm. in Molnar, 2011) noted similarity of its
posterior cervicals to Austrosaurus.
Relationships- Stated by Zhao (1983) to be a homalosauropodoid, in which
he includes peg-toothed taxa like diplodocoids and titanosaurs. Lambert (1990),
Olshevsky (1991) and provisionally Glut (1997) listed it as a diplodocid. Olshevsky
(online, 1999) also suggested it may be mamenchisaurid, perhaps based on the
name, though Zhao did place Mamenchisaurus in Homalosauropodoidea too.
Weishampel et al. (2004) questioningly listed it as a lithostrotian.
As no morphological data has been released on "Megacervixosaurus",
its relationships must necessarily remain tentative. The absence of Cretaceous
or Asian diplodocids renders that identification suspect, while mamenchisaurids
are also unknown from the Cretaceous. Indeed, the only Late Cretaceous sauropods
are rebbachisaurids and somphospondyls, and only the latter are known from East
Asia. "Megacervixosaurus" is here considered a probable somphospondylan.
References- Zhao, 1983. Phylogeny and evolutionary stages of Dinosauria.
Acta Palaeontologica Polonica. 28(1-2), 295-306.
Yang, 1986. The Cretaceous System. in Yang, Cheng and Wang (eds.). The Geology
of China. Clarendon Press. 153-167.
Zhao, 1986. The Cretaceous biota of China: Reptilia. in Hao, Su, Yu and Li (eds.).
The Cretaceous System of China. Stratigraphy of China. 12, 67-73, plates XI,
XII.
Molnar, 2011. New morphological information about Cretaceous sauropod dinosaurs
from the Eromanga Basin, Queensland, Australia. Alcheringa. 35, 329-339.
Vahiny Curry Rogers and Wilson,
2014
V. depereti Curry Rogers and Wilson, 2014
Middle Maastrichtian, Late Cretaceous
Anembalemba Member of the Maevarano Formation, Madagascar
Holotype- (UA 9940) incomplete braincase
Paratype- (FMNH PR 3046) (juvenile) supraoccipital
Diagnosis- (after Curry Rogers and Wilson, 2014) basal tubera composed
mainlyof basioccipital, with reduced contribution from basisphenoid; thick web
of bone separating basipterygoid embayment from basal tubera, visible posteriorly;
paired blind fossae present between basipterygoid processes and parasphenoid
rostrum; ventrally keeled and dorsally troughed parasphenoid rostrum; foramina
for cranial nerves II, III and V co-linear; metotic foramen displaced ventrally
from cranial nerves II, III and V.
Comments- The holotype was discovered in 2005. Note that based on the
presence of only two titanosaur morphotypes in the Maevarano Formation, it's
nearly certain Vahiny belongs to the postcrania informally called 'Malagasy
taxon B'. In addition, a partial mid caudal (UCB 92305) in the syntype series
of Titanosaurus madagascariensis "exhibits a distinctive, dorsoventrally
compressed morphology that characterizes caudal vertebrae of 'Malagasy Taxon
B'" according to Curry Rogers (2009). Since the other morphotype had already
been named Rapetosaurus krausei in 2001, 'Malagasy taxon B' should have
madagascariensis as its species name. Contra Curry Rogers, being chimarical
does not invalidate a species name, so the correct name for 'Malagasy taxon
B' should now be Vahiny madagascariensis with UCB 92305 as the lectotype.
Curry Rogers and Wilson propose Vahiny is sister to Jainosaurus,
which has yet to be placed precisely within Titanosauria.
References- Curry Rogers, 2009. The postcranial osteology of Rapetosaurus
krausei (Sauropoda: Titanosauria) from the Late Cretaceous of Madagascar.
Journal of Vertebrate Paleontology. 29(4), 1046-1086.
Curry Rogers and Wilson, 2014. Vahiny depereti, gen. et sp. nov., a new
titanosaur (Dinosauria, Sauropoda) from the Upper Cretaceous Maevarano Formation,
Madagascar. Journal of Vertebrate Paleontology. 34(3), 606-617.
"Yunxianosaurus" Li, 2001
"Y. hubeinensis" Li, 2001
Coniacian-Campanian, Late Cretaceous
Majiacun Group, Hubei, China
Holotype- (Museum of CUG coll.) (juvenile) skeleton including cervical vertebrae,
dorsal vertebrae, ribs, sacral vertebrae, caudal vertebrae, ilium, pubis and
tibia
....(SYF9) femur
....(SYH28) humerus
....(SYR25) radius
....(SYS7) scapula
....(SYU22) ulna
Comments- Li (2001) erected this taxon in a brief description consisting
largely of comparison to various sauropod families. In this it is noted as having
opisthocoelous dorsals and a humerofemoral ratio of 71%, lacking bifurcated
presacral neural spines and procoelous caudals. Li thus tentatively assigned
it to his typological Titanosauridae, but also stated the name is only intended
as temporary pending further excavation and research. This makes the name a
nomen nudum following ICZN Article 8.3. "Yunxiansaurus hubei" appears
in Zhou (2005) as a figure caption for a skeleton mounted at the CUG, but is
the same taxon. Li's paper would seem to indicate only a single individual is
present, with each element having its own SY number (not a museum abbreviation,
as in the same paper Bactrosaurus sp. elements are OB while "Lijiagousaurus"
elements are OL).
References- Li, 2001. Distribution, burying and classification of dinosaur
fossils in Upper Cretaceous strata at Meipu Town, Yunxian County of Hubei Province.
Hubei Geology & Mineral Resources. 15(4), 25-31.
Zhou, 2005. The Dinosaur Egg Fossils in Nanyang, China. China University of
Geosciences Press. 145 pp.
unnamed titanosaurian (Alifanov and Bolotsky, 2010a)
Late Maastrichtian, Late Cretaceous
Udurchukan Formation of the Tsagayan Group, Russia
Material- (AEHM 1/316; paratype of Arkharavia heterocoelica) incomplete
tooth
Comments- This was referred to Alifanov and Bolotsky's (2010a, b) new
supposed titanosauriform Arkharavia, but Mannion et al. (2013) correctly
noted it's from another locality and cannot be compared to the holotype vertebra.
They assigned it to Titanosauria based on its cylindrical shape.
References- Alifanov and Bolotsky, 2010a. Arkharavia heterocoelica
gen. et sp. nov., a new sauropod dinosaur from the Upper Cretaceous of far earstern
Russia. Paleontologicheskii Zhurnal. 2010(1), 76-83.
Alifanov and Bolotsky, 2010b. Arkharavia heterocoelica gen. et sp. nov.,
a new sauropod dinosaur from the Upper Cretaceous of the far East of Russia.
Paleontological Journal. 44(1), 84-91.
Mannion, Upchurch, Barnes and Mateus, 2013. Osteology of the Late Jurassic Portuguese
sauropod dinosaur Lusotitan atalaiensis (Macronaria) and the evolutionary
history of basal titanosauriforms. Zoological Journal of the Linnean Society.
168(1), 98-206.
Titanosauria indet. (Malkani, 2004)
Maastrichtian, Late Cretaceous
Vitakri Formation, Pakistan
Material- ?(GSP/MSM-56-1; intended paratype of "Vitakridrinda sulaimani") incomplete ?axis (Malkani, 2006)
(GSP/MSM-53-2; intended paratype of "Vitakridrinda sulaimani") incomplete caudal vertebra (Malkani, 2006)
(GSP/MSM-54-2; intended paratype of "Vitakridrinda sulaimani") incomplete caudal vertebra (Malkani, 2006)
(GSP/MSM-55-2; intended paratype of "Vitakridrinda
sulaimani") incomplete caudal vertebrae (87 mm) (Malkani, 2006)
?(GSP/MSM-57-3; intended paratype of "Vitakridrinda sulaimani") ?dorsal centrum (83
mm) (Malkani, 2006)
?(GSP/MSM-58-15; intended paratype of "Vitakridrinda sulaimani") proximal caudal centrum
(79 mm) (Malkani, 2006)
?(GSP/MSM-62-19; intended syntype of "Vitakridrinda sulaimani") partial ?braincase (Malkani, 2004)
Comments- Malkani (2004) first
mentions "thick basioccipital condyle articulated with posterior
braincase" as material of his supposed abelisaurid "Vitakridrinda"
(here referred to Coelurosauria), and it was later (Malkani, 2006)
misidentified as GSP/MSM-61-19 and listed as a holotype specimen of the
taxon. As noted in its entry here it would be better designated a
syntype given the uncertain association, being discovered 100 meters
away. Indeed, Malkani (2014) later says of "Vitakridrinda" "Its
braincase and basioccipital can be assigned to Maojandino alami (because of the yellow brown matrix covered on supposed braincase is same as on associated vertebrae and limb elements of Maojandino alami)
if character supports after specimen preparation." Malkani (2019
online, 2020a) later referred it to his sauropod
taxon "Gspsaurus pakistani" instead, which he synonymized "Maojandino"
with. The supposed anterior view is similar in rough
outline to Abelisaurus (assuming the supposed basipterygoid processes
are basal tubera instead) except for the decurved and much taller paroccipital
processes. The latter features match titanosaur braincases better, though in
that case everything beneath the occipital condyle area would be broken off.
In either case, it's only the shape that is similar, and since there are no
obvious surface features or natural edges, it could just as easily be part of
an ilium, vertebra, etc.. The posterior view does have the rough shape of an occipital
condyle, but no obvious foramen magnum above it. The possibility it could be
a non-fossil rock with a rounded protruding area should not be excluded.
Malkani (2006) intended to make the vertebrae paratypes of his new
abelisaur "Vitakridrinda", but they were referred to supposed
abelisauroid Vitakrisaurus
(here made Archosauria indet.) later in Malkani (2019, 2020b).
Their size and anteriorly placed neural arches (in the caudals) suggest
referral to Titanosauria. GSP/MSM-56-1 was originally called a
caudal vertebra (Malkani, 2006a)
and later an anterior dorsal (Malkani, 2020). The lack of a large
diapophysis suggests this may be an axis instead, tentatively assigned
to Titanosauria based on the broad, rounded and posteriorly sloped
neural spine. This would make Malkani's supposed posterior side
anterior, and note his labeled hyposphenes don't match the narrow
structure ventral to more horizontal postzygapophyseal surfaces seen in
abelisaurids. GSP/MSM-58-15 was identified as a caudal and later
(2020) as a proximal caudal, which seems correct. The size and
anteriorly placed neural arch resemble titanosaurs. Note the
supposed
transverse process is a broken posterodorsal corner while the
supposed
chevron facets are also broken surfaces.
References-
Malkani, 2004. Saurischian dinosaurs from Late Cretaceous of Pakistan. in Hussain
and Akbar (eds.). Abstract volume of Fifth Pakistan Geological Congress, Islamabad,
Pakistan. 71-73.
Malkani, 2006. Biodiversity of saurischian dinosaurs from the Latest Cretaceous
park of Pakistan. Journal of Applied and Emerging Sciences. 1(3), 108-140.
Malkani, 2009. New Balochisaurus (Balochisauridae,
Titanosauria, Sauropoda) and Vitakridrinda (Theropoda) remains from Pakistan.
Sindh University Research Journal (Science Series). 41(2), 65-92.
Malkani, 2014. Theropod dinosaurs and mesoeucrocodiles from the
terminal Cretaceous of Pakistan. The second International Symposium of
International Geoscience Programme (IGCP) Project 608, abstract volume.
169-172.
Malkani, 2019. Vitakrisaurus saraiki theropod from south Asia. Open Journal of Geology. 9, 643-645.
Malkani, 2019 online. Revision, discussion and diagnostic features of
valid titanosaurs (Sauropoda, Dinosauria) from Indo-Pakistan landmass.
[no longer uploaded] DOI: 10.13140/RG.2.2.25076.81287
Malkani, 2020a. First skull of medium sized titanosaur in Indo-Pakistan
subcontinent found from the Latest Maastrichtian Vitakri Formation of
Pakistan; Associated cranial and postcranial skeletons of Gspsaurus pakistani (Poripuchia, stocky Titanosauria, Sauropoda) from Pakistan and India. Open Journal of Geology. 9, 631-634.
Malkani, 2020b. Theropods, mesoeucrocodiles and pterosaurs found from
the latest Maastrichtian Vitakri Formation of Balochistan, Pakistan;
Description with large photographs and comparison with coeval taxa from
Indo-Pakistan subcontinent. Open Journal of Geology. 10, 510-551.
Titanosauria indet. (Huene and Matley, 1933)
Maastrichtian, Late Cretaceous
Lameta Formation, India
Material- (GSI K20/315) caudal vertebra (130 mm)
....(GSI K20/316) caudal vertebra (100 mm)
....(GSI K20/318) two dorsal chevron fragments
Comments- GSI K20/316 and K20/318 were referred to Titanosaurus indicus by Huene and Matley (1933), but referred to supposed abelisaurid Vitakrisaurus by Malkani (2019, 2020). Malkani may have decided this based on the amphicoelous centrum of GSI
K20/316 unlike the usual procoelous titanosaur caudals, but Huene and
Matley noted it was found with procoelous GSI K20/315 and "are,
therefore, almost certainly adjoining vertebrae", and indeed several
articulated examples of mixed coely in titanosaur tails have been
described since. The large size and anteriorly placed neural arch
further confirm titanosaur affinities.
References- Huene and Matley, 1933. The Cretaceous Saurischia and Ornithischia of the central
provinces of India. Palaeontologica Indica. 21, 1-74.
Malkani, 2019. Vitakrisaurus saraiki theropod from south Asia. Open Journal of Geology. 9, 643-645.
Malkani, 2020. Theropods, mesoeucrocodiles and pterosaurs found from
the latest Maastrichtian Vitakri Formation of Balochistan, Pakistan;
Description with large photographs and comparison with coeval taxa from
Indo-Pakistan subcontinent. Open Journal of Geology. 10, 510-551.
unnamed titanosaurian (De Carles, 1912)
Turonian-Maastrichtian, Late Cretaceous
Puerto Yerua Formation, Entre Rios, Argentina
Material- (MACN coll.) humerus (~1.12 m)
Comments- Originally reported by De Carles (1912) as being questionably
from the Asencio Formation. This specimen was referred to Argyrosaurus superbus
by Huene (1929), but Mannion and Otero (2012) noted it lacked the medial ridge
diagnostic for that taxon so referred it to Titanosauria indet.. Powell (2003)
similarly referred it to Titanosauridae indet.. Mannion and Otero's identification
was based on the medially projecting deltopectoral crest and square proximolateral
corner, only known to cooccur in titanosaurians/somphospondylans.
References- De Carles, 1912. Contribucion al estudio de las geodas ferruginosas
de Entre Rios, Corrientes, Misiones, etc., � prop�sito de varias,
coleccionadas por la Comisi�n Explorada � la laguna Iber�,
bajo los ausp�cios de la Sociedad Cientifica Argentina. Anales del Museo
Nacional de Historial Natural de Buenos Aires. 33, 411-415.
Huene, 1929. Los Saurisquios y Ornitisquios del Cretaceo Argentino. Anales del
Museo de La Plata. 3, 1-196.
Powell, 2003. Revision of South American titanosaurid dinosaurs: Palaeobiological,
palaeobiogeographical and phylogenetic aspects. Records of the Queen Victoria
Museum. 111, 173 pp.
Mannion and Otero, 2012. A reappraisal of the Late Cretaceous Argentinean sauropod
dinosaur Argyrosaurus superbus, with a description of a new titanosaur
genus. Journal of Vertebrate Paleontology. 32(3), 614-638.
unnamed probable Titanosauria (Huene, 1929)
Campanian, Late Cretaceous
Palacio Formation, Uruguay
Material- (MMAB coll.) rib fragment, distal humerus, partial radius
Comments- This was assigned to Argyrosaurus by Huene (1929) based
on its large size, but Powell's examination of the material could not justify
a more precise identity than Titanosauridae indet.. Mannion and Otero (2012)
did not study the material, so assigned it merely to Sauropoda indet.. Interestingly,
the material should be comparable to Argyrosaurus' holotype if reexamined
for the autapomorphies of that genus identified by the latter authors.
References- Huene, 1929. Terrestrische Oberkreide in Uruguay. Zentralblatt
fur Mineralogie, Geologie und Palaontologie Abteilung B. 1929, 107-112.
Powell, 2003. Revision of South American titanosaurid dinosaurs: Palaeobiological,
palaeobiogeographical and phylogenetic aspects. Records of the Queen Victoria
Museum. 111, 173 pp.
Mannion and Otero, 2012. A reappraisal of the Late Cretaceous Argentinean sauropod
dinosaur Argyrosaurus superbus, with a description of a new titanosaur
genus. Journal of Vertebrate Paleontology. 32(3), 614-638.
unnamed probable titanosaur (Gemmellaro, 1921)
Late Campanian, Late Cretaceous
Duwi Formation, Al Sharauna, Gebel Duwi and/or Gebel Nakheil, Egypt
Material- (MGUP MEGA004) pedal ungual (~42 mm)
Comments- Gemmellaro (1921) referred several remains to Megalosaurus crenatissimus
(Abelisauridae indet. here) from three localities in eastern
Egypt. He described two unguals and figured one, comparing the latter
favorably to crenatissimus
syntype FSL 92.290, but the preserved section of the figured specimen is
less tapered in side view with no obvious vascular groove. This
holds true as well for other Majungasaurus pedal unguals (e.g. FMNH PR 2434, MSNM V6418, V6419) as well as Masiakasaurus(e.g.
FMNH PR 2135) and indeed most theropods. Photos provided by Di
Patti (pers. comm. 6-2023) indicate this specimen (MGUP MEGA004) is
actually a sauropod pedal ungual and was illustrated poorly. This
matches the strong "asymmetry of the two sides" (translated) noted by
Gemmellaro and reveals the supposedly lacking proximal end is merely
poorly developed and anterodorsally angled as in e.g. Cedarosaurus' pedal ungual I and to a lesser extent Rapetosaurus.
Other similarities are the deep body with strongly curved dorsal and
ventral edges, vascular groove only present laterally (making this a
left ungual) and subparalell sides as in Cedarosaurus'
pedal ungual IV. The age would indicate titanosaurian
affinities. The second specimen is here provisionally considered
a hadrosauroid manual ungual I.
Reference- Gemmellaro, 1921. Rettili ma�strichtiani di Egitto. Giornale
di Scienze Naturali ed Economiche. 32, 339-351.
Andesauroidea Calvo and Bonaparte, 1991
vide Salgado, 2003
Definition- (Andesaurus delgadoi <- Saltasaurus loricatus)
(modified from Salgado, 2003)
= Laurasiformes Royo-Torres, 2009
Definition- (Tastavinsaurus sanzi <- Saltasaurus loricatus)
(Royo-Torres, 2009)
References- Calvo and Bonaparte, 1991. Andesaurus delgadoi gen.
et sp. nov. (Saurischia-Sauropoda), dinosaurio Titanosauridae de la Formacion
Rio Limay (Albiano-Cenomaniano), Neuqu�n, Argentina. Ameghiniana. 28(3-4),
303-310.
Salgado, 2003. Should we abandon the name Titanosauridae? Some comments on the
taxonomy of titanosaurian sauropods (Dinosauria). Revista Espa�ola de
Paleontolog�a. 18(1), 15-21.
Royo-Torres, 2009. El saur�podo de Pe�arroya de Tastavins. Monograpf�as
Turonlenses. 6, 548 pp.
Chubutisauridae Corro, 1974
?= Huanghetitanidae Lu, Xu, Zhang, Hu, Wu, Jia and Ji, 2007
References- Corro, 1974. Un nuevo sauropodo del Cretacico Chubutisaurus
insignis gen. et sp. nov. (Saurischia- Chubutisauridae nov.) del Cretacico
Superior (Chubutiano), Chubut, Argentina. Actas I Congreso Argentino de Paleontolog�a
y Bioestratigraf�a, Tucuman. 2, 229-240.
Lu, Xu, Zhang, Hu, Wu, Jia and Ji, 2007. A new gigantic sauropod dinosaur with
the deepest known body cavity from the Cretaceous of Asia. Acta Geologica Sinica.
81(2), 167-176.
Angolatitan Mateus, Jacobs,
Schulp, Polcyn, Tavares, Neto, Morais and Antunes, 2011
A. adamastor Mateus, Jacobs, Schulp, Polcyn, Tavares, Neto, Morais
and Antunes, 2011
Late Turonian, Late Cretaceous
Itombe Formation, Angola
Holotype- (MGUAN-PA-003) incomplete scapula (~1.05 m), humerus (1.l m),
radius (630 mm), ulna (690 mm), metacarpal I (300 mm), metacarpal III (309 mm),
metacarpal IV (285 mm)
Diagnosis- (after Mateus et al., 2011) posteroventral eminence of scapula
in proximal one quarter, anterior to acromial ridge; acute medioproximal margin
of humerus; rectangular lateral corner of proximal humerus; proximal outline
of ulna with anteromedial ridge (process); posterior facet of distal epiphysis
of metacarpal I with two small splint-like projections.
Comments- The holotype was discovered in 2005 and recovered as a non-lithostrotian
somphospondylan less derived than Euhelopus using a version of Wilson's
sauropod matrix joined with Upchurch et al.'s forelimb characters. D'Emic (2012)
stated it was a lithostrotian based on the anteriorly undivided radial condyle.
Mannion et al.'s (2013) analyses find Angolatitan to be a somphospondylan,
usually chubutisaurid.
References- Mateus, Jacobs, Schulp, Polcyn, Tavares, Neto, Morais and
Antunes, 2011. Angolatitan adamastor, a new sauropod dinosaur and the
first record from Angola. Anais da Academia Brasileira de Ci�ncias. 83(1),
1-13.
D'Emic, 2012. The early evolution of titanosauriform sauropod dinosaurs. Zoological
Journal of the Linnean Society. 166, 624-671.
Mannion, Upchurch, Barnes and Mateus, 2013. Osteology of the Late Jurassic Portuguese
sauropod dinosaur Lusotitan atalaiensis (Macronaria) and the evolutionary
history of basal titanosauriforms. Zoological Journal of the Linnean Society.
168(1), 98-206.
Chubutisaurus
Huanghetitan
Wintontitan
Andesauridae Calvo and Bonaparte, 1991 vide
Bonaparte and Coria, 1993
= Andesaurinae Calvo and Bonaparte, 1991
References- Calvo and Bonaparte, 1991. Andesaurus delgadoi gen.
et sp. nov. (Saurischia-Sauropoda), dinosaurio Titanosauridae de la Formacion
Rio Limay (Albiano-Cenomaniano), Neuqu�n, Argentina [Andesaurus delgadoi
gen. et sp. nov. (Saurischia-Sauropoda), a titanosaurid dinosaur from Rio Limay
Formation (Albian-Cenomanian), Neuqu�n, Argentina]. Ameghiniana. 28(3-4),
303-310.
Bonaparte and Coria, 1993. Un neuvo y gigantesco saur�podo titanosaurio
de la Formaci�n Rio Limay (Albanio-Cenomaniano) de la Provincia del Neuqu�n,
Argentina [A new and huge titanosaur sauropod from the Rio Limay Formation (Albian-Cenomanian)
of Neuqu�n Province, Argentina]. Ameghiniana. 30(3), 271-282.
Sauroposeidon Wedel, Cifelli
and Sanders, 2000a
= Paluxysaurus Rose, 2007
S. proteles Wedel, Cifelli and Sanders, 2000a}
= Paluxysaurus jonesi Rose, 2007
Aptian-Albian, Early Cretaceous
Antlers Formation, Oklahoma, US
Holotype- (OMNH 53062) fifth cervical vertebra with fused rib, sixth cervical
vertebra with fused rib (1.22 mm, rib 3.42 m), seventh cervical vertebra with
fused rib (1.23 mm, rib ~3.3 m), eighth cervical vertebra with fused rib (1.25
m, rib ~3.1 m)
Late Aptian-Early Albian, Early Cretaceous
Twin Mountains Formation of the Trinity Group, Texas, US
Referred- (FWMSH 93B-10-1; paratype of Paluxysaurus jonesi) (at
least four individuals) incomplete metacarpal III (Rose, 2007)
..(FWMSH 93B-10-2; paratype of Paluxysaurus jonesi) humerus (Rose, 2007)
..(FWMSH 93B-10-3; paratype of Paluxysaurus jonesi) femur (1.469 m) (Rose,
2007)
..(FWMSH 93B-10-4; paratype of Paluxysaurus jonesi) two chevrons (143,
157 mm) (Rose, 2007)
..(FWMSH 93B-10-5; paratype of Paluxysaurus jonesi) tooth (Rose, 2007)
..(FWMSH 93B-10-6; paratype of Paluxysaurus jonesi) tooth (Rose, 2007)
..(FWMSH 93B-10-7; paratype of Paluxysaurus jonesi) humeri (1.294, ~1.26
m), radius (789 mm), ulna (830 mm), femur (~1.52 m) (Rose, 2007)
..(FWMSH 93B-10-8; paratype of Paluxysaurus jonesi) (adult) incomplete
~fifth cervical vertebra and fused rib (~730 mm) (Rose, 2007)
..(FWMSH 93B-10-9; paratype of Paluxysaurus jonesi) partial dorsal vertebra
(Rose, 2007)
..(FWMSH 93B-10-10; paratype of Paluxysaurus jonesi) metacarpal IV (357
mm) (Rose, 2007)
..(FWMSH 93B-10-11; paratype of Paluxysaurus jonesi) first dorsal vertebra
(217 mm) (Rose, 2007)
..(FWMSH 93B-10-12; paratype of Paluxysaurus jonesi) two mid caudal vertebrae
(98, 103 mm) (Rose, 2007)
..(FWMSH 93B-10-13; paratype of Paluxysaurus jonesi) ?third and ?fourth
dorsal vertebrae (270 mm), dorsal rib fragment (Rose, 2007)
..(FWMSH 93B-10-14; paratype of Paluxysaurus jonesi) two proximal caudal
vertebrae (97, 106 mm) (Rose, 2007)
..(FWMSH 93B-10-15; paratype of Paluxysaurus jonesi) tibia (887 mm),
fibula (911 mm) (Rose, 2007)
..(FWMSH 93B-10-16; paratype of Paluxysaurus jonesi) metatarsal I (or
V?) (172 mm), metatarsal II (190 mm) (Rose, 2007)
..(FWMSH 93B-10-17; paratype of Paluxysaurus jonesi) eight distal caudal
vertebrae (100, 102, 89, 88, 86, 90 mm) (Rose, 2007)
..(FWMSH 93B-10-18; holotype of Paluxysaurus jonesi) maxilla, nasal (194
mm) (Rose, 2007)
..(FWMSH 93B-10-19; paratype of Paluxysaurus jonesi) cervical vertebra
and fused rib fragment (Rose, 2007)
..(FWMSH 93B-10-20; paratype of Paluxysaurus jonesi) dorsal rib fragment
(Rose, 2007)
..(FWMSH 93B-10-21; paratype of Paluxysaurus jonesi) five mid caudal
vertebrae (137, 131, 128 mm) (Rose, 2007)
..(FWMSH 93B-10-22; paratype of Paluxysaurus jonesi) metacarpal II (311
mm) (Rose, 2007)
..(FWMSH 93B-10-23; paratype of Paluxysaurus jonesi) dorsal rib fragment
(Rose, 2007)
..(FWMSH 93B-10-24; paratype of Paluxysaurus jonesi) incomplete scapula
(1.432 m), sternal plate (583 mm) (Rose, 2007)
..(FWMSH 93B-10-25; paratype of Paluxysaurus jonesi) femur (1.442 m),
fibula (Rose, 2007)
..(FWMSH 93B-10-26; paratype of Paluxysaurus jonesi) metatarsal IV (195
mm) (Rose, 2007)
..(FWMSH 93B-10-27; paratype of Paluxysaurus jonesi) last five dorsal
vertebrae (204, 195, ~260, ~280, ~220 mm), sacrum (?, ?, ?, 210, 139 mm), first
caudal vertebra (72 mm), second caudal vertebra (98 mm), third caudal vertebra,
fourth caudal vertebra (~109 mm), fifth caudal vertebra (~119 mm), sixth caudal
vertebra, eighth caudal vertebra (~120 mm), ninth caudal vertebra (115 mm),
tenth caudal vertebra, eleventh caudal vertebra, twelfth caudal vertebra, thirteenth
caudal vertebra, ilia, pubis, ischium, femur (Rose, 2007)
..(FWMSH 93B-10-28-32; paratypes of Paluxysaurus jonesi) (adult) third-ninth
cervical vertebrae (446, 661, ?, 665, ?, ~740, ~830 mm), cervical rib fragments
(Rose, 2007)
..(FWMSH 93B-10-33; paratype of Paluxysaurus jonesi) tooth (Rose, 2007)
..(FWMSH 93B-10-34; paratype of Paluxysaurus jonesi) coracoid (357 mm)
(Rose, 2007)
..(FWMSH 93B-10-35; paratype of Paluxysaurus jonesi) pubes (966 mm),
ischium (Rose, 2007)
..(FWMSH 93B-10-36; paratype of Paluxysaurus jonesi) radius (~790 mm),
metacarpal I (321 mm), partial metacarpal II, metacarpal III (375 mm), partial
metacarpal IV (Rose, 2007)
..(FWMSH 93B-10-37; paratype of Paluxysaurus jonesi) distal caudal vertebra
(114 mm) (Rose, 2007)
..(FWMSH 93B-10-38; paratype of Paluxysaurus jonesi) distal caudal vertebra
(83 mm) (Rose, 2007)
..(FWMSH 93B-10-39; paratype of Paluxysaurus jonesi) coracoid (309 mm)
(Rose, 2007)
..(FWMSH 93B-10-40; paratype of Paluxysaurus jonesi) tooth (Rose, 2007)
..(FWMSH 93B-10-41; paratype of Paluxysaurus jonesi) dsital caudal vertebra
(109 mm) (Rose, 2007)
..(FWMSH 93B-10-42; paratype of Paluxysaurus jonesi) distal caudal vertebra
(71 mm) (Rose, 2007)
..(FWMSH 93B-10-43; paratype of Paluxysaurus jonesi) distal caudal vertebra
(103 mm) (Rose, 2007)
..(FWMSH 93B-10-44; paratype of Paluxysaurus jonesi) distal caudal vertebra
(107 mm) (Rose, 2007)
..(FWMSH 93B-10-45; paratype of Paluxysaurus jonesi) tibia (889 mm) (Rose,
2007)
..(FWMSH 93B-10-46; paratype of Paluxysaurus jonesi) tibia (880 mm) (Rose,
2007)
..(FWMSH 93B-10-47; paratype of Paluxysaurus jonesi) metacarpal(s) (Rose,
2007)
..(FWMSH 93B-10-48; paratype of Paluxysaurus jonesi) dorsal vertebra
(Rose, 2007)
..(FWMSH 93B-10-49; paratype of Paluxysaurus jonesi) tooth (Rose, 2007)
..(FWMSH 93B-10-50; paratype of Paluxysaurus jonesi) tooth (Rose, 2007)
..(FWMSH 93B-10-51; paratype of Paluxysaurus jonesi) pubis, ischium (Rose,
2007)
..(TMM 42488; paratype of Paluxysaurus jonesi) three dorsal vertebrae
(196 mm), dorsal ribs, more than ten caudal vertebrae, two chevrons (293 mm),
incomplete scapulocoracoid (cor 329 mm), humerus (1.2 m), pubis, two ischia,
two femora, tibia (823 mm) (Rose, 2007)
.. several scapulae or scapulocoracoids, ischium, four femora (one lost?) (Rose,
1007)
Late Albian, Early Cretaceous
Cloverly Formation, Wyoming, US
(YPM 5147) incomplete posterior dorsal vertebra (290 mm), two fused incomplete
proximal caudal vertebrae (99, 99 mm) (Ostrom, 1970)
?...(YPM 5087) chevron (D'Emic and Foreman, 2012)
?...(YPM 5089) chevron (D'Emic and Foreman, 2012)
?...(YPM 5096) chevron (D'Emic and Foreman, 2012)
?...(YPM 5097) chevron (D'Emic and Foreman, 2012)
?...(YPM 5100) dorsal rib (D'Emic and Foreman, 2012)
?...(YPM 5103) mid-distal caudal centrum (101 mm) (Ostrom, 1970)
?...(YPM 5104) incomplete mid-distal caudal vertebra (101 mm) (Ostrom, 1970)
?...(YPM 5450) tibia (985 mm) (Ostrom, 1970)
(YPM 5294) (juvenile) cervical centrum (485 mm) (Ostrom, 1970)
?...(UM 20800) incomplete scapula, incomplete coracoid (200 mm) (D'Emic and
Foreman, 2012)
?...(YPM 5123) chevron (D'Emic and Foreman, 2012)
?...(YPM 5126) dorsal rib fragments (D'Emic and Foreman, 2012)
?...(YPM 5152) distal caudal centrum (102 mm) (Ostrom, 1970)
?...(YPM 5451) femur (~1.46 m) (Ostrom, 1970)
....(YPM 5449) incomplete ?second dorsal vertebra (190 mm), incomplete ?third
dorsal vertebra, incomplete ?fourth dorsal vertebra, three incomplete posterior
dorsal vertebrae (190, 202 mm), dorsal rib fragments, sternal plates (390 mm),
distal radius, ulna (600 mm), incomplete pubes (Ostrom, 1970)
Diagnosis- (after Wedel et al., 2000a) cervical diapophysis located approximately
one third of centrum length behind anterior condyle; deeply excavated cervical
neural spines which are perforate in anterior cervicals; hypertrophied central
pneumatic fossae that extend posteriorly to cotyles; cervical neural spines
occupy anterior nine-tenths of centra and are not bifurcate; cervical ribs slender
and elongated, with long, robust anterior processes that extend nearly to anterior
condyles; total length of each cervical rib equals or exceeds 3 centrum lengths
(also in Euhelopus).
(after Rose, 2007) broad nasal process of maxilla; strong lateral curvature
of premaxillary process of nasal; differs from Giraffititan in having
shorter, non-arching premaxillary nasal process and more pronounced lacrimal
process of maxilla; differs from Pleurocoelus in shape of posteroventral
margin of maxilla, shape of distal scapular blade, and shape of proximal tibial
condyle; differs from Euhelopus zdanskyi in nasal process of maxilla
rises from the middle of bone; distinguished from Giraffatitan by distinct
intrapostzygapophyseal lamina on cervical neural arches; deep postspinal fossa
on posterior surface of cervical neural arches; accessory laminae on dorsal
neural arches; dorsal vertebrae that lack postspinal lamina; mid caudal centra
articular faces angle anteriorly; proximal and mid- caudal neural arches with
intraprezygapophyseal laminae that form, with spinoprezygapophyseal laminae,
a prespinal fossa above neural canal; transversely expanded proximal and mid
caudal neural spines; distal scapular blade broadly expanded on both acromial
and glenoid sides; banteroposteriorly compressed femoral shaft.
(after D'Emic and Foreman, 2012) mid cervical vertebral elongation index greater
than six; mid cervical pneumatic fossa developed posteriorly to cotyle; mid
and posterior dorsal neural spines taper transversely distally; proximal caudal
vertebral centra squared off in anteroposterior views; mid caudal vertebrae
with wide spinoprezygapophyseal fossa and spinoprezygapophyseal laminae that
meet at angle greater than 50 degrees; scapula with two processes at base of
blade on ventral side; humerus gracile (length/midshaft width >7.5).
Other diagnoses- D'Emic (2013) suggested the "cervical centra extremely
elongate (length more than five times posterior centrum height)" was also
found in Euhelopus and Erketu, the camellate internal structure
was also found in e.g. Euhelopus and Chubutisaurus, the "well
defined centroparapophyseal laminae that extend to posterior ends of centra"
was also found in Giraffatitan and Euhelopus.
Comments- Ostrom (1970) described several ?titanosaurid specimens from
the Cloverly Formation, including cervical YPM 5294 that Wedel et al. (2000a,
b) referred to Sauroposeidon. D'Emic and Foreman (2012) referred most
of these to Sauroposeidon, reinterpreting the supposed ischia as pubes
and identifying new elements (e.g. radius, ribs, chevrons).
The holotype was discovered in 1994. It was described in Wedel's (1997) thesis,
then published and named in Wedel et al. (2000). Sauroposeidon was hypothesized
to be a brachiosaurid.
Paluxysaurus' type material was discovered in 1985 and described as a
new taxon by Rose (2007). Entered into a version of Wilson's sauropod analysis,
it emerged as a brachiosaurid.
D'Emic and Foreman (2012) synonymized these taxa.
References- Ostrom, 1970. Stratigraphy and paleontology of the Cloverly
Formation (Lower Cretaceous) of the Bighorn Basin area, Wyoming and Montana.
Bulletin of the Peabody Museum of Natural History. 35, 1-234.
Wedel, 1997. A new sauropod dinosaur from the Early Cretaceous of Oklahoma.
BS Thesis, University of Oklahoma. 31 pp.
Wedel, Cifelli and Sanders, 2000a. Sauroposeidon proteles, a new sauropod
from the Early Cretaceous of Oklahoma. Journal of Vertebrate Paleontolog. 20(1),
109-114.
Wedel, Cifelli and Sanders, 2000b. Osteology, paleobiology, and relationships
of the sauropod dinosaur Sauroposeidon. Acta Palaeontologica Polonica.
45(4), 343-388.
Wedel and Cifelli, 2005. Sauroposeidon: Oklahoma's native giant. Oklahoma
Geology Notes 65, 40-57.
Rose, 2007. A new titanosauriform sauropod (Dinosauria: Saurischia) from the
Early Cretaceous of central Texas and its phylogenetic relationships. Palaeontologia
Electronica. 10(2), 8A:65p.
D'Emic and Foreman, 2012. The beginning of the sauropod dinosaur hiatus in North
America: Insights from the Lower Cretaceous Cloverly Formation of Wyoming. Journal
of Vertebrate Paleontology. 32(4), 883-902.
D'Emic, 2013. Revision of the sauropod dinosaurs of the Lower Cretaceous Trinity
Group, southern USA, with the description of a new genus. Journal of Systematic
Palaeontology. 11(6), 707-726.
Winkler, Polcyn and Jacobs, 2013. New sauropod dinosaur material from Jones
Ranch: A large Comanchean nonmammalian tetrapod from Texas. Earth and Environmental
Science Transactions of the Royal Society of Edinburgh. 103(3-4), 459-469.
Tastavinsaurus
Andesaurus
Ligabuesaurus
Titanosauroidea Lydekker, 1885 vide Upchurch,
1995
Definition- (Titanosaurus indicus <- Andesaurus delgadoi)
(suggested)
Other definitions- (Titanosaurus indicus <- Brachiosaurus
altithorax) (modified from Upchurch, 1998)
(Saltasaurus loricatus <- Andesaurus delgadoi) (modified from
Salgado, 2003)
Comments- I modify this definition to be equivalent to Salgado's, but
using the eponymous genus as the internal specifier.
References- Lydekker, 1885. The Reptilia & Amphibia of the Maleria
& Denwa Groups. Memoirs of the Geological Survey of India. Palaeontologia
Indica, Series IV. Indian Pretertiary Vertebrata. 1(5), 1-38.
Upchurch, 1995. The evolutionary history of sauropod dinosaurs. Philosophical
Transactions of the Royal Society of London B. 349, 365-390.
Upchurch, 1998. The phylogenetic relationships of sauropod dinosaurs. Zoological
Journal of the Linnean Society. 124, 43-103.
Salgado, 2003. Should we abandon the name Titanosauridae? Some comments on the
taxonomy of titanosaurian sauropods (Dinosauria). Revista Espa�ola de
Paleontolog�a. 18(1), 15-21.
Tambatitanis Saegusa and
Ikeda, 2014
T. amicitiae Saegusa and Ikeda, 2014
Early Albian, Early Cretaceous
Lower Formation of the Sasayama Group, Japan
Holotype- (MNHAH D-1029280; Tamba sauropod) braincase, partial dentary,
eleven teeth, partial atlas, fragmentary cervical vertebra, cervical rib fragments,
two dorsal centra, presacral vertebral fragments, twelve incomplete dorsal ribs,
five fused sacral neural spines, first sacral ribs, second sacral rib, third
sacral rib, fourth sacral rib, fifth sacral rib, partial first caudal vertebra,
incomplete second caudal vertebra (100 mm), incomplete third caudal vertebra
(101 mm), incomplete fourth caudal vertebra (112 mm), incomplete fifth caudal
vertebra (89 mm), incomplete sixth caudal vertebra (99 mm), incomplete seventh
caudal vertebra (91 mm), incomplete eighth caudal vertebra (110 mm), ninth caudal
vertebra (100 mm), tenth caudal vertebra (94 mm), eleventh caudal vertebra (109
mm), eleven distal caudal vertebrae (~120, 112, 112, 113, 105, 101, 103, 89,
84, 77, 75 mm), seventeen chevrons (102->410 mm), incomplete ilium, pubis
Diagnosis- (after Saegusa and Ikeda, 2014) dorsal border of paroccipital
process shaft that forms ventral margin of posttemporal fenestra short mediolaterally
and V-shaped in posterior view; postzygapophysis and apex of proximal caudal
neural spine located posterior to centrum; proximal caudal neural spine curved
strongly anteriorly and bow-shaped in lateral view; proximal caudal neural spine
expanded and hemispherical with anterior face excavated by posterodorsal extension
of deep and narrow spinoprezygapophyseal fossa; proximal caudal transverse process
short and L-shaped; proximal chevrons longest among sauropods in proportion
to centrum height (over twice); ventral end of proximal chevrons rod-shaped;
ventral ends of mid chevrons transversely thin and long anteroposteriorly, without
anterior process.
Comments- The holotype was discovered in 2006, the teeth described in
Saegusa et al. (2010), and most of the material described when the taxon was
named in Saegusa and Ikeda (2014). Some material (cervical vertebra, presacral
fragments, pubis) remained unprepared and undescribed. Saegusa and Ikeda recovered
Tambatitanis as a euhelopodid somphospondylan, which in their topology
is outside Titanosauria and includes Qiaowanlong, Erketu, Daxiatitan,
Tangvayosaurus and Phuwiangosaurus as well. These are a grade
of non-lithostrotian titanosauroids in the topology used here, so Tambatitanis
is assumed to be in this region of the tree as well.
References- Saegusa, Tanaka and Ikeda, 2010. Preliminary observations
on the dinosaur teeth from the Lower Cretaceous Sasayama Group in Tamba City,
Hyogo Prefecture and additional notes on the pneumaticity of the postcranial
skeleton of Tamba sauropod. Journal of Fossil Research. 42, 52-65.
Saegusa and Tomida, 2011. Titanosauriform teeth from the Cretaceous of Japan.
Anais da Academia Brasileira de Ci�ncias. 83, 247-265.
Saegusa and Ikeda, 2014. A new titanosauriform sauropod (Dinosauria: Saurischia)
from the Lower Cretaceous of Hyogo, Japan. Zootaxa. 3848(1), 1-66.
Astrophocaudia D'Emic,
2012
A. slaughteri D'Emic, 2012
Early Albian, Early Cretaceous
Paluxy Formation of the Trinity Group, Texas, US
Holotype- (SMU 203/73655) tooth (~15x5.6x6.4 mm)
....(SMU 61732) incomplete mid-posterior cervical vertebra (480 mm), several
fragmentary posterior cervical or anterior dorsal vertebrae, ~twenty fragmentary
dorsal ribs, incomplete ~eighth caudal vertebra (72 mm), partial ~tenth caudal
vertebra, partial ~eleventh caudal vertebra, ~twelfth caudal vertebra (97 mm),
~thirteenth caudal vertebra (91 mm), ~fourteenth caudal vertebra (87 mm), ~fifteenth
caudal vertebra (74 mm), ~nineteenth caudal vertebra (88 mm), ~twentieth caudal
vertebra (91 mm), ~twenty-second caudal vertebra (86 mm), ~twenty-third caudal
vertebra (87 mm), ~twenty-fourth caudal vertebra (80 mm), incomplete ~twenty-fifth
caudal vertebra (73 mm), incomplete ~twenty-sixth caudal vertebra (72 mm), ~twenty-seventh
caudal vertebra (81 mm), incomplete ~thirtieth caudal vertebra (78 mm), ~thirty-first
caudal vertebra (72 mm), ~thirty-third caudal vertebra (68 mm), ~thirty-seventh
caudal vertebra (69 mm), ~thirty-eighth caudal vertebra (65 mm), ~thirty-ninth
caudal vertebra (57 mm), incomplete ~forty-third caudal vertebra (~51 mm), ~forty-sixth
caudal vertebra (45 mm), partial ~forty-eighth caudal vertebra ( mm), caudal
vertebral fragments, two incomplete proximal chevrons, partial scapula, partial
ilium
Diagnosis- (after D'Emic, 2013) proximal-mid caudal vertebrae with planar
hyposphene-hypantrum articulations set off from zygapophyses; proximal-mid caudal
vertebrae with prespinal lamina contacting intraprezygapophyseal lamina.
Comments- D'Emic's description naming the taxon was published electronically
in 2012 with a ZooBank registration, but not physically until 2013. The two
teeth mentioned by Rose (2007) are mid caudal prezygapophyses (D'Emic, 2013).
This sauropod was first mentioned by Bilelo (1969) and was discovered between
1964 and 1967. Langston (1974) initially described and referred this specimen
to Pleurocoelus sp., but both Salgado et al. (1995) and Tidwell et al.
(1999) disagreed and placed it closer to titanosaurs. D'Emic (2013) fully described
and named the specimen, finding it to be a somphospondylan more closely related
to titanosaurs than Ligabuesaurus. D'Emic (2012) previously found the
specimen to be a somphospondylan as derived as Sauroposeidon but outside
Titanosauria. Mannion et al. (2013) recovered it as a titanosauriform outside
Sauroposeidon+ or as a basal titanosauroid sister to Brontomerus,
depending on assumptions used.
References- Bilelo, 1969. The fossil fish Lepidotes in the Paluxy
Formation, north-central Texas. American Midland Naturalist. 81, 405-411.
Langston, 1974. Nonmammalian Comanchean tetrapods. Geoscience and Man. 8, 77-102.
Salgado, Calvo and Coria, 1995. Relaciones filogen�ticas de Pleurocoelus
Marsh (Sauropoda). Jornadas Argentinas de Paleontolog�a de Vertebrados
11. 34.
Tidwell, Carpenter and Brooks, 1999. New sauropod from the Lower Cretaceous
of Utah, USA. Oryctos. 2, 21-37.
Rose, 2007. A new titanosauriform sauropod (Dinosauria: Saurischia) from the
Early Cretaceous of central Texas and its phylogenetic relationships. Palaeontologia
Electronica. 10(2), 8A:65p.
D'Emic, 2012. The early evolution of titanosauriform sauropod dinosaurs. Zoological
Journal of the Linnean Society. 166, 624-671.
D'Emic, 2013. Revision of the sauropod dinosaurs of the Lower Cretaceous Trinity
Group, southern USA, with the description of a new genus. Journal of Systematic
Palaeontology. 11(6), 707-726.
Mannion, Upchurch, Barnes and Mateus, 2013. Osteology of the Late Jurassic Portuguese
sauropod dinosaur Lusotitan atalaiensis (Macronaria) and the evolutionary
history of basal titanosauriforms. Zoological Journal of the Linnean Society.
168(1), 98-206.
Brontomerus Taylor, Wedel
and Cifelli, 2011
B. mcintoshi Taylkor, Wedel and Cifelli, 2011
Aptian-Albian, Early Cretaceous
Ruby Ranch Member of Cedar Mountain Formation, Utah, US
Holotype- (OMNH 66430) incomplete ilium (405 mm)
Referred- ?...(OMNH 61248) incomplete mid caudal vertebra (110 mm) (Taylor
et al., 2011)
?...(OMNH 66429) presacral centrum (140 mm) (Taylor et al., 2011)
?...(OMNH 66431) sternal plate (150 mm) (Taylor et al., 2011)
?...(OMNH 66432) partial sternal (Taylor et al., 2011)
?(OMNH 27761) incomplete scapula (~1.21 m) (Taylor et al., 2011)
?...(OMNH 27762) dorsal rib shaft (Taylor et al., 2011)
?...(OMNH 27763) dorsal rib fragment (Taylor et al., 2011)
?...(OMNH 27764) dorsal rib fragment (Taylor et al., 2011)
?...(OMNH 27765) dorsal rib fragment (Taylor et al., 2011)
?...(OMNH 27766) first sacral rib (760 mm) (Taylor et al., 2011)
?...(OMNH 27767) dorsal rib fragment (Taylor et al., 2011)
?...(OMNH 27768) dorsal rib fragment (Taylor et al., 2011)
?...(OMNH 27769) ?ischial fragment (Taylor et al., 2011)
?...(OMNH 27770) dorsal rib fragment (Taylor et al., 2011)
?...(OMNH 27771) dorsal rib fragment (Taylor et al., 2011)
?...(OMNH 27772) dorsal rib fragment (Taylor et al., 2011)
?...(OMNH 27773) dorsal rib fragment, two flat bone fragments (Taylor et al.,
2011)
?...(OMNH 27774-27783) fragments (Taylor et al., 2011)
?...(OMNH 27784) twenty-one fragments (Taylor et al., 2011)
?...(OMNH 27785-27793) fragments (Taylor et al., 2011)
?...(OMNH 27794) partial distal caudal centrum (Taylor et al., 2011)
?...(OMNH 27795-27800) fragments (Taylor et al., 2011)
Diagnosis- (after Taylor et al., 2011) presacral vertebrae camellate;
mid to distal caudal vertebrae with elongate pre- and postzygapophyses, having
the postzygapophyseal facets hanging below level of ramus; acromion expansion
pronounced and steep, but not forming acromial fossa; dorsal and ventral margins
of scapular blade 'stepped'; sternal plates crescentic and three times as long
as broad; preacetabular process directed anterolaterally at 30 degrees relative
to the sagittal plane, but straight in dorsal view and vertically oriented.
Other diagnoses- D'Emic (2012) correctly noted that since the postacetabular
process is not preserved, several characters listed by Taylor et al. cannot
be evaluated- preacetabular process 55% of ilial length; postacetabular process
reduced to near absence; ilium proportionally tall - height is 52% of total
length. D'Emic claims the other two ilial characters are "present in a
variety of taxa (e.g. Tastavinsaurus, Royo-Torres, 2009; Giraffatitan,
Janensch, 1961)", however both have a more laterally angled preacetabular
process (~40 degrees) and Giraffatitan's is twisted to be diagonally
oriented anteriorly. The ischial peduncle is larger than interpreted by Taylor
et al. though, as they only treat the ventralmost laterally bulging portion
as this structure, while the portion ventral to any posterior flaring of the
postacetabular process is also part of the peduncle. Thus their character "ischiadic
peduncle reduced to very low bulge" is not correct.
Taylor et al. also listed "first dorsal rib with expanded, dorsally oriented
articular facts, laterally curving shaft, and distally directed pneumatic foramen
in head", but Saegusa and Ikeda (2014) reidentified this element as a first
sacral rib based on similarity to that element in Tambatitanis.
Comments- Discovered in 1994-1995. Taylor et al.'s analysis using Harris'
sauropod matrix found it to be a camarasauromorph outside Brachiosauridae and
Saltasaurinae. D'Emic (2012) claimed Brontomerus was an indeterminate
potential chimaera including titanosauriform vertebrae and ribs, as the type
quarry contains individuals of different sizes and no overlap with the holotype
ilium. However, it is common for sauropodomorphs to be based on disarticulated
elements of multiple individuals from one quarry and the elements are all congruent
with a basal macronarian phylogenetic position. Thus, Taylor et al.'s referrals
are provisionally accepted here. As noted above, one of the ilial autapomorphies
used by Taylor et al. survives comparison to at least Tastavinsaurus
and Giraffatitan. Mannion et al. (2013) recovered Brontomerus
as a somphospondylan sister to laurasiforms plus more derived taxa, or sister
to Astrophocaudia as a basal titanosauroid, depending on the assumptions
made.
References- Kirkland, Britt, Burge, Carpenter, Cifelli, DeCourten, Eaton,
Hasiotis and Lawton, 1997. Lower to middle Cretaceous dinosaur faunas of the
Central Colorado Plateau: A key to understanding 35 million years of tectonics,
sedimentology, evolution, and biogeography. Brigham Young University Geology
Studies. 42, 69-103.
Taylor, Wedel and Cifelli, 2011. Brontomerus mcintoshi, a new sauropod
dinosaur from the Lower Cretaceous Cedar Mountain Formation, Utah, USA. Acta
Palaeontologica Polonica. 56(1), 75-98.
D'Emic, 2012. The early evolution of titanosauriform sauropod dinosaurs. Zoological
Journal of the Linnean Society. 166, 624-671.
Mannion, Upchurch, Barnes and Mateus, 2013. Osteology of the Late Jurassic Portuguese
sauropod dinosaur Lusotitan atalaiensis (Macronaria) and the evolutionary
history of basal titanosauriforms. Zoological Journal of the Linnean Society.
168(1), 98-206.
Saegusa and Ikeda, 2014. A new titanosauriform sauropod (Dinosauria: Saurischia)
from the Lower Cretaceous of Hyogo, Japan. Zootaxa. 3848(1), 1-66.
Australodocus
Euhelopodidae Romer, 1956 vide Kuhn, 1965
= Helopodidae Wiman, 1929
Definition- (Euhelopus zdanskyi <- Neuquensaurus australis)
(D'Emic, 2012)
References- Wiman, 1929. Die Kreide-Dinosaurier aus Shantung [The Cretaceous
dinosaurs of Shantung]. Palaeontologia Sinica, Series C. 6(1), 1-67.
Romer, 1956. Osteology of the Reptiles. University of Chicago Press. 772 pp.
Kuhn, 1965. Saurischia. Fossilium Catalogus, I: Animalia. 109, 94 pp.
D'Emic, 2012. The early evolution of titanosauriform sauropod dinosaurs. Zoological
Journal of the Linnean Society. 166, 624-671.
Euhelopodinae Romer, 1956
= Helopodinae Wiman, 1929 vide Huene, 1932
References- Wiman, 1929. Die Kreide-Dinosaurier aus Shantung [The Cretaceous
dinosaurs of Shantung]. Palaeontologia Sinica, Series C. 6(1), 1-67.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte
[The fossil reptile order Saurischia, their development and history]. Monographien
zur Geologie und Palaeontologie, serie 1. 4(1-2), 361 pp.
Romer, 1956. Osteology of the Reptiles. University of Chicago Press. 772 pp.
Euhelopus
Gannansaurus Lu, Yi, Zhong
and Wei, 2013
G. sinensis Lu, Yi, Zhong and Wei, 2013
Campanian-Maastrichtian, Late Cretaceous
Nanxiong Formation, Jiangxi, China
Holotype- (GMNH F10001-1) incomplete posterior dorsal vertebra (330 mm inc.
condyle)
....(GMNH F10001-2) incomplete mid caudal vertebra (135 mm)
Diagnosis- (after Lu et al., 2013) two centroprezygapophyseal lamina
fossae, the neural canal and distinct openings separately located at the bottom
and upper portion of large, square-shaped concavity, which are formed by centroprezygapophyseal
laminae and dorsal margin of centrum; large infraparapophyseal fossa; lateral
concavity with eye-shaped pleurocoel large and round, 65% of central length;
anterior and posterior centroparapophyseal laminae poorly developed; posterior
centrodiapophyseal lamina separated into two branches by infradiapophyseal fossa
dorsally; parapophyseal and diapophyseal laminae cross to form 'K' configuration
(also in Euhelopus); wide and shallow ventral concavity on caudal vertebra.
Comments- More of the specimen was originally present, but accidentally
destroyed during a construction project. Lu et al. (2013) assigned Gannansaurus
to Somphospondyli and proposed a relationship with Euhelopus based on
the K-shaped lamina configuration.
Reference- Lu, Yi, Zhong and Wei, 2013. A new somphospondylan sauropod
(Dinosauria, Titanosauriformes) from the Late Cretaceous of Ganzhou, Jiangxi
Province of southern China. Acta Geologica Sinica (English Edition). 87(3),
678-685.
Fukuititan Azuma and Shibata,
2010
?= "Sugiyamasaurus" Lambert, 1990
F. nipponensis Azuma and Shibata, 2010
Barremian, Early Cretaceous
Kitadani Formation of the Akaiwa Subgroup of the Tetori Group, Japan
Holotype- (FPDM V8468) three teeth, incomplete cervical neural arch, three
distal caudal vertebrae (30 mm), proximal scapula, incomplete humeri (~910 mm),
incomplete radii (~600 mm), metacarpal IV (295 mm), incomplete ischia, partial
femur (~1.14 m), partial tibia, fibula, pedal phalanx (130 mm), fragments
Referred- ?(material of "Sugiyamasaurus"; Sugiyama-ryu) three
teeth (Azuma, 1991)
? five teeth (Azuma, 1991)
?(FPDM 1080417) tooth (Goto et al., 2002)
?(FPDM 1080417-A) tooth (Goto et al., 2002)
?(FPDM 1080757) tooth (Goto et al., 2002)
?(FPDM 1080920) tooth (Goto et al., 2002)
?(FPDM 1080942) tooth (Goto et al., 2002)
?(FPDM 1080944) tooth (Goto et al., 2002)
Diagnosis- (after Azuma and Shibata, 2010) tooth crowns asymmetrical
and elongated with weak or absent labial groove and no lingual concavity; stalk-like
cervical epipophyses; humerus proximal width 32% of length; metacarpal IV 48%
of radial length; ischia slightly expanded distally.
Comments- The holotype was discovered in 2007 and described as a basal
titanosauriform. D'Emic (2012) assigned it to Macronaria based on its long metacarpus.
Mannion et al. (2013) recovered it either as a titanosauriform outside the Sauroposeidon+
clade or as a basal titanosauroid.
Teeth were first discovered from the quarry in 1989. Three teeth were informally
called "Sugiyama-ryu", as found in Azuma (1991) and referred to Camarasauridae
by this author and Dong et al. (1990). Lambert (1990) inappropriately made it
into a genus name, listing it as "Sugiyamasaurus" in a childrens'
book. Azuma (1991) lists five additional sauropod teeth as family indet. A and
B, but he later (2003) placed all the teeth in Brachiosauridae. Kobayashi et
al. (2006) referred the specimens to Titanosauriformes indet. based on near
identical morphology to teeth from the Kuwajima Formation. Goto et al. (2002)
illustrate a tooth. As the "Sugiyamasaurus" teeth are generally similar
to Fukuititan's in morphology (D-shaped section; wrinkled enamel; parallel
mesial and distal edges- Kobayashi et al., 2006) and found in the same quarry,
they may belong to the same taxon. They should be examined for Fukuititan's
supposed dental autapomorphies, which should be checked for positional variation
and compared to other taxa as well.
References- Dong, Hasegawa and Azuma, 1990. The Age of Dinosaurs in Japan
and China. Fukui, Japan: Fukui Prefectural Museum. 65 pp.
Lambert, 1990. The Dinosaur Data Book. New York: Avon Books, 66. ISBN 0-380-75896-3.
Azuma, 1991. Early Cretaceous Dinosaur Fauna from the Tetori Group, central
Japan. Research on Dinosaurs from the Tetori Group (1). Professor S. Miura Memorial
Volume, 55-69.
Azuma, Kawagoshi and Miyagawa, 1995. Dinosaurs of the Tetori Group in Japan.
Fukui Prefectural Museum. 158 pp.
Azuma and Tomida, 1995. Early Cretaceous dinosaur fauna of the Tetori Group
in Japan. in Sun and Wang (eds.). Sixth Symposium on Mesozoic Terrestrial Ecosystems
and Biota, Short Papers. China Ocean Press, Beijing. 125-131.
Goto, Yabe and Sano, 2002. The research report of the Dinosaur Fossil Exploratory
Excavation held by Fukui Prefecture in 2001. Memoir of the Fukui Prefectural
Dinosaur Museum. 1, 102-118.
Azuma, 2003. Early Cretaceous vertebrates from Katsuyama City, Fukui Prefecture,
Japan. Memoir of the Fukui Prefectural Dinosaur Museum. 2, 17-21.
Kobayashi, Manabe, Ikegami, Tomida and Hayakawa, 2006. Dinosaurs from Japan.
in Lu, Kobayashi, Huang and Lee (eds.). Papers from the 2005 Heyuan International
Dinosaur Symposium. Geological Publishing House, Beijing. 87-102.
Shibata and Goto, 2008. Report of the 3rd Dinosaur Excavation Project in Katsuyama,
Fukui, 2007. Memoir of the Fukui Prefectural Dinosaur Museum. 7, 109-116.
Azuma and Shibata, 2010. Fukuititan nipponensis, a new titanosauriform
sauropod from the Early Cretaceous Tetori Group of Fukui Prefecture, Japan.
Acta Geologica Sinica. 84(3), 454-462.
D'Emic, 2012. The early evolution of titanosauriform sauropod dinosaurs. Zoological
Journal of the Linnean Society. 166, 624-671.
Mannion, Upchurch, Barnes and Mateus, 2013. Osteology of the Late Jurassic Portuguese
sauropod dinosaur Lusotitan atalaiensis (Macronaria) and the evolutionary
history of basal titanosauriforms. Zoological Journal of the Linnean Society.
168(1), 98-206.
unnamed Titanosauroidea (Huene, 1929)
Mid Cenomanian-Turonian, Late Cretaceous
Bajo Barreal Formation?, Chubut, Argentina
Material- (FMNH 13018) femur (2 m)
(FMNH 13019) femur (1.77 m)
(FMNH 13020) tibia (124 mm)
Comments- FMNH 13018 was referred to Argyrosaurus superbus by
Huene (1929), but cannot be compared to the holotypic forelimb of that taxon
and was assigned merely to Titanosauridae indet. or cf. Argyrosaurus
by Powell (2003) and Titanosauria indet. by Mannion and Otero (2012) in their
revision of the genus. Similarly, FMNH 13019 and 13020 were referred to Antarctosaurus
?wichmannianus by Huene, and assigned merely to Titanosauridae indet. or
cf. Argyrosaurus by Powell. Mannion and Otero again noted these cannot
be compared with Argyrosaurus, assigning them to Titanosauria indet.,
but did state the tibia lacks Antarctosaurus' subcircular distal end.
The femora were assigned to Titanosauria based on the distal articular surface
extending up the anterior face of the femur, which is present in Phuwiangosaurus
but lacking in e.g. Sauroposeidon and Euhelopus, so these materials
are assigned to Titanosauroidea here.
References- Huene, 1929. Los Saurisquios y Ornitisquios del Cretaceo
Argentino. Anales del Museo de La Plata. 3, 1-196.
Bonaparte and Gasparini, 1979. Los sauropodos de los grupos Neuquen y Chubut
y sus relaciones cronologicas. Actas V Congreso Geologico Argentino, Neuquen.
2, 393-406.
Powell, 2003. Revision of South American titanosaurid dinosaurs: Palaeobiological,
palaeobiogeographical and phylogenetic aspects. Records of the Queen Victoria
Museum. 111, 173 pp.
Mannion and Otero, 2012. A reappraisal of the Late Cretaceous Argentinean sauropod
dinosaur Argyrosaurus superbus, with a description of a new titanosaur
genus. Journal of Vertebrate Paleontology. 32(3), 614-638.
Phuwiangosaurus
Quiaowanlong
Yunmenglong Lu, Xu, Pu,
Zhang, Zhang, Jia, Chang, Zhang and Wei, 2013
Y. ruyangensis Lu, Xu, Pu, Zhang, Zhang, Jia, Chang, Zhang and
Wei, 2013
Aptian-Albian, Early Cretaceous
Haoling Formation, Henan, China
Holotype- (41HIII-0006) axis (~310 mm), third cervical vertebra (450 mm),
fourth cervical vertebra with fused rib (580 mm, rib 1.4 m), fifth cervical
vertebra with fused rib (~610 mm), sixth cervical vertebra with fused rib (~660
mm), seventh cervical vertebra with fused rib (600 mm excl. condyle), partial
eighth cervical vertebra, ninth cervical vertebra (520 mm), tenth cervical vertebra
(500 mm), incomplete mid-posterior dorsal vertebra (300 mm), two dorsal ribs,
first caudal vertebra (160 mm), second caudal vertebra (155 mm), third caudal
vertebra (180 mm), fourth caudal vertebra (190 mm), femur (1.92 m)
Diagnosis- (after Lu et al., 2013) distinct pleurocoel and fossa under
base of axial diapophysis; axial centroparapophyseal lamina parallel to long
axis of centrum; axial parapophysis situated at middle level of centrum; extremely
elongated cervical vertebrae; postaxial epipophyses very elongated and rod-like;
two fossae present on lateral longitudinal depression on cervical centra, anterior
one shallow and posterior one deep and pierces centrum; dorsal neural canal
triangular in posterior view and located within large concavity formed by centropostzygapophyseal
laminae and dorsal margin of neural canal; distal end of caudal neural spine
ball-shaped with coarse surfaces, but without postspinal fossa.
Comments- Lu et al. (2013) added Yunmenglong to Wilson's sauropod
analysis, finding it to be sister to Qiaowanlong then Erketu,
with this clade as somphospondylans sister to Titanosauria.
Reference- Lu, Xu, Pu, Zhang, Zhang, Jia, Chang, Zhang and Wei, 2013.
A new sauropod dinosaur (Dinosauria, Sauropoda) from the late Early Cretaceous
of the Ruyang Basin (central China). Cretaceous Research. 44, 202-213.
Malarguesaurus
Tangvayosaurus
Eutitanosauria sensu Carballido, Otero, Mannion, Salgado and Perez Moreno, 2022
Definition- (Saltasaurus loricatus + Patagotitan mayorum)
Reference- Carballido, Otero, Mannion, Salgado and Perez Moreno, 2022.
Titanosauria: A critical reappraisal of its systematics and the
relevance of the South American record. In Otero, Carballido and Pol
(eds.). South American Sauropodomorph Dinosaurs. Springer Earth System
Sciences. 269-298.
Antarctosauridae Olshevsky, 1978
= "Antarctosaurinae" Olshevsky, 1978 vide Powell, 1986
= Antarctosaurinae Olshevsky, 1978 vide Powell, 1986 vide Bonaparte, 1987
Comments- Olshevsky (1978) first used Antarctosauridae, and Powell (1986)
later used the subfamily version in his thesis. As theses do not count for nomenclature
under the ICZN, Bonaparte (1987) was the first to technically publish Antarctosaurinae,
though he did credit Powell for the name.
References- Olshevsky, 1978. The Archosaurian Taxa (excluding the Crocodylia).
Mesozoic Meanderings. 1, 50 pp.
Powell, 1986. Revisi�n de los titanosaurios de Am�rica del Sur.
PhD Thesis. Universidad Nacional de Tucum�n. 340 pp.
Bonaparte, 1987. The Late Cretaceous fauna of Los Alamitos, Patagonia, Argentina.
Parts I to IX. Revista Museo Argentino de Ciencias Naturales "Bernandino
Rivadavia". Paleontolog�a. 3(3), 103-178.
Antarctosaurus
Argentinosaurus
Brasilotitan Machado, Avilla,
Nava, Campos and Kellner, 2013
B. nemophagus Machado, Avilla, Nava, Campos and Kellner, 2013
Turonian-Santonian, Late Cretaceous
Adamantina Formation, Brazil
Holotype- (MPM 125R) partial dentary, incomplete anterior cervical vertebra,
incomplete posterior cervical vertebra with fused rib, several fragmentary cervical
ribs, three partial sacral vertebrae, ilial fragment, ischial fragment, partial
ungual, fragments
Paratype- ?...(MPM 126R) tooth (?x6x5 mm)
Diagnosis- (after Machado et al., 2013) dorsal symphyseal region of dentary
twisted medially; dorsal portion of dentary symphyseal contact thicker anteroposteriorly
than ventral part; anteriorly directed accessory prezygapophyseal articulation
surface on cervical vertebrae; cervical intraprezygapophyseal laminae V-shaped
in dorsal view.
Comments- The holotype was discovered in 2000. Machado et al. (2013)
believed Brasilotitan was a lithostrotian outside of Lognkosauria, Nemegtosauridae
and Saltasaurinae. The dentary was said to be most similar to Antarctosaurus
and Bonitasaura, which are lithostrotians or near the clade in this site's
phylogeny.
Reference- Machado, Avilla, Nava, Campos and Kellner, 2013. A new titanosaur
sauropod from the Late Cretaceous of Brazil. Zootaxa. 3701(3), 301-321.
Rukwatitan Gorscak, O'Connor,
Stevens and Roberts, 2014
R. bisepultus Gorscak, O'Connor, Stevens and Roberts, 2014
Aptian-Cenomanian, Early-Late Cretaceous
Namba Member of Galula Formation, Tanzania
Holotype- (RRBP 07409) incomplete third to last cervical vertebra with fused
ribs (492 mm ex. condyles), incomplete penultimate cervical vertebra with fused
partial ribs (478 mm ex. condyles), last partial cervical vertebra with fused
ribs, partial anterior dorsal neural arch, several partial dorsal ribs, proximal
caudal vertebra (104 mm), partial proximal caudal vertebra, incomplete proximal
caudal vertebra (107 mm), six mid caudal vertebrae (last incomplete; 113, 116,
121, 120, 131, 107 mm), two chevrons (one partial), partial scapula, partial
coracoids, incomplete humerus (910 mm), incomplete ulna, incomplete ilium, proximal
pubis
Paratype- (RRBP 03151) humerus (1.015 m)
Diagnosis- (after Gorscak et al., 2014) ventral surface of posterior
cervical vertebrae with weakly developed carotid processes posteriorly; posterior
cervicals with paired ventral shallow fossae anteriorly that are not separated
by thin keel; posterior cervical vertebrae with anteroposteriorly elongate and
undivided fossa posterior to anterior centrodiapophyseal lamina that deeply
invades ventral surface of diapophysis; accessory tubercle on ventral surface
of cervical rib capitulum, lateral to capituloparapophyseal suture; deep coracobrachialis
fossa on proximal humerus bounded by strong anteriorly projecting deltopectoral
crest and strong anteromedial ridge, extending to approximately humeral midshaft;
flattened anterior, medial, lateral, and posterior surfaces at humeral midshaft,
resulting in subquadrangular cross-section; pubic peduncle of ilium teardrop-shaped,
with 'tail' curving posteriorly, slightly longer transversely than anteroposteriorly.
Comments- Gorscak et al. (2014) recovered Rukwatitan as a titanosauroid
sister to Lithostrotia.
Reference- Gorscak, O'Connor, Stevens and Roberts, 2014. The basal titanosaurian
Rukwatitan bisepultus (Dinosauria, Sauropoda) from the middle Cretaceous
Galula Formation, Rukwa Rift Basin, southwestern Tanzania. Journal of Vertebrate
Paleontology. 34(5), 1133-1154.
Sonidosaurus
Colossosauria Gonz�lez Riga, Lamanna, Otero, Ortiz David, Kellner and Ibiricu, 2019
Official Definition- (Mendozasaurus neguyelap <- Saltasaurus loricatus, Epachthosaurus sciuttoi)
(Silva Junior, Martinelli, Lori, Marinho, Hechenleitner and Langer,
2022; originally Gonz�lez Riga, Lamanna, Otero, Ortiz David,
Kellner and Ibiricu, 2019; Registration Number 551)
Other definition- (Patagotitan mayorum <- Saltasaurus loricatus) (Carballido, Otero, Mannion, Salgado and Perez Moreno, 2022)
= Colossosauria sensu Carballido, Otero, Mannion, Salgado and Perez Moreno, 2022
Definition- (Patagotitan mayorum <- Saltasaurus loricatus)
References- Gonz�lez Riga,
Lamanna, Otero, Ortiz David, Kellner and Ibiricu, 2019. An overview of
the appendicular skeletal anatomy of South American titanosaurian
sauropods, with definition of a newly recognized clade. Anais da
Academia Brasileira de Ci�ncias. 91(suppl 2), e20180374.
Carballido, Otero, Mannion, Salgado and Perez Moreno, 2022.
Titanosauria: A critical reappraisal of its systematics and the
relevance of the South American record. In Otero, Carballido and Pol
(eds.). South American Sauropodomorph Dinosaurs. Springer Earth System
Sciences. 269-298.
Silva Junior, Martinelli, Lori, Marinho, Hechenleitner and Langer, 2022 (online 2021). Reassessment of Aeolosaurus maximus,
a titanosaur dinosaur from the Late Cretaceous of southeastern Brazil.
Historical Biology. 34(3), 403-411.
Bonitasaura
Lognkosauria Calvo, Porfiri, Gonz�lez
Riga and Kellner, 2007
Official Definition- (Futalognkosaurus dukei + Mendozasaurus neguyelap)
(Navarro, Ghilardi, Aureliano, D�ez D�az, Bandeira, Cattaruzzi, Iori,
Martine, Carvalho, Anelli, Fernandes and Zaher, 2022; Calvo et al., 2007; Registration Number 730)
References- Calvo, Porfiri, Gonz�lez Riga and Kellner, 2007. A
new Cretaceous terrestrial ecosystem from Gondwana with the description of a
new sauropod dinosaur. Anais da Academia Brasileira de Ci�ncias. 79(3),
529-541.
Navarro, Ghilardi, Aureliano, D�ez D�az, Bandeira, Cattaruzzi, Iori,
Martine, Carvalho, Anelli, Fernandes and Zaher, 2022. A new nanoid
titanosaur (Dinosauria: Sauropoda) from the Upper Cretaceous of Brazil.
Ameghiniana. 59(5), 317-354.
Drusilasaura Navarrete,
Casal and Martinez, 2011
D. deseadensis Navarrete, Casal and Martinez, 2011
Cenomanian-Turonian, Late Cretaceous
Upper Bajo Barreal Formation, Santa Cruz, Argentina
Holotype- (MPM-PV 2097/1; MPM-PV 2097/19) four dorsal vertebrae, partial
dorsal rib, sacral vertebra, ~first caudal neural arch, partial ~second caudal
vertebra, incomplete ?third caudal vertebra, incomplete ?fourth caudal vertebra,
incomplete ?fifth caudal vertebra, incomplete ?sixth caudal vertebra, two incomplete
chevrons, scapula (1.43 m), fragments
Diagnosis- (after Navarrete et al., 2011) two robust anterior dorsal
spinodiapophyseal laminae, one anterior and other posterior, which delimit an
elongate and deep supradiapophyseal cavity; small anterior dorsal circumneural
lamina surrounding the neural canal in posterior view; last sacral vertebra
with postspinal lamina expanded toward neural spine apex; prespinal laminae,
at least until ?fifth caudal vertebra, expanded toward neural spine apex; proximal
caudal tuberopostzygapophyseal laminae; ventral foramina, at least until ?fourth
caudal vertebrae; prezygapophyseal tuberosity jointed the prespinal lamina until
?fourth caudal vertebra.
Comments- Navarrete et al. (2011) considered this taxon close to Baurutitan
and Lognkosauria.
References- Navarrete, Casal and Mart�nez, 2008. Nuevos materiales
de Titanosauridae (Lognkosauria?) de la Formaci�n Bajo Barreal, Cret�cico
Superior, Santa Cruz, Argentina. In Congreso Latinoamericano de paleontologia
de vertebrados. 3, 176.
Navarrete, Casal and Martinez, 2011. Drusilasaura deseadensis gen. et
sp. nov., un nuevo titanosaurio (Dinosauria-Sauropoda), de la Formacion Bajo
Barreal, Cretacico Superior del norte de Santa Cruz, Argentina. Revista Brasileira
de Paleontologia. 14(1), 1-14.
Futalognkosaurus
Mendozasaurus
Quetecsaurus Gonzalez Riga
and David, 2014
Q. rusconii Gonzalez Riga and David, 2014
Mid-Late Turonian, Late Cretaceous
Cerro Lisandro Formation, Mendoza, Argentina
Holotype- (UNCUYO-LD-300) postorbital, two teeth (40x5x? mm), incomplete
atlas (30 mm), incomplete ~eleventh cervical vertebra (400 mm), partial anterior
dorsal vertebra, eight dorsal ribs (1.32 m), proximal caudal centrum (160 mm),
incomplete coracoid (250 mm deep), proximal humerus, distal radius, distal ulna,
metacarpal I (285 mm), metacarpal II (312 mm), metacarpal III (292 mm), incomplete
metacarpal IV (303 mm), incomplete metacarpal V (251 mm)
Diagnosis- (after Gonzalez Riga and David, 2014) atlantal intercentrum
with prominent anteroventral border and expanded posteroventral processes; posterior
cervical neural spines with incipient lateral laminae; humerus with strongly
sigmoid proximal border; rounded humeral proximomedial corner; angular humeral
proximolateral corner.
Comments- Gonzalez Riga and David (2014) recovered Quetecsaurus
as a eutitanosaur sister to Lognkosauria.
Reference- Gonzalez Riga and David, 2014. A new titanosaur (Dinosauria,
Sauropoda) from the Upper Cretaceous (Cerro Lisandro Formation) of Mendoza Province,
Argentina. Ameghiniana. 51(1), 3-25.
Traukutitan Valieri and Calvo,
2011
T. eocaudata Valieri and Calvo, 2011
Early Santonian, Late Cretaceous
Bajo de la Carpa Formation of the Rio Colorado Subgroup, Neuquen, Argentina
Holotype- (MUCPv 204) partial ?first caudal vertebra, partial ?second caudal
vertebra, partial ?fourth caudal vertebra, partial ?fifth caudal vertebra, ?sixth
caudal centrum, five partial mid caudal vertebrae, pubic fragments (lost), femora
(1.85 m)
Diagnosis- proximal caudal centra higher than wider; proximal caudal
centra strongly procoelous, with condyle placed in middle-dorsal part of centrum;
first caudal prezygodiapophyseal lamina subvertical (also in Futalognkosaurus);
first caudal vertebra with posteroventral placement of transverse process (also
in Malawisaurus, Mendozasaurus, Futalognkosaurus and Elaltitan);
single deep foramen ventral to base of first caudal transverse process (also
in Futalognkosaurus); mid caudal vertebrae wider than long, with procoelous-opisthoplatyan
centra, with upper border of posterior face slightly enlarged.
Comments- Salgado and Calvo (1993) initially described this as a basal
titanosaur of the same grade as Janenschia and Macrurosaurus.
The specimen was redescribed and named by Valieri and Calvo (2011), who noted
Salgado and Calvo's assignment of the material to the later Rio Colorado Formation
was incorrect. Note Valieri and Calvo report thirteen caudals are preserved,
but only illustrate ten, which is the number reported by Salgado and Calvo.
Valieri and Calvo assigned Traukutitan to Lognkosauria based on the combination
of strongly procoelous proximal caudals with weakly procoelous to platycoelous
mid caudals, and a few characters shared with Futalognkosaurus (see diagnosis),
but this has yet to be tested in a phylogenetic analysis.
References- Salgado and Calvo, 1993. Report of a sauropod with amphiplatyan
mid-caudal vertebae from the Late Cretaceous of Neuqu�n province (Argentina).
Ameghiniana. 30, 215-218.
Valieri and Calvo, 2011. Revision of MUCPv 204, a Senonian basal titanosaur
from northern Patagonia. Dinosaurios y Paleontolog�a desde Am�rica
Latina. Anales del III Congreso Latinoamericano de Paleontolog�a. Editorial
de la Universidad Nacional de Cuyo, Neuqu�n. 143-152.
Lithostrotia Upchurch, Barrett and Dodson,
2004 vide Wilson and Upchurch, 2003
Official Definition- (Malawisaurus dixeyi + Saltasaurus loricatus)
(Navarro, Ghilardi, Aureliano, D�ez D�az, Bandeira, Cattaruzzi, Iori,
Martine, Carvalho, Anelli, Fernandes and Zaher, 2022; Wilson and
Upchurch, 2003; Registration Number 728)
References- Wilson and Upchurch, 2003. A revision of Titanosaurus
Lydekker (Dinosauria - Sauropoda), the first dinosaur genus with a 'Gondwanan'
distribution. Journal of Systematic Palaeontology. 1(3), 125-160.
Upchurch, Barrett and Dodson, 2004. Sauropoda. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria (2nd edition). University of California Press.
259-322.
Navarro, Ghilardi, Aureliano, D�ez D�az, Bandeira, Cattaruzzi, Iori,
Martine, Carvalho, Anelli, Fernandes and Zaher, 2022. A new nanoid
titanosaur (Dinosauria: Sauropoda) from the Upper Cretaceous of Brazil.
Ameghiniana. 59(5), 317-354.
Malawisaurus
Lithostrotia incertae sedis
Dreadnoughtus Lacovara, Lamanna,
Ibiricu, Poole, Schroeter, Ullmann, Voegele, Boles, Carter, Fowler, Egerton,
Moyer, Coughenour, Schein, Harris, Martinez and Novas, 2014
D. schrani Lacovara, Lamanna, Ibiricu, Poole, Schroeter, Ullmann,
Voegele, Boles, Carter, Fowler, Egerton, Moyer, Coughenour, Schein, Harris,
Martinez and Novas, 2014
Campanian-Maastrichtian, Late Cretaceous
Cerro Fortaleza Formation, Santa Cruz, Argentina
Holotype- (MPM-Pv 1156) (~26 m, 59.3 tons, subadult) maxillary fragment,
tooth (35 mm), incomplete ~ninth cervical vertebra (1.13 m), cervical rib fragments,
~fourth dorsal vertebra (400 mm), ~fifth dorsal vertebra (470 mm), ~sixth dorsal
vertebra (200 mm), ~seventh dorsal vertebra (~300 mm), ~eighth dorsal vertebra
(350 mm), ~ninth dorsal centrum (410 mm), ~tenth dorsal centrum (330 mm), dorsal
vertebra, incomplete to fragmentary dorsal ribs, partial third to sixth sacral
vertebrae (~850 mm), partial first caudal vertebra (200 mm without condyle),
second caudal vertebra (350 mm), third caudal vertebra (300 mm), partial fourth
caudal vertebra (330 mm), fifth caudal vertebra (295 mm), sixth caudal vertebra
(313 mm), seventh caudal vertebra (292 mm), eighth caudal vertebra (265 mm),
ninth caudal vertebra (293 mm), tenth caudal vertebra (291 mm), eleventh caudal
vertebra (272 mm), twelfth caudal vertebra (295 mm), thirteenth caudal vertebra
(291 mm), fourteenth caudal vertebra (253 mm), fifteenth caudal vertebra (261
mm), sixteenth caudal vertebra (260 mm), seventeeth caudal vertebra (261 mm),
eighteenth caudal vertebra (245 mm), nineteenth caudal vertebra (251 mm), twentieth
caudal vertebra (246 mm), twenty-first caudal vertebra (225 mm), twenty-second
caudal vertebra (240 mm), twenty-third caudal vertebra (240 mm), twenty-fourth
caudal vertebra (235 mm), twenty-fifth caudal vertebra (215 mm), twenty-sixth
caudal vertebra (201 mm), twenty-seventh caudal vertebra (211 mm), twenty-eighth
caudal vertebra (206 mm), twenty-ninth caudal vertebra (183 mm), thirtieth caudal
vertebra (205 mm), thirty-first caudal vertebra (200 mm), thirty-second caudal
vertebra (175 mm), eighteen chevrons (35-441 mm), scapula (1.74 m), coracoid
(580 mm deep), sternal plates (1.14, 1.12 m), humerus (1.6 m), radius (950 mm),
ulna (1.01 m), incomplete ilia (1.31 m), pubes (1.4, 1.26 m), ischia (1.02,
1.01 m), femur (1.91 m), tibiae (1.2, 1.09 m), fibula (1.03 m), astragalus (230
mm trans), metatarsal I (210 mm), pedal ungual I (230 mm), metatarsal II (250
mm)
Paratype- (MPM-Pv 3546) partial ~fourth cervical vertebra, (some dorsal
vertebrae listed under holotype?), incomplete to fragmentary dorsal ribs, sacrum,
seven caudal vertebrae, five chevrons, incomplete ilia, pubes (1.2, 1 m), ischia,
femur (1.29 m)
Diagnosis- (after Lacovara et al., 2014) first caudal centrum with median
ventral keel; proximal caudal neural spines with extensively subdivided fossa
between spinoprezygapophyseal and spinopostzygapophyseal laminae; proximal caudal
neural spines with hypertrophied prespinal and postspinal laminae; middle caudal
neural spines with triangular anterodorsal process that extends well beyond
anterior margin of centrum; proximal chevrons with paddle-shaped ventral expansion;
posterodorsally-anteroventrally oriented ridge on medial surface of anterior
end of scapular blade; posteromedial surface of radius concave in proximal view;
distal end of radius nearly square, with subequal anteroposterior and mediolateral
dimensions.
Comments- Assignment of dorsal vertebrae to either holotype or paratype
is uncertain, although Lacovara et al. note the ~sixth and ~tenth belong to
the holotype.
The authors recovered Dreadnoughtus as a titanosauroid sister to Lithostrotia.
Reference- Lacovara, Lamanna, Ibiricu, Poole, Schroeter, Ullmann, Voegele,
Boles, Carter, Fowler, Egerton, Moyer, Coughenour, Schein, Harris, Martinez
and Novas, 2014. A gigantic, exceptionally complete titanosaurian sauropod dinosaur
from southern Patagonia, Argentina. Scientific Reports. 4, 6196.
Notocolossus Gonzalez
Riga, Lamanna, Ortiz Davis, Calvo and Coria, 2016
N. gonzalezparejasi Gonzalez Riga, Lamanna, Ortiz Davis, Calvo
and Coria, 2016
Late Coniacian-Early Santonian, Late Cretaceous
Plottier Formation of the Neuquen Group, Mendoza, Argentina
Holotype- (UNCUYO-LD 301) (~60 tons) incomplete anterior dorsal vertebra,
incomplete proximal caudal vertebra (302 mm), humerus (1.76 m), proximal pubis
(femur ~2.17 m)
Paratype- (UNCUYO-LD 302) seven partial proximal caudal vertebrae (207,
188 mm), seven chevrons (six partial), (femur ~1.28 m) astragalus (67 mm wide),
metatarsal I (164 mm), phalanx I-1 (45 mm), pedal ungual I (64 mm), metatarsal
II (185 mm), phalanx II-1 (96 mm), pedal ungual II (102 mm), metatarsal III
(197 mm), phalanx III-1 (115 mm), pedal ungual III (80 mm), metatarsal IV (218
mm), phalanx IV-1 (98 mm), phalanx IV-2 (37 mm), metatarsal V (196 mm)
Diagnosis- (after Gonzalez Riga et al., 2016) anterior dorsal vertebra
with parapophyseal centrodiapophyseal fossa subdivided by two accessory laminae
(one subvertical and visible in anterior and lateral views, the other anterodorsally
oriented and visible only in lateral view); proximal caudal vertebrae with laminae
that converge ventrally on anterior surface of neural spine, not reaching prezygapophyses
and forming a V-shaped conformation in anterior view; humerus with
greatly expanded proximomedial process, the proximal apex of which lies well
medial to humeral midshaft; proportionally wide proximal humerus (proximal width:
midshaft width = 2.9); proximolaterally-distomedially oriented ridge bounding
distal edge of humeral coracobrachialis fossa; metatarsal I with
proximal dorsoventral diameter greater than proximodistal length of element;
short metatarsal III (1.2 times length of metatarsal I); proximal pedal phalanges
more than half as wide as their corresponding metatarsals are long; pedal unguals
reduced, rugose, and distally truncated.
Comments- Gonzalez Riga et al. (2016) recovered this as a non-saltasaurid
lithostrotian more derived than Malawisaurus, sister to Dreadnoughtus.
Reference- Gonzalez Riga, Lamanna, Ortiz Davis, Calvo and Coria, 2016.
A gigantic new dinosaur from Argentina and the evolution of the sauropod hind
foot. Scientific Reports. 6:19165.
"Sauropodus" Simon vide Anonymous,
2001
Late Cenomanian, Late Cretaceous
Huincul Formation of the Rio Limay Subgroup of the Neuquen Group, Neuquen, Argentina
Material- (~20 m) tooth, vertebra (1 m), ribs?, scapula, humerus (1.7
m), femur, tibiae, fibula
Comments- Discovered in 1999, this is supposed to be a derived titanosaur,
so may be a lithostrotian. The name may be a mistranslation of "sauropod",
as it does not appear in the Spanish article (Chavez, 2001) and Simon only says
it may be a new taxon.
Reference- Anonymous, 2001. Argentine find may be previously unknown
herbivore. Reuters. 2-8-2001.
Chavez, 2001. Rescatan los restos f�siles de un gran dinosaurio herb�voro.
La Nacion. 2-9-2001.
Simon, 2001. A giant sauropod from the Upper Cretaceous of El Chocon. Ameghiniana.
38(4S), 19R.
unnamed lithostrotian (Huene, 1929)
Cenomanian-Turonian, Late Cretaceous
Cerro Castillo or Bajo Barreal Formation, Chubut, Argentina
Material- (DGM coll.) proximal caudal centrum (250 mm)
Comments- Huene (1929) referred this to Argyrosaurus, but it cannot
be compared to the holotypic forelimb of that taxon and was assigned merely
to Lithostrotia indet. by Mannion and Otero (2012) in their revision of the
genus. This is based on the strongly procoelous centrum.
References- Huene, 1929. Los Saurisquios y Ornitisquios del Cretaceo
Argentino. Anales del Museo de La Plata. 3, 1-196.
Bonaparte and Gasparini, 1979. Los sauropodos de los grupos Neuquen y Chubut
y sus relaciones cronologicas. Actas V Congreso Geologico Argentino, Neuquen.
2, 393-406.
Mannion and Otero, 2012. A reappraisal of the Late Cretaceous Argentinean sauropod
dinosaur Argyrosaurus superbus, with a description of a new titanosaur
genus. Journal of Vertebrate Paleontology. 32(3), 614-638.
unnamed lithostrotian (Huene, 1929)
Maastrichtian, Late Cretaceous
Chorrillo Formation, Santa Cruz, Argentina
Material- (MACN 5205) three mid-distal caudal vertebrae (230, 220, 220 mm)
Comments- Huene (1929) referred these to Argyrosaurus, but they
cannot be compared to the holotypic forelimb of that taxon and was assigned
merely to Lithostrotia indet. by Mannion and Otero (2012) in their revision
of the genus. This is based on the strongly procoelous centrum.
References- Huene, 1929. Los Saurisquios y Ornitisquios del Cretaceo
Argentino. Anales del Museo de La Plata. 3, 1-196.
Powell, 2003. Revision of South American titanosaurid dinosaurs: Palaeobiological,
palaeobiogeographical and phylogenetic aspects. Records of the Queen Victoria
Museum. 111, 173 pp.
Mannion and Otero, 2012. A reappraisal of the Late Cretaceous Argentinean sauropod
dinosaur Argyrosaurus superbus, with a description of a new titanosaur
genus. Journal of Vertebrate Paleontology. 32(3), 614-638.
Saltasauroidea Powell, 1992 vide Fran�a, J�lio, Riff, Hsiou and Langer, 2016
Official Definition- (Lirainosaurus astibiae + Opisthocoelicaudia skarzynskii + Saltasaurus loricatus) (Navarro, Ghilardi, Aureliano, D�ez D�az, Bandeira, Cattaruzzi, Iori,
Martine, Carvalho, Anelli, Fernandes and Zaher, 2022; Registration Number 719)
Other definitions- (Saltasaurus loricatus <- Nemegtosaurus mongoliensis) (modified from Fran�a, J�lio, Riff, Hsiou and Langer, 2016)
(Saltasaurus loricatus <- Patagotitan mayorum) (Carballido, Otero, Mannion, Salgado and Perez Moreno, 2022)
Comments- Note Carballido et
al.'s (2022) definition of Saltasauroidea cannot be applied to this
cladogram given the uncertain position of lognkosaurs (and thus Patagotitan).
References- Powell, 1992. Osteologia de Saltasaurus loricatus
(Sauropoda - Titanosauridae) del Cret�cico Superior del noroeste Argentino. In Sanz and Buscalioni (eds.).
Los Dinosaurios y Su Entorno Biotico: Actas del Segundo Curso de Paleontologia
in Cuenca. Institutio "Juan de Valdes", Cuenca, Argentina. 165-230.
Fran�a, J�lio, Riff, Hsiou and Langer, 2016. New lower jaw and teeth referred to Maxakalisaurus topai (Titanosauria: Aeolosaurini) and their implications for the phylogeny of titanosaurid sauropods. PeerJ. 4, e2054.
Carballido, Otero, Mannion, Salgado and Perez Moreno, 2022.
Titanosauria: A critical reappraisal of its systematics and the
relevance of the South American record. In Otero, Carballido and Pol
(eds.). South American Sauropodomorph Dinosaurs. Springer Earth System
Sciences. 269-298.
Navarro, Ghilardi, Aureliano, D�ez D�az, Bandeira, Cattaruzzi, Iori,
Martine, Carvalho, Anelli, Fernandes and Zaher, 2022. A new nanoid
titanosaur (Dinosauria: Sauropoda) from the Upper Cretaceous of Brazil.
Ameghiniana. 59(5), 317-354.
Aegyptosaurus
Atacamatitan Kellner, Rubilar-Rogers,
Vargas and Suarez, 2011
A. chilensis Kellner, Rubilar-Rogers, Vargas and Suarez, 2011
Late Cretaceous
Tolar Formation, Chile
Holotype- (SGO-PV-961) two partial mid-posterior dorsal vertebrae, dorsal
ribs, two mid-distal caudal vertebrae (86 mm), caudal fragments, ?sternal fragment,
proximal humerus, incomplete femur (1.1 m)
Diagnosis- dorsal centra pierced by pleurocoels, which are rounded or
elliptical but do not acuminate posteriorly; ventral surface of dorsal vertebra
strongly concave; distal caudals with laterally compressed neural spine that
bears blade-like anterior margin; femur with shaft getting gradually narrower
until two thirds of its length.
Comments- The holotype was discovered in 2001. Kellner et al. (2011)
believed Atacamatitan was a lithostrotian more derived than Malawisaurus,
but not an aeolosaurin or saltasaurine. Rubilar-Rogers et al. (2012) added Atacamatitan
to Wilson's sauropod matrix and found it to be a lithostrotian more derived
than Malawisaurus but outside Saltasauridae.
References- Kellner, Rubilar-Rogers, Vargas and Suarez, 2011. A new titanosaur
sauropod from the Atacama desert, Chile. Anais da Academia Brasileira de Ci�ncias.
83(1), 211-219.
Rubilar-Rogers and Gutstein, 2012. Los titanosaurios de Chile y su contexto
filogen�tico y biogeogr�fico. Bolet�n del Museo Nacional
de Historia Natural (Santiago). 61, 55-73.
Rubilar-Rogers, Otero, Yury-Yanez, Vargas and Gutstein, 2012. An overview of
the dinosaur fossil record from Chile. Journal of South American Earth Sciences.
37, 242-255.
Isisaurus
Paralititan
Petrobrasaurus Filippi,
Canudo, Salgado, Garrido, Garcia, Cerda and Otero, 2011
P. puestohernandezi Filippi, Canudo, Salgado, Garrido, Garcia,
Cerda and Otero, 2011
Santonian, Late Cretaceous
Plottier Formation, Neuquen, Argentina
Holotype- (MAU-Pv-PH-449) two teeth, partial ?mid cervical centrum, cervical
rib fragments, three incomplete posterior dorsal vertebrae, incomplete posterior
dorsal neural arch, incomplete dorsal centrum, several dorsal rib fragments,
incomplete first caudal centrum, five proximal caudal vertebrae, chevron fragments,
sternal plates, humerus (1.2 m), metacarpal II (395 mm), metacarpal IV (350
mm), metacarpal V (315 mm), incomplete metacarpal, ilial fragment, pubis, femora
(1.572, 1.575 m), partial tibia, distal tibia, fragments
Diagnosis- (after Filippi et al., 2011) posterior dorsal accessory posterior
centrodiapophyseal lamina more developed than posterior centrodiapophyseal lamina;
posterior dorsal posterior centroparapophyseal lamina extends from point of
contact with accessory posterior centrodiapophyseal lamina; posterior dorsal
vertebrae with broad concave surface delimited by bony rim of posterior centroparapophyseal
lamina and rim located on dorsal edge of pleurocoel; presence of deep fossa,
subcircular in outline, on slightly concave surface delimited by posterior dorsal
accessory posterior centrodiapophyseal lamina, posterior centrodiapophyseal
lamina, and bony rim corresponding to extension of posterior centroparapophyseal
lamina; T-shaped anteroposterior expansion on laminae of posterior dorsal neural
arch (pcdl and apcdl); deep fossa between proximal caudal spinoprezygapophyseal
lamina and postzygapophysis, bearing small fossae; proximal caudal prespinal
lamina distally thickened; subcircular small fossae on both sides of base of
proximal caudal prespinal lamina; T-shaped anteroposterior expansion in proximal
caudal spinoprezygapophyseal lamina.
Comments- The holotype was discovered in 2006-2007. Filippi et al. (2011)
reported that adding Petrobrasaurus to Calvo et al.'s titanosaur analysis
resulted in a polytomy in Eutitanosauria, though they referred the taxon to
Titanosauria incertae sedis.
Reference- Filippi, Canudo, Salgado, Garrido, Garcia, Cerda and Otero,
2011. A new sauropod titanosaur from the Plottier Formation (Upper Cretaceous)
of Patagonia (Argentina). Geologica Acta. 9(1), 1-12.
Titanosauridae Lydekker, 1885
Other definitions- (Argentinosaurus huinculensis + Malawisaurus
dixeyi + Trigonosaurus pricei + Epachthosaurus scuttoi + Alamosaurus
sanjuanensis + Opisthocoelicaudia skarzynskii + Aeolosaurus rionegrinus
+ Saltasaurus loricatus) (modified from Salgado et al., 1997)
(Argentinosaurus huinculensis + Malawisaurus dixeyi + Epachthosaurus
scuttoi + Alamosaurus sanjuanensis + Opisthocoelicaudia skarzynskii
+ Aeolosaurus rionegrinus + Saltasaurus loricatus) (modified from
Gonzalez-Riga, 2003)
(Epachthosaurus scuttoi + Saltasaurus loricatus) (modified from
Salgado, 2003)
= Titanosaurinae Lydekker, 1885 vide Nopcsa, 1928
Comments- This family was abandoned by Wilson and Upchurch (2003) because
they view Titanosaurus indicus as indeterminate, but the ICZN does not
dictate family-level taxa must be based on diagnostic species. Unfortunately,
the phylogenetic position of Titanosaurus within Titanosauroidea has
not been examined recently, so whether Titanosauridae is a senior synonym of
Saltasauridae, Nemegtosauridae, etc. is still unknown. None of the proposed
definitions use Titanosaurus indicus as an internal specifier, so are
problematic.
Contra Olshevsky (1991), Marsh's (1895) Titanosauridae was not a homonym based
on his 1877 genus Titanosaurus (preoccupied by Lydekker's genus and replaced
with Atlantosaurus). Instead, it is merely a reuse of Lydekker's Titanosauridae
as demonstrated by the listed genera (Titanosaurus, Argyrosaurus)
and characters (e.g. biconvex first caudal; procoelous more distal caudals;
rectangular coracoid).
References- Lydekker, 1885. The Reptilia & Amphibia of the Maleria
& Denwa Groups. Memoirs of the Geological Survey of India. Palaeontologia
Indica, Series IV. Indian Pretertiary Vertebrata. 1(5), 1-38.
Marsh, 1895. On the affinities and classification of the dinosaurian reptiles.
American Journal of Science. 50(300), 483-498.
Nopcsa, 1928. The genera of reptiles. Palaeobiologica. 1, 163-188.
Olshevsky, 1991. A Revision of the Parainfraclass Archosauria Cope, 1869, Excluding
the Advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.
Salgado, Coria and Calvo, 1997. Evolution of titanosaurid sauropods. I: Phylogenetic
analysis based on the postcranial evidence. Ameghiniana. 34(1), 3-32.
Gonz�lez-Riga, 2003. A new titanosaur (Dinosauria, Sauropoda) from the
Upper Cretaceous of Mendoza Province, Argentina. Ameghiniana. 40(2), 155-172.
Salgado, 2003. Should we abandon the name Titanosauridae? Some comments on the
taxonomy of titanosaurian sauropods (Dinosauria). Revista Espa�ola de
Paleontolog�a. 18(1), 15-21.
Wilson and Upchurch, 2003. A revision of Titanosaurus Lydekker (Dinosauria
- Sauropoda), the first dinosaur genus with a 'Gondwanan' distribution. Journal
of Systematic Palaeontology. 1(3), 125-160.
Titanosaurus
Lirainosaurinae D�ez D�az, Garcia, Pereda Suberbiola, Jentgen-Ceschino, Stein, Godefroit and Valentin, 2018
Official Definition- (Lirainosaurus astibiae + Ampelosaurus atacis)
(Navarro, Ghilardi, Aureliano, D�ez D�az, Bandeira, Cattaruzzi, Iori,
Martine, Carvalho, Anelli, Fernandes and Zaher, 2022; D�ez D�az,
Garcia, Pereda Suberbiola, Jentgen-Ceschino, Stein, Godefroit and
Valentin, 2018; Registration Number 727)
References- D�ez D�az, Garcia, Pereda Suberbiola, Jentgen-Ceschino, Stein, Godefroit and Valentin, 2018. The titanosaurian dinosaur Atsinganosaurus velauciensis
(Sauropoda) from the Upper Cretaceous of southern France: New material,
phylogenetic affinities, and palaeobiogeographical implications. Cretaceous Research. 91, 429-456.
Navarro, Ghilardi, Aureliano, D�ez D�az, Bandeira, Cattaruzzi, Iori,
Martine, Carvalho, Anelli, Fernandes and Zaher, 2022. A new nanoid
titanosaur (Dinosauria: Sauropoda) from the Upper Cretaceous of Brazil.
Ameghiniana. 59(5), 317-354.
Lirainosaurus Sanz, Powell,
Le Loueff, Martinez and Pereda-Suberbiola, 1999
L. astibiae Sanz, Powell, Le Loueff, Martinez and Pereda-Suberbiola,
1999
Late Campanian-Early Maastrichtian, Late Cretaceous
Lano quarry, Burgos, Spain
Holotype- (MCNA 7458) (at least four individuals; ~4-6 m, ~1.7-4 tons) first
caudal vertebra
Paratypes- ..(MCNA 1812) distal caudal vertebra (118 mm)
..(MCNA 3160) incomplete femur
..(MCNA 7439) (subadult) braincase fragment
..(MCNA 7440) tooth
..(MCNA 7441) tooth
..(MCNA 7442) mid-posterior dorsal vertebra
..(MCNA 7443) ~ninth dorsal vertebra (144 mm)
..(MCNA 7444) dorsal vertebra
..(MCNA 7445) incomplete ~first dorsal vertebra
..(MCNA 7446) partial posterior dorsal vertebra (125 mm)
..(MCNA 7447) incomplete anterior dorsal vertebra
..(MCNA 7448) dorsal vertebra
..(MCNA 7449) dorsal vertebra
..(MCNA 7450) dorsal vertebra
..(MCNA 7451) proximal caudal vertebra (111 mm)
..(MCNA 7452) distal caudal vertebra (102 mm)
..(MCNA 7453) distal caudal vertebra
..(MCNA 7454) distal caudal vertebra (88 mm)
..(MCNA 7455) proximal caudal vertebra
..(MCNA 7456) partial dorsal vertebra
..(MCNA 7457) proximal caudal vertebra (133 mm)
..(MCNA 7459) incomplete scapula
..(MCNA 7460) incomplete coracoid
..(MCNA 7461) partial sternal plate
..(MCNA 7462) incomplete humerus
..(MCNA 7463) humerus (531 mm)
..(MCNA 7464) incomplete humerus
..(MCNA 7465) incomplete humerus
..(MCNA 7466) ilial fragment
..(MCNA 7467) incomplete pubis
..(MCNA 7468) femur (682 mm)
..(MCNA 7470) femur
..(MCNA 7471) tibia
..(MCNA 7472) incomplete fibula
..(MCNA 7473) osteoderm
..(MCNA 7474) osteoderm
Referred- ..(MCNA 1816) dorsal neural arch (Diez Diaz et al., 2013a)
..(MCNA 1817) incomplete femur (Diez Diaz et al., 2013b)
..(MCNA 1818) dorsal neural arch (Diez Diaz et al., 2013a)
..(MCNA 1846) incomplete coracoid (Diez Diaz et al., 2013b)
..(MCNA 1854) (adult) tooth (?x6.6x5.9 mm) (Diez Diaz et al., 2012)
..(MCNA 1882) proximal caudal vertebra (117 mm) (Diez Diaz et al., 2013a)
..(MCNA 1883) partial dorsal vertebra (Diez Diaz et al., 2013a)
..(MCNA 2203) incomplete tibia (Diez Diaz et al., 2013b)
..(MCNA 2207) distal caudal vertebra (Diez Diaz et al., 2013a)
..(MCNA 2208) caudal vertebra (Diez Diaz et al., 2013a)
..(MCNA 2211) distal caudal vertebra (Diez Diaz et al., 2013a)
..(MCNA 3157) ulna (114.3 mm) (Diez Diaz et al., 2013b)
..(MCNA 3158) coracoid (233 mm) (Diez Diaz et al., 2013b)
..(MCNA 3159) incomplete anterior dorsal vertebra (Diez Diaz et al., 2013a)
..(MCNA 7458) first caudal vertebra (136 mm) (Sanz et al., 1999)
..(MCNA 7495) distal caudal vertebra (Sanz et al., 1999)
..(MCNA 8551) incomplete posterior dorsal vertebra (Diez Diaz et al., 2013a)
..(MCNA 8604) dorsal vertebral fragment (Diez Diaz et al., 2013a)
..(MCNA 8605) distal caudal vertebra (122 mm) (Diez Diaz et al., 2013a)
..(MCNA 8606) mid caudal vertebra (Diez Diaz et al., 2013a)
..(MCNA 8607) mid caudal vertebra (125 mm) (Diez Diaz et al., 2013a)
..(MCNA 8608) partial dorsal vertebra (Diez Diaz et al., 2013a)
..(MCNA 8609) ilial fragment (Diez Diaz et al., 2013b)
..(MCNA 9410) incomplete fibula (Diez Diaz et al., 2013b)
..(MCNA 9647) distal caudal vertebra (57 mm) (Diez Diaz et al., 2013a)
..(MCNA 12402) (juvenile) tooth (10.2x2.3x2.3 mm) (Diez Diaz et al., 2012)
..(MCNA 13388) mid caudal vertebra (73 mm) (Diez Diaz et al., 2013a)
..(MCNA 13851) distal caudal vertebra (114 mm) (Diez Diaz et al., 2013a)
..(MCNA 13852) dorsal vertebral fragment (Diez Diaz et al., 2013a)
..(MCNA 13853) mid caudal vertebra (Diez Diaz et al., 2013a)
..(MCNA 13854) distal caudal vertebra (Diez Diaz et al., 2013a)
..(MCNA 13855) incomplete scapula (Diez Diaz et al., 2013b)
..(MCNA 13856) dorsal vertebral fragment (Diez Diaz et al., 2013a)
..(MCNA 13857) distal caudal vertebra (172 mm) (Diez Diaz et al., 2013a)
..(MCNA 13858) partial dorsal vertebra (Diez Diaz et al., 2013a)
..(MCNA 13859) incomplete posterior dorsal vertebra (151 mm) (Diez Diaz et al.,
2013a)
..(MCNA 13860) tibia (412 mm) (Diez Diaz et al., 2013b)
..(MCNA 13861) ilial fragment (Diez Diaz et al., 2013b)
..(MCNA 13864) (juvenile) tooth (Diez Diaz et al., 2012)
..(MCNA 13865) (juvenile) tooth (9.2x2.6x2.3 mm) (Diez Diaz et al., 2012)
..(MCNA 13866) (juvenile) tooth (?x2x2 mm) (Diez Diaz et al., 2012)
..(MCNA 13867) (adult) tooth (?x7.3x6.5 mm) (Diez Diaz et al., 2012)
..(MCNA 13868) (juvenile) tooth (12.6x2.6x2.3 mm) (Diez Diaz et al., 2012)
..(MCNA 13869) (juvenile) tooth (10.5x2x2 mm) (Diez Diaz et al., 2012)
..(MCNA 13870) (juvenile) tooth (?x2.4x2.2 mm) (Diez Diaz et al., 2012)
..(MCNA 13871) (juvenile) tooth (?x1.7x1.6 mm) (Diez Diaz et al., 2012)
..(MCNA 13872) (juvenile) tooth (?x2.7x2.4 mm) (Diez Diaz et al., 2012)
..(MCNA 13873) (juvenile) tooth (?x~2.3x~2.1 mm) (Diez Diaz et al., 2012)
..(MCNA 13874) (juvenile) tooth (?x~2.1x~2 mm) (Diez Diaz et al., 2012)
..(MCNA 13875) (juvenile) tooth (?x2.3x2 mm) (Diez Diaz et al., 2012)
..(MCNA 13876) (juvenile) tooth (?x1.7x1.6 mm) (Diez Diaz et al., 2012)
..(MCNA 13877) (juvenile) tooth (13.3x2.3x2.2 mm) (Diez Diaz et al., 2012)
..(MCNA 13878) (adult) tooth (16.9x4.2x3.5 mm) (Diez Diaz et al., 2012)
..(MCNA 13913) (subadult) incomplete braincase (Diez Diaz et al., 2011)
..(MCNA 13914) (juvenile) tooth (?x3.2x2.4 mm) (Diez Diaz et al., 2012)
..(MCNA 14231) (juvenile) tooth (10.7x2.7x2.3 mm) (Diez Diaz et al., 2012)
..(MCNA 14232) (juvenile) tooth (?x~2.4x~2.3 mm) (Diez Diaz et al., 2012)
..(MCNA 14233) (juvenile) tooth (?x~1.9x~1.9 mm) (Diez Diaz et al., 2012)
..(MCNA 14234) (juvenile) tooth (9x2x1.9 mm) (Diez Diaz et al., 2012)
..(MCNA 14235) (juvenile) tooth (?x2.1x2 mm) (Diez Diaz et al., 2012)
..(MCNA 14236) (juvenile) tooth (?x~2.5x~2.3 mm) (Diez Diaz et al., 2012)
..(MCNA 14237) (juvenile) tooth (8x2.1x1.9 mm) (Diez Diaz et al., 2012)
..(MCNA 14238) (adult) tooth (24.8x7.1x7.1 mm) (Diez Diaz et al., 2012)
..(MCNA 14239) (adult) tooth (?x7.4x7.3 mm) (Diez Diaz et al., 2012)
..(MCNA 14429) cervical centrum (~180 mm) (Diez Diaz et al., 2013a)
..(MCNA 14430) dorsal neural arch (Diez Diaz et al., 2013a)
..(MCNA 14431) dorsal neural arch (Diez Diaz et al., 2013a)
..(MCNA 14432) dorsal neural arch (Diez Diaz et al., 2013a)
..(MCNA 14433) dorsal neural arch (Diez Diaz et al., 2013a)
..(MCNA 14434) dorsal neural arch (Diez Diaz et al., 2013a)
..(MCNA 14435) distal caudal vertebra (100 mm) (Diez Diaz et al., 2013a)
..(MCNA 14436) caudal vertebra (Diez Diaz et al., 2013a)
..(MCNA 14437) distal caudal vertebra (116 mm) (Diez Diaz et al., 2013a)
..(MCNA 14438) distal caudal vertebra (Diez Diaz et al., 2013a)
..(MCNA 14439) proximal caudal vertebra (126 mm) (Diez Diaz et al., 2013a)
..(MCNA 14440) distal caudal vertebra (Diez Diaz et al., 2013a)
..(MCNA 14441) proximal caudal vertebra (Diez Diaz et al., 2013a)
..(MCNA 14442) caudal vertebra (Diez Diaz et al., 2013a)
..(MCNA 14443) proximal caudal vertebra (140 mm) (Diez Diaz et al., 2013a)
..(MCNA 14444) proximal caudal vertebra (54 mm) (Diez Diaz et al., 2013a)
..(MCNA 14445) proximal caudal vertebra (Diez Diaz et al., 2013a)
..(MCNA 14446) proximal caudal vertebra (152 mm) (Diez Diaz et al., 2013a)
..(MCNA 14447) distal caudal vertebra (103 mm) (Diez Diaz et al., 2013a)
..(MCNA 14448) distal caudal vertebra (90 mm) (Diez Diaz et al., 2013a)
..(MCNA 14449) distal caudal vertebra (73 mm) (Diez Diaz et al., 2013a)
..(MCNA 14450) distal caudal vertebra (135 mm) (Diez Diaz et al., 2013a)
..(MCNA 14451) proximal caudal vertebra (Diez Diaz et al., 2013a)
..(MCNA 14452) distal caudal vertebra (Diez Diaz et al., 2013a)
..(MCNA 14453) caudal vertebra (Diez Diaz et al., 2013a)
..(MCNA 14454) caudal vertebra (Diez Diaz et al., 2013a)
..(MCNA 14455) caudal vertebra (Diez Diaz et al., 2013a)
..(MCNA 14456) fragmentary dorsal rib (Diez Diaz et al., 2013a)
..(MCNA 14457) fragmentary dorsal rib (Diez Diaz et al., 2013a)
..(MCNA 14458) fragmentary dorsal rib (Diez Diaz et al., 2013a)
..(MCNA 14459) fragmentary dorsal rib (Diez Diaz et al., 2013a)
..(MCNA 14460) chevron (Diez Diaz et al., 2013a)
..(MCNA 14461) scapula (722 mm) (Diez Diaz et al., 2013b)
..(MCNA 14462) partial scapula (Diez Diaz et al., 2013b)
..(MCNA 14463) partial humerus (Diez Diaz et al., 2013b)
..(MCNA 14464) partial ilium (Diez Diaz et al., 2013b)
..(MCNA 14465) femur (680 mm) (Diez Diaz et al., 2013b)
..(MCNA 14466) femur (816 mm) (Diez Diaz et al., 2013b)
..(MCNA 14467) partial femur (Diez Diaz et al., 2013b)
..(MCNA 14468) incomplete femur (Diez Diaz et al., 2013b)
..(MCNA 14469) tibia (Diez Diaz et al., 2013b)
..(MCNA 14470) partial tibia (Diez Diaz et al., 2013b)
..(MCNA 14471) fibula (423 mm) (Diez Diaz et al., 2013b)
..(MCNA 14473) incomplete osteoderm (Diez Diaz et al., 2013b)
..(MCNA 14474) metatarsal III (198 mm), incomplete osteoderm (Diez Diaz et al.,
2013b)
Late Campanian-Early Maastrichtian, Late Cretaceous
Sierra Perenchiza Formation, Valencia, Spain
(CMPGC 49) caudal vertebra (Company et al., 2009)
(MGUV 16438) proximal caudal vertebra (Company et al., 2009)
(MGUV 16446) sternal fragment (Company et al., 2009)
(MGUV 16448) caudal vertebra (Company et al., 2009)
(MGUV 16449) humeral fragment (Company et al., 2009)
(MGUV 16450) femur (Company et al., 2009)
(MGUV 16454) ?humeral fragment (Company et al., 2009)
(MGUV 17016) distal caudal vertebra (Company et al., 2009)
(MGUV 17027) caudal vertebra (Company et al., 2009)
(MGUV 17028) caudal vertebra (Company et al., 2009)
(MGUV 17044) caudal vertebra (Company et al., 2009)
(MGUV 17045) ulna (Company et al., 2009)
(MGUV 17090) caudal vertebra (Company et al., 2009)
(MGUV 17092) caudal centrum (Company et al., 2009)
(MGUV 17093) caudal centrum (Company et al., 2009)
(MGUV 17094) caudal vertebra (Company et al., 2009)
(MGUV 17095) caudal vertebra (Company et al., 2009)
(MGUV 17096) caudal vertebra (Company et al., 2009)
(MGUV 17097) caudal fragment (Company et al., 2009)
(MGUV 17098) caudal fragment (Company et al., 2009)
(MGUV 17100) caudal vertebra (Company et al., 2009)
(MGUV 17101) caudal vertebra (Company et al., 2009)
(MGUV 17105) caudal vertebra (Company et al., 2009)
(MGUV 17165) humeral fragment (Company et al., 2009)
(MGUV 17166) humeral fragment (Company et al., 2009)
(MGUV 17168) chevron (Company et al., 2009)
(MGUV 17169) caudal vertebra (Company et al., 2009)
(MGUV 17192) chevron fragment (Company et al., 2009)
(MGUV 17194) femur (Company et al., 2009)
(MGUV 17229) dorsal neural arch (Company et al., 2009)
(MGUV 17235) femur (Company et al., 2009)
(MGUV 17238) coracoids (Company et al., 2009)
(MGUV 17243) caudal vertebra (Company et al., 2009)
(MGUV 17272) caudal vertebra (Company et al., 2009)
(MGUV 17274) ulna (Company et al., 2009)
(MGUV 17280) caudal vertebra (Company et al., 2009)
(MGUV 17286) distal humerus (Company et al., 2009)
(MGUV 17288) caudal vertebra (Company et al., 2009)
(MGUV 17294) femur (Company et al., 2009)
(MGUV 17295) dorsal vertebra (Company et al., 2009)
(MGUV 17314) proximal caudal vertebra (Company et al., 2009)
(MGUV 17315) dorsal vertebra (Company et al., 2009)
(MGUV 17320) dorsal vertebra (Company et al., 2009)
(MGUV 17332) chevron (Company et al., 2009)
(MGUV 17337) caudal fragment (Company et al., 2009)
(MGUV 17374) dorsal rib fragment (Company et al., 2009)
(MGUV 17539) caudal vertebra (Company et al., 2009)
(MGUV 17552) humerus (Company et al., 2009)
(MGUV 17645) scapula (Company et al., 2009)
(MGUV 17683) humeral fragment (Company et al., 2009)
(MGUV 17705) caudal vertebra (Company et al., 2009)
Early Campanian-Late Maastrichtian, Late Cretaceous
Villalba de la Sierra Formation, Guadalajara, Spain
?(MCCM-BD 001) distal caudal centrum (Ortega and Perez Garcia, 2009)
Diagnosis- (after Sanz et al., 1999) accessory lamina in proximal caudal
interzygapophyseal fossa; spinopostzygapophyseal structure not posteriorly projected
in distal caudal vertebrae; ridge on ventromedial margin of scapular blade (also
in Ampelosaurus); anterolateral sternal process.
(after Company et al., 2009) coracoid foramen located close to scapular border
and dorsal margin of coracoid.
(after Diez Diaz et al., 2011) presence of foramen in distal surface of basal
tuber; absence of median subcondylar foramina in basioccipital, below occipital
condyle and between basal tubera (may be ontogenetic).
Other diagnoses- Sanz et al. (1999) cited "restricted articular
condyle in distal caudal vertebrae" as diagnostic for Lirainosaurus,
but Diez Diaz et al. (2013a) noted this is also present in several other titanosaurs
(e.g. Paludititan, Rocasaurus, Saltasaurus). Similarly,
the "sagittal groove on posterior articular surface of distal caudal vertebrae"
used by Sanz et al. was said by Diez Diaz et al. to be present other titanosaurs
(e.g. Magyarosaurus) and vary within Lirainosaurus as well.
Comments- The type material was discovered in 1984. After a preliminary
description by Sanz et al. (1999), the teeth were described by Diez Diaz et
al. (2012), the braincases by Diez Diaz et al. (2011), the axial skeleton by
Diez Diaz et al. (2013a) and the appendicular skeleton and armor by Diez Diaz
et al. (2013b).
Sanz et al. initially recovered Lirainosaurus as a eutitanosaur in a
very small analysis, similar to Upchurch et al. (2004) who found it as a lithostrotian
sister to an Alamosaurus+Pellegrinisaurus+saltasaurid clade. Calvo
et al. (2007) and Hocknull et al. (2009) recovered it as sister to saltasaurids,
while Gonzalez-Riga et al. (2009) found it sister to saltasaurids+epachthosaurines.
References- Sanz, Powell, Le Loueff, Martinez and Pereda-Suberbiola,
1999. Sauropod remains from the Upper Cretaceous of La�o (Northcentral
Spain). Titanosaur phylogenetic relationships. Estudios del Museo de Ciencias
Naturales de �lava. 14(N�mero Especial 1), 235-255.
Upchurch, Barrett and Dodson, 2004. Sauropoda. In Weishampel, Dodson and Osm�lska
(eds.). The Dinosauria. Second ed. University of California Press. 259-324.
Calvo, Porfiri, Gonz�lez-Riga and Kellner, 2007. A new Cretaceous terrestrial
ecosystem from Gondwana with the description of a new sauropod dinosaur. Anais
da Academia Brasileira de Ci�ncias. 79(3), 529-541.
Company, Pereda Suberbiola and Ruiz-Ome�aca, 2009. Nuevos restos f�siles
del dinosaurio Lirainosaurus (Sauropoda, Titanosauria) en el Cret�cico
Superior (Campaniano-Maastrichtiano) de la Pen�nsula Ib�rica.
Ameghiniana. 46(2), 391-405.
Gonz�lez Riga, Previtera and Pirrone, 2009. Malarguesaurus florenciae
gen. et sp. nov., a new titanosauriform (Dinosauria, Sauropoda) from the Upper
Cretaceous of Mendoza, Argentina. Cretaceous Research. 30, 135-148.
Hocknull, White, Tischler, Cook, Calleja, Sloan and Elliott, 2009. New Mid-Cretaceous
(Latest Albian) dinosaurs from Winton, Queensland, Australia. PLoS ONE. 4(7),
e6190.
Ortega and P�rez Garc�a, 2009. cf. Lirainosaurus sp. (Dinosauria:
Titanosauria) en el Cret�cico Superior de Saced�n (Guadalajara).
Geogaceta. 46, 87-90.
Company, 2011. Bone histology of the titanosaur Lirainosaurus astibiae
(Dinosauria: Sauropoda) from the Latest Cretaceous of Spain. Naturwissenschaften.
98, 67-78.
D�ez D�az, Pereda Suberbiola and Sanz, 2011. Braincase anatomy
of the titanosaurian sauropod Lirainosaurus astibiae from the Late Cretaceous
of the Iberian Peninsula. Acta Palaeontologica Polonica. 56(3), 521-533.
D�ez D�az, Pereda Suberbiola and Sanz, 2012. Juvenile and adult
teeth of the titanosaurian dinosaur Lirainosaurus (Sauropoda) from the
Late Cretaceous of Iberia. Geobios. 45, 265-274.
D�ez D�az, Pereda Suberbiola and Sanz, 2013a. The axial skeleton
of the titanosaur Lirainosaurus astibiae (Dinosauria: Sauropoda) from
the latest Cretaceous of Spain. Cretaceous Research. 43, 145-160.
D�ez D�az, Pereda Suberbiola and Sanz, 2013b. Appendicular skeleton
and dermal armour of the Late Cretaceous titanosaur Lirainosaurus astibia
(Dinosauria: Sauropoda) from Spain. Palaeontologia Electronica. 16(2), 19A
Atsinganosaurus Garcia,
Amico, Fournier, Thouand and Valentin, 2010
A. velauciensis Garcia, Amico, Fournier, Thouand and Valentin,
2010
Late Campanian, Late Cretaceous
La Bastide Neuve, France
Holotype- (VBN.93.01.a) (at least two adult individuals) four posterior
dorsal vertebrae (three incomplete)
Paratypes- ..(VBN.00.01) caudal vertebra
..(VBN.00.02) caudal vertebra
..(VBN.00.03) caudal vertebra
..(VBN.00.12) humerus
..(VBN.02.03) tooth
..(VBN.02.22) tooth
..(VBN.02.53) tooth
..(VBN.02.78a, b) scapula, coracoid
..(VBN.02.82) sacrum, ossified tendons
..(VBN.02.90) metacarpal
..(VBN.02.99) anterior-mid dorsal vertebra
..(VBN.02.109) tibia (530 mm)
..(VBN.02.110) mid caudal vertebra
..(VBN.93.03) caudal vertebra
....(VBN.93.04) caudal vertebra
....(VBN.93.05) caudal vertebra
....(VBN.93.06) caudal vertebra
....(VBN.93.07) caudal vertebra
....(VBN.93.08) caudal vertebra
..(VBN.93.10) metatarsal
..(VBN.93.11) scapula
..(VBN.93.12a-c) three anterior cervical vertebrae with fused partial cervical
ribs (one incomplete; 230-275 mm)
..(VBN.93.MHNA.99.21) tooth
..(VBN.93.MHNA.99.32) caudal vertebra
..(VBN.93.MHNA.99.33) distal caudal vertebra
..(VBN.93.MHNA.99.34) caudal vertebra
..(VBN.93.MHNA.99.52) humerus
Diagnosis- (after Garcia et al., 2010) teeth with cylindrical root and
slightly spatulate crown; anterior cervical vertebrae without marked pleurocoels
(also in Malawisaurus); posterior dorsal vertebrae with accessory lamina
connecting posterior centrodiapophyseal and postzygodiapophyseal laminae (also
in Malawisaurus); absence of hyposphene-hypantrum articulation in posterior
dorsal vertebrae; sacrum with five opisthocoelous fused vertebrae; sacral pleurocoels
present only on first vertebra; strongly procoelous proximal caudal vertebrae;
amphiplatyan to slightly opisthocoelous distal caudal vertebrae (also in Malawisaurus);
dorsal edge of scapular blade perpendicular to proximal expansion of scapula;
coracoid foramen equidistant from dorsal and ventral margins of coracoid; distal
end of humerus very slightly transversely expanded.
Comments- At least some of the specimens were found in 1993. The material
was originally described in Thouand's (2004) thesis before being officially
described and named by Garcia et al. (2010). Tortosa (2006) recovered the then-unnamed
taxon as a lithostrotian in a phylogenetic analysis. Garcia et al. concluded
Atsinganosaurus as a lithostrotian closest to Malawisaurus based
on shared vertebral characters (see diagnosis).
References- Thouand, 2004. Biodiversit� reptilienne et pal�oenvironnement
d'un gisement campanien du bassin d'Aix-en-Provence: La Bastide Neuve (Velaux).
MS thesis. Universit� de Poitiers. [pp?]
Tortosa, 2006. Diversit� des Titanosauria du Cr�tac� terminal
europ�en: Cadre chronologique, phylog�nie et pal�obiog�ographie.
MS thesis. University of Monpellier II. [pp?]
Garcia, Amico, Fournier, Thouand and Valentin, 2010. A new titanosaur genus
(Dinosauria, Sauropoda) from the Late Cretaceous of southern France and its
paleobiogeographic implications. Bulletin de la Soci�t� g�ologique
de France. 181(3), 269-277.
Epachthosaurinae Salgado, 2003
Definition- (Epachthosaurus sciuttoi <- Saltasaurus loricatus)
(modified from Salgado, 2003)
Reference- Salgado, 2003. Should we abandon the name Titanosauridae?
Some comments on the taxonomy of titanosaurian sauropods (Dinosauria). Revista
Espa�ola de Paleontolog�a. 18(1), 15-21.
Epachthosaurus
Diamantinasauria Poropat, Kundr�t, Mannion, Upchurch, Tischler and Elliott, 2021
Definition- (Diamantinasaurus matildae <- Saltasaurus loricatus) (Poropat, Kundr�t, Mannion, Upchurch, Tischler and Elliott, 2021)
Reference- Poropat, Kundr�t,
Mannion, Upchurch, Tischler and Elliott, 2021. Second specimen
of the Late Cretaceous Australian sauropod dinosaur Diamantinasaurus matildae
provides new anatomical information on the skull and neck of early
titanosaurs. Zoological Journal of the Linnean Society. 192(2), 610-674.
Sarmientosaurus
Diamantinasaurus
Nemegtosauridae Upchurch, 1995
Definition- (Nemegtosaurus mongoliensis <- Saltasaurus loricatus)
(modified from Apesteguia, 2004)
Other definitions- (Nemegtosaurus mongoliensis <- Diplodocus
longus) (modified from Upchurch et al., 2004)
?= "Argyrosaurinae" Powell, 1986
?= Argyrosaurinae Powell, 1986 vide Bonaparte, 1987
= "Nemegtosaurinae" Yu, 1993
= Nemegtosaurinae Upchurch, 1995 vide Yu, 1993 vide Salgado and Calvo, 1997
Comments- Yu (1993) was the first to separate Nemegtosaurus and
Quaesitosaurus from dicraeosaurids and other taxa and name the clade,
which he called Nemegtosaurinae. Being a thesis, Yu's work is not available
for nomenclature under the ICZN. Upchurch (1995) first validly published a name
for this group, as the family Nemegtosauridae. The first valid publication of
the subfamily version was Salgado and Calvo (1997), who cited Yu's thesis as
the source. Because of the ICZN's principle of coordination though, Upchurch
(1995) is still the author of Nemegtosaurinae.
My addition of Argyrosaurus to Mannion et al.'s macronarian matrix resulted
in it grouping with supposed nemegtosaurids Rapetosaurus and Mongolosaurus.
Note Argyrosauridae has priority over Nemegtosauridae, though given the preliminary
placement of Argyrosaurus, this is not formalized here.
References- Powell, 1986. Revisi�n de los titanosaurios de Am�rica
del Sur. PhD Thesis. Universidad Nacional de Tucum�n. 340 pp.
Bonaparte, 1987. The Late Cretaceous fauna of Los Alamitos, Patagonia, Argentina.
Parts I to IX. Revista Museo Argentino de Ciencias Naturales "Bernandino
Rivadavia". Paleontolog�a. 3(3), 103-178.
Yu, 1993. Reconstruction of the skull of Diplodocus and the phylogeny of the
Diplodocidae (Dinosauria - Sauropoda). PhD thesis. University of Chicago. 143
pp.
Upchurch, 1995. The evolutionary history of sauropod dinosaurs. Philosophical
Transactions of the Royal Society of London B. 349, 365-390.
Salgado and Calvo, 1997. Evolution of titanosaurid sauropods. II: The cranial
evidence. Ameghiniana. 34(1), 33-48.
Apestegu�a, 2004. Bonitasaura salgadoi gen. et sp. nov.: A beaked
sauropod from the Late Cretaceous of Patagonia. Naturwissenschaften. 91(10),
493-497.
Upchurch, Barrett and Dodson, 2004. Sauropoda. in Weishampel, Dodson and Osmolska
(eds.). The Dinosauria (2nd edition). University of California Press, Berkeley.
259-322.
Argyrosaurus Lydekker, 1893
A. superbus Lydekker, 1893
Campanian-Maastrichtian, Late Cretaceous
Upper Bajo Barreal Formation of the Chubut Group, Chubut, Argentina
Holotype- (MLP 77-V-29-1) incomplete humerus (1.37 m), radius (858 mm),
incomplete ulna (965 mm), two carpals (lost), metacarpal I (480 mm), metacarpal
II (~513 mm), metacarpal III (~504 mm), metacarpal IV (420 mm), metacarpal V
(398 mm)
Diagnosis- (after Mannion and Otero, 2012) medial margin of humerus forms
transversely wide ridge that projects prominently anteriorly; midshaft of humerus
extremely compressed anteroposteriorly (transverse to anteroposterior width
ratio 2.6); transverse width of distal radius only slightly greater than midshaft
width (ratio 1.3); radius subtriangular in distal view; ossified carpals; extreme
elongation of metacarpals (longest metacarpal to radius length ratio 0.6); metacarpals
II and III longest.
Other diagnoses- Lydekker (1893) originally distinguished Argyrosaurus
from Saltasaurus based on- large size; proximal humerus more squared;
more strongly developed and shorter deltopectoral crest; "postaxial"
border of humeral shaft less curved.
Bonaparte and Gasparini (1979) diagnosed Argyrosaurus based on a robust
humerus and antebrachium, which was rejected by Mannion and Otero (2012).
Mannion and Otero dismiss Powell's (2003) diagnostic characters as common in
titanosaurs- huge-sized; stout humerus with broad proximal end that has straight
edge perpendicular to long axis; deltopectoral crest more prominent and projected
anteriorly and medially; ulna with extremely robust proximal end showing prominent
edges delimiting markedly concave facets; stout metacarpals approximately one-third
the length of humerus.
Comments- The holotype was found in 1888, and was originally a complete
skeleton but only the forelimb survived excavation (Salgado, 2007).
Traditionally assigned to Titanosauridae in precladistic works, Argyrosaurus
has been included in relatively few cladistic analyses as the material was only
recently redescribed (Mannion and Otero, 2012). Apesteguia (2004) recovered
the holotype as a non-saltasaurine titanosaurian (under an uncertain definition
of the latter). Contra Mannion and Otero, Curry Rogers (2005) did not recover
Argyrosaurus as sister to Paralititan, the tree in question being
an Adams consensus. Her actual strict consensus trees only placed the genus
in a huge polytomy of non-saltasaurine somphospondylans, as Curry Rogers never
attempted a posteriori deletion of taxa to resolve its relations further. Mannion
and Upchurch's (2011) reanalysis of her matrix recovered Argyrosaurus
as a somphospondylan (probably titanosauroid based on the position of Euhelopus
and Phuwiangosaurus) outside the epachthosaurine+eutitanosaur clade.
However, Curry Rogers did code for a scapula (possibly Elaltitan's),
making her OTU a composite. Coding Argyrosaurus for Mannion et al.'s
macronarian matrix resolves it as a nemegtosaurid within Lithostrotia.
Not Argyrosaurus- Lydekker (1893) initially referred an incomplete
femur (MLP 21) and two distal caudal centra (MLP 22) to the genus based on size,
but these are not comparable to the holotype and were identified as Titanosauriformes
indet. (Camarasauromorpha here) and Lithostrotia indet. respectively by Mannion
and Otero (2012). Huene (1929) referred a number of additional specimens to
Argyrosaurus, but Mannion and Otero rejected them all- a proximal caudal
centrum (DGM coll.) from Chubut and two mid-distal caudals (MACN 5205) from
Santa Cruz cannot be compared and are Lithostrotia indet., two femora and a
tibia from Chubut (FMNH 13018-13020) cannot also not be compared and are Titanosauroidea
indet., a femur (MLP 27) from Neuquen cannot be compared and is Camarasauromorpha
indet., a humerus (MACN 5017) probably from Neuquen lacks Argyrosaurus'
diagnostic medial ridge and is Titanosauriformes indet., and another humerus
(MACN coll.) from Entre Rios also lacks this ridge and is Titanosauria indet..
Huene (1929b) referred a fragmentary forelimb and rib fragment (MMAB coll.)
from the Palacio Formation of Uruguay to Argyrosaurus based on its size,
but this has not been verified and it is treated as Titanosauria indet. here
(Powell, 2003; Mannion and Otero, 2012). Finally, Powell (1986, 2003) referred
a partial skeleton (PVL 4628) from Chubut to Argyrosaurus ?superbus,
but this was redescribed by Mannion and Otero as the new taxon Elaltitan
lilloi.
References- Lydekker, 1893. The dinosaurs of Patagonia. Anales del Museo
de la Plata, Seccion de Paleontologia. 2, 1-14.
Huene, 1929a. Los Saurisquios y Ornitisquios del Cretaceo Argentino. Anales
del Museo de La Plata. 3, 1-196.
Huene, 1929b. Terrestrische Oberkreide in Uruguay. Zentralblatt fur Mineralogie,
Geologie und Palaontologie Abteilung B. 1929, 107-112.
Bonaparte and Gasparini, 1979. Los sauropodos de los grupos Neuquen y Chubut
y sus relaciones cronologicas. Actas V Congreso Geologico Argentino, Neuquen.
2, 393-406.
Powell, 1986. Revisi�n de los titanosaurios de Am�rica del Sur.
PhD Thesis. Universidad Nacional de Tucum�n. 340 pp.
Powell, 2003. Revision of South American titanosaurid dinosaurs: Palaeobiological,
palaeobiogeographical and phylogenetic aspects. Records of the Queen Victoria
Museum. 111, 173 pp.
Apesteguia, 2004. An approach to the phylogenetic position of Argyrosaurus
(Sauropoda, Titanosauria). Ameghiniana. 41, 34A.
Curry Rogers, 2005. Titanosauria: Aphylogenetic overview. In Curry Rogers and
Wilson (eds.). The Sauropods: Evolution and Paleobiology. University of California
Press. 50-103.
Salgado, 2007. Patagonia and the study of its Mesozoic reptiles - a brief history.
In Gasparini, Salgado and Coria (eds.). Patagonian Mesozoic Reptiles. Indiana
University Press. 1-28.
Mannion and Upchurch, 2011. A re-evaluation of the 'mid- Cretaceous sauropod
hiatus' and the impact of uneven sampling of the fossil record on patterns of
regional dinosaur extinction. Palaeogeography, Palaeoclimatology, Palaeoecology.
299, 529-540.
Mannion and Otero, 2012. A reappraisal of the Late Cretaceous Argentinean sauropod
dinosaur Argyrosaurus superbus, with a description of a new titanosaur
genus. Journal of Vertebrate Paleontology. 32(3), 614-638.
Erketu
Nemegtosaurus
Rapetosaurus
Tapuiasaurus Zaher, Pol, Carvalho,
Nascimento, Riccomini, Larson, Juarez-Valieri, Pires-Domingues, da Silva and
de Almeida Campos, 2011
T. macedoi Zaher, Pol, Carvalho, Nascimento, Riccomini, Larson,
Juarez-Valieri, Pires-Domingues, da Silva and de Almeida Campos, 2011
Aptian, Early Cretaceous
Quirico Formation, Brazil
Holotype- (MZSP-PV 807) incomplete skull, sclerotic ossicles, mandibles,
hyoids (176, 173 mm), atlas, axis, five cervical vertebrae, five dorsal vertebrae,
dorsal ribs, incomplete coracoid, incomplete sternal plate, humerus, radius,
ulna, metacarpals, incomplete femora, fibula, incomplete pes
Diagnosis- (after Zaher et al., 2011) hook-shaped posteroventral quadratojugal
process; anterior jugal process tapering and forming most of ventral antorbital
fenestra margin; anterolateral tip of pterygoid contacts the medial surface
of ectopterygoid.
Comments- Zaher et al. (2011) added Tapuiasaurus to a version
of Wilson's sauropod analysis and recovered it as a nemegtosaurid closer to
Rapetosaurus than Nemegtosaurus.
Reference- Zaher, Pol, Carvalho, Nascimento, Riccomini, Larson, Juarez-Valieri,
Pires-Domingues, da Silva and de Almeida Campos, 2011. A complete skull of an
Early Cretaceous sauropod and the evolution of advanced titanosaurians. PLoS
ONE. 6(2), e16663.
Eutitanosauria Sanz, Powell, Le Loueff,
Martinez and Pereda-Suberbiola, 1999
Official Definition- (Saltasaurus loricatus <- Epachthosaurus sciuttoi)
(Navarro, Ghilardi, Aureliano, D�ez D�az, Bandeira, Cattaruzzi, Iori,
Martine, Carvalho, Anelli, Fernandes and Zaher, 2022; modified from Salgado, 2003; Registration Number 729)
Other definitions- (Saltasaurus loricatus + Argyrosaurus superbus
+ Lirainosaurus astibiae + "the Peiropolis titanosaur") (modified
from Sanz et al., 1999)
(Saltasaurus loricatus + Patagotitan mayorum) (Carballido, Otero, Mannion, Salgado and Perez Moreno, 2022)
= Saltasauroidea sensu Fran�a, J�lio, Riff, Hsiou and Langer, 2016
Definition- (Saltasaurus loricatus <- Nemegtosaurus mongoliensis) (modified)
Comments- Sanz et al. (1999) originally gave this clade a complex node-based
definition including "the Peiropolis titanosaur" as described by Powell
(1987) as an internal specifier. However, the latter material includes the holotypes
of Baurutitan britoi, Trigonosaurus pricei and two unnamed specimens
(MCT 1487-R and MCT 1489-R). Due to this and the use of the poorly resolved
Argyrosaurus in the definition, Salgado's (2003) later definition is
used, which is equivalent to the same clade in Sanz et al.'s topology except
for being stem-based.
References- Powell, 1987. Morfologia del esqueleto axial de los dinosaurios
titanos�uridos (Saurischia, Sauropoda) del Estado de Minas Gerais, Brasil.
Anais do X Congreso Brasiliero de Paleontologia, Rio de Janeiro. 155-171.
Sanz, Powell, Le Loueff, Martinez and Pereda-Suberbiola, 1999. Sauropod remains
from the Upper Cretaceous of La�o (Northcentral Spain). Titanosaur phylogenetic
relationships. Estudios del Museo de Ciencias Naturales de �lava. 14(N�mero
Especial 1), 235-255.
Salgado, 2003. Should we abandon the name Titanosauridae? Some comments on the
taxonomy of titanosaurian sauropods (Dinosauria). Revista Espa�ola de
Paleontolog�a. 18(1), 15-21.
Fran�a, J�lio, Riff, Hsiou and Langer, 2016. New lower jaw and teeth referred to Maxakalisaurus topai (Titanosauria: Aeolosaurini) and their implications for the phylogeny of titanosaurid sauropods. PeerJ. 4, e2054.
Carballido, Otero, Mannion, Salgado and Perez Moreno, 2022.
Titanosauria: A critical reappraisal of its systematics and the
relevance of the South American record. In Otero, Carballido and Pol
(eds.). South American Sauropodomorph Dinosaurs. Springer Earth System
Sciences. 269-298.
Navarro, Ghilardi, Aureliano, D�ez D�az, Bandeira, Cattaruzzi, Iori,
Martine, Carvalho, Anelli, Fernandes and Zaher, 2022. A new nanoid
titanosaur (Dinosauria: Sauropoda) from the Upper Cretaceous of Brazil.
Ameghiniana. 59(5), 317-354.
Alamosaurus
Elaltitan Mannion and Otero, 2012
E. lilloi Mannion and Otero, 2012
Middle Cenomanian-Turonian, Late Cretaceous
Lower Bajo Barreal Formation, Chubut, Argentina
Holotype- (MACN-CH 217) incomplete ~fifth dorsal vertebra (295 mm excl.
condyle), incomplete ~ninth dorsal vertebra (254 mm excl. condyle), incomplete
~tenth dorsal vertebra (237 mm excl. condyle), fragmentary ?first caudal vertebra
....(PVL 4628) rib fragments, ~second caudal vertebra (133 mm excl. condyle),
scapula (1.49 m), humerus (1.3 m), radius (730 mm), ulnae (827, 880 mm), pubis
(1.13 m), proximal femur, distal tibia, incomplete fibula, incomplete astragalus,
calcaneum (71 mm trans)
Diagnosis- (after Upchurch et al., 2004; as Argyrosaurus) dorsoventrally
tall neural arch restricted to anterior half of centrum (excluding condyle)
in proximal-most caudal vertebrae.
(after Mannion and Otero, 2012) mid-posterior dorsal spinopostzygapophyseal
laminae bifurcate into medial and lateral branches; astragalar ascending process
does not extend to posterior margin of astragalus; calcaneum present.
Other diagnoses- Novas (2009) listed "mild opisthocoely present
in posterior dorsal vertebrae" as diagnostic of Argyrosaurus, which
he assigned the specimen to. However, Mannion and Otero (2012) noted this is
only true of one of three preserved dorsals so rejected the character.
Comments- The holotype was discovered in 1976, and initially ascribed
to Antarctosaurus sp. by Bonaparte and Gasparini (1979) without justification
(though Powell 2003 claimed it was based on the iliofibularis tuber of the fibula).
Powell (1986, 2003) described the specimen as Argyrosaurus ?superbus
based on "general morphology, humeral proportions, and particularly the
ulna", but Mannion and Otero (2012) redescribed both PVL 4628 and Argyrosaurus,
finding both have autapomorphies and do not share obvious characters within
Titanosauria. They thus erected the taxon Elaltitan lilloi for PVL 4628,
referring it to Lithostrotia.
Note Bonaparte and Gasparini did not list the dorsals and listed another radius,
while Powell listed a third caudal, and neither publication listed the calcaneum.
As Mannion and Otero note, several elements are catalogued as MACN-CH 217 because
they were transferred to that repository.
Elaltitan is tentatively referred to Eutitanosauria here, as the prominent
muscle scar on the posterolateral surface of the deltopectoral crest noted by
Mannion and Otero is only present in this clade in Mannion et al.'s macronarian
matrix.
References- Bonaparte and Gasparini, 1979. Los sauropodos de los grupos
Neuquen y Chubut y sus relaciones cronologicas. Actas V Congreso Geologico Argentino,
Neuquen. 2, 393-406.
Powell, 1986. Revisi�n de los titanosaurios de Am�rica del Sur.
PhD Thesis. Universidad Nacional de Tucum�n. 340 pp.
Powell, 2003. Revision of South American titanosaurid dinosaurs: Palaeobiological,
palaeobiogeographical and phylogenetic aspects. Records of the Queen Victoria
Museum. 111, 173 pp.
Novas, 2009. The Age of Dinosaurs in South America. Indiana University Press.
452 pp.
Mannion and Otero, 2012. A reappraisal of the Late Cretaceous Argentinean sauropod
dinosaur Argyrosaurus superbus, with a description of a new titanosaur
genus. Journal of Vertebrate Paleontology. 32(3), 614-638.
Yongjinglong Li, Li, You and
Dodson, 2014
Y. datangi Li, Li, You and Dodson, 2014
Early Cretaceous
Upper Hekou Group, Gansu, China
Holotype- (GSGM ZH(08)-04) three teeth (81x20.6x19.8, 51x19x13, 28x17x11.5
mm), incomplete posterior cervical vertebra (~210 mm excl. condyle), first dorsal
vertebra (200 mm excl. condyle), three incomplete anterior dorsal vertebrae
(~160, 150, 173 mm excl. condyle), three incomplete mid dorsal vertebrae (158,
178, ~166 mm excl. condyle), fragmentary dorsal rib, scapulocoracoid (1.88 mm;
scap 1.55 m, cor 390 mm), radius (550 mm), ulna (590 mm)
Diagnosis- (after Li et al., 2014) long-crowned, spoon-shaped premaxillary
tooth; large, deep pleurocoels spanning entirety of lateral surfaces of cervical
and cervicodorsal centra; complex 'XI'- and 'IX'-shaped configuration of laminae
on left and right lateral surfaces respectively of articulated middle dorsal
vertebrae; low unbifurcated mid dorsal neural spine that along with postzygapophyses,
forms anteriorly pointing, triangular
plate; very long scapular blade with exceptionally straight anterior and posterior
edges.
Comments- The holotype was discovered in 2008. Li et al. (2014) recovered
it as a titanosauroid/lithostrotian, either outside Saltasauridae or inside
Saltasauridae as an opisthocoelicaudiine depending on the analysis used.
Reference- Li, Li, You and Dodson, 2014. A new titanosaurian sauropod
from the Hekou Group (Lower Cretaceous) of the Lanzhou-Minhe basin, Gansu Province,
China. PLoS ONE. 9(1), e85979.
Aeolosaurini Franco-Rosas, Salgado, Rosas
and Carvalho, 2004
Official Definition- (Aeolosaurus rionegrinus + Gondwanatitan faustoi) (Silva Junior, Martinelli, Lori, Marinho, Hechenleitner and Langer, 2022; Registration Number 433)
Other definition- (Aeolosaurus rionegrinus, Gondwanatitan faustoi
<- Opisthocoelicaudia skarzynskii, Saltasaurus loricatus) (Franco-Rosas,
Salgado, Rosas and Carvalho, 2004)
References- Franco-Rosas, Salgado, Rosas and Carvalho, 2004. Nuevos materiales
de titanosaurios (Sauropoda) en el Cret�cico Superior de Mato Grosso,
Brasil [New materials of titanosaurs (Sauropoda) from the Upper Cretaceous of
Mato Grosso, Brazil]. Revista Brasileira de Paleontologia. 7(3), 329-336.
Silva Junior, Martinelli, Lori, Marinho, Hechenleitner and Langer, 2022 (online 2021). Reassessment of Aeolosaurus maximus,
a titanosaur dinosaur from the Late Cretaceous of southeastern Brazil.
Historical Biology. 34(3), 403-411.
unnamed aeolosaurin (Almeida, Avilla and Candeiro, 2003)
Turonian-Santonian, Late Cretaceous
Adamantina Formation of the Bauru Group, Brazil
Material- (UFRJ-DG 270-R) caudal vertebra, chevron
Reference- Almeida, Avilla and Candeiro, 2003. Restos caudais associados
a Titanosauridae do Cretaceo Superior da Formacao Adamantina, Bacia Bauru, Municipio
do Prata - MG. XVIII Congresso Brasileiro de Paleontologia. Buletim de Resumos.
36-37.
Panamericansaurus Calvo
and Porfiri, 2010
P. schroederi Calvo and Porfiri, 2010
Late Campanian, Late Cretaceous
Allen Formation, Neuquen, Argentina
Holotype- (MUCPv-417) dorsal rib fragments, incomplete sacral vertebra,
five caudal vertebrae, two chevrons, humerus (1.23 m)
Diagnosis- (after Calvo and Porfiri, 2010) mid distal caudal prezygapophyses
directed anterodorsaly with the length equal to or shorter than the centrum;
mid distal caudal postzygapophyses placed in the middle of the centrum; spinoprezygapophyseal
lamina expanded dorsoventrally in mid caudals; hueral robusticity index less
than 0.40.
Comments- Discovered in 2003 and first assigned to Aeolosaurus.
Calvo and Porfiri (2010) assign it to Aeolosaurini.
References- Calvo and Porfiri, 2003. Primer registro de Aeolosaurus
en la provincia de Neuqu�n. Ameghiniana. 40(30), 34-60.
Calvo and Porfiri, 2010. Panamericansaurus schroederi gen. nov. sp. nov.
Un nuevo Sauropoda (Titanosauridae-Aeolosaurini) de la Provincia del Neuqu�n,
Cret�cico Superior de Patagonia, Argentina. Brazilian Geographical Journal:
Geosciences and Humanities. 1, 100-115.
Gondwanatitan
Aeolosaurus
Rinconsauria Calvo, Gonz�lez Riga
and Porfiri, 2007
Official Definition- (Rinconsaurus caudamirus + Muyelensaurus pecheni)
(Silva Junior, Martinelli, Lori, Marinho, Hechenleitner and Langer,
2022; modified from Calvo et al., 2007; Registration Number 434)
References- Calvo, Gonz�lez Riga and Porfiri, 2007. A new titanosaur
sauropod from the Late Cretaceous of Neuqu�n, Patagonia, Argentina. Arquivos
do Museu Nacional, Rio de Janeiro. 65(4), 485-504.
Silva Junior, Martinelli, Lori, Marinho, Hechenleitner and Langer, 2022 (online 2021). Reassessment of Aeolosaurus maximus,
a titanosaur dinosaur from the Late Cretaceous of southeastern Brazil.
Historical Biology. 34(3), 403-411.
Muyelensaurus
Rinconsaurus
Saltasauridae Powell, 1992 vide Sereno,
1998
Official Definition- (Opisthocoelicaudia skarzynskii + Saltasaurus loricatus)
(Navarro, Ghilardi, Aureliano, D�ez D�az, Bandeira, Cattaruzzi, Iori,
Martine, Carvalho, Anelli, Fernandes and Zaher, 2022; Wilson and
Upchurch, 2003; modified from Sereno, 1998; Registration Number 721)
?= Huabeisauridae Pang and Cheng, 2000
References- Powell, 1992. Osteologia de Saltasaurus loricatus
(Sauropoda - Titanosauridae) del Cret�cico Superior del noroeste Argentino. In Sanz and Buscalioni (eds.).
Los Dinosaurios y Su Entorno Biotico: Actas del Segundo Curso de Paleontologia
in Cuenca. Institutio "Juan de Valdes", Cuenca, Argentina. 165-230.
Sereno, 1998. A rationale for phylogenetic definitions, with application to
the higher-level taxonomy of Dinosauria. Neues Jahrbuch f�r Geologie und
Pal�ontologie Abhandlungen. 210, 41-83.
Pang and Cheng, 2000. A new family of sauropod dinosaur from the Upper Cretaceous
of Tianzhen, Shanxi province, China. Acta Geologica Sinica. 74(2), 117-125.
Wilson and Upchurch, 2003. A revision of Titanosaurus Lydekker (Dinosauria
- Sauropoda), the first dinosaur genus with a 'Gondwanan' distribution. Journal
of Systematic Palaeontology. 1(3), 125-160.
Navarro, Ghilardi, Aureliano, D�ez D�az, Bandeira, Cattaruzzi, Iori,
Martine, Carvalho, Anelli, Fernandes and Zaher, 2022. A new nanoid
titanosaur (Dinosauria: Sauropoda) from the Upper Cretaceous of Brazil.
Ameghiniana. 59(5), 317-354.
Opisthocoelicaudiinae McIntosh,
1990
Official Definition- (Opisthocoelicaudia skarzynskii <- Saltasaurus
loricatus)
(Navarro, Ghilardi, Aureliano, D�ez D�az, Bandeira, Cattaruzzi, Iori,
Martine, Carvalho, Anelli, Fernandes and Zaher, 2022; Wilson and
Upchurch, 2003; modified from Sereno, 1998; Registration Number 726)
References- McIntosh, 1990. Sauropoda. In Weishampel, Dodson and Osmolska
(eds). The Dinosauria, Berkeley: University of California Press. 345-401.
Sereno, 1998. A rationale for phylogenetic definitions, with application to
the higher-level taxonomy of Dinosauria. Neues Jahrbuch f�r Geologie und
Pal�ontologie Abhandlungen. 210, 41-83.
Wilson and Upchurch, 2003. A revision of Titanosaurus Lydekker (Dinosauria
- Sauropoda), the first dinosaur genus with a 'Gondwanan' distribution. Journal
of Systematic Palaeontology. 1(3), 125-160.
Navarro, Ghilardi, Aureliano, D�ez D�az, Bandeira, Cattaruzzi, Iori,
Martine, Carvalho, Anelli, Fernandes and Zaher, 2022. A new nanoid
titanosaur (Dinosauria: Sauropoda) from the Upper Cretaceous of Brazil.
Ameghiniana. 59(5), 317-354.
Liubangosaurus
Opisthocoelicaudia
Saltasaurinae Powell, 1992
Official Definition- (Saltasaurus loricatus <- Opisthocoelicaudia
skarzynskii)
(Navarro, Ghilardi, Aureliano, D�ez D�az, Bandeira, Cattaruzzi, Iori,
Martine, Carvalho, Anelli, Fernandes and Zaher, 2022; Wilson and
Upchurch, 2003; modified from Sereno, 1998; Registration Number 722)
Other definition- (Saltasaurus loricatus + Neuquensaurus australis)
(modified from
Salgado, Coria and Calvo, 1997)
= "Saltasaurinae" Powell, 1986
Comments- Note Saltasaurinae
was originally used in Powell's (1986) thesis, which is invalid as
theses are not considered published by the ICZN, then mentioned by
Powell (1987) but without a diagnosis so was still invalid under
Article 13.1.1.
References- Powell, 1986. Revisi�n de los titanosaurios de Am�rica
del Sur. PhD Thesis. Universidad Nacional de Tucum�n. 340 pp.
Powell,
1987. Hallazgo de un dinosaurio hadrosaurido (Ornithischia,
Ornithopoda) en la Formaci�n Allen (Cret�cico Superior) de Salitral
Moreno, Provincia de R�o Negro, Argentina. Congreso Geol�gico Argentino
10(3), 149-152.
Powell, 1992. Osteologia de Saltasaurus loricatus (Sauropoda - Titanosauridae)
del Cret�cico Superior del noroeste Argentino. In Sanz and Buscalioni (eds.). Los Dinosaurios y Su Entorno Biotico:
Actas del Segundo Curso de Paleontologia in Cuenca. Institutio "Juan de
Valdes", Cuenca, Argentina. 165-230.
Salgado, Coria and Calvo, 1997. Evolution of titanosaurid sauropods. I: Phylogenetic
analysis based on the postcranial evidence. Ameghiniana. 34(1), 3-32.
Sereno, 1998. A rationale for phylogenetic definitions, with application to
the higher-level taxonomy of Dinosauria. Neues Jahrbuch f�r Geologie und
Pal�ontologie Abhandlungen. 210, 41-83.
Wilson and Upchurch, 2003. A revision of Titanosaurus Lydekker (Dinosauria
- Sauropoda), the first dinosaur genus with a 'Gondwanan' distribution. Journal
of Systematic Palaeontology. 1(3), 125-160.
Navarro, Ghilardi, Aureliano, D�ez D�az, Bandeira, Cattaruzzi, Iori,
Martine, Carvalho, Anelli, Fernandes and Zaher, 2022. A new nanoid
titanosaur (Dinosauria: Sauropoda) from the Upper Cretaceous of Brazil.
Ameghiniana. 59(5), 317-354.
Normanniasaurus Le Loeuff,
Suteethorn and Buffetaut, 2013
N. genceyi Le Loeuff, Suteethorn and Buffetaut, 2013
Early-Middle Albian, Early Cretaceous
Poudingue Ferrugineux Formation, Seine-Maritime, France
Holotype- (MHNH-2013.2.1) two presacral prezygapophyses, three fused sacral
centra, incomplete proximal caudal vertebra (141 mm), incomplete mid caudal
vertebra (101 mm), partial scapula, two ilial fragments, ischia (one partial,
one fragmentary), femoral fragment, fibular fragment
Late Albian, Early Cretaceous
Poudingue Ferrugineux Formation?, Seine-Maritime, France
Paratype- ?(MHNR coll.) mid caudal centrum (101 mm) (Buffetaut, 1984)
Middle Albian, Early Cretaceous
Moru quarry, Oise, France
Referred- ?(MNHN coll.) ten mid caudal vertebrae (Lapparent, 1946)
Diagnosis- (after Le Loeuff et al., 2013) proximal caudal vertebrae with
antepostzygapophysial foramen; proximal caudal vertebrae with deep postspinal
and prespinal fossae; proximal caudal vertebrae with axially elongated neural
spine; dorsal projection of spinoprezygapophysial lamina in mid caudals.
Other diagnoses- Most of the supposedly diagnostic characters listed
by Le Loeuff et al. (2013) are either primitive for titanosauriforms (presacral
vertebrae with hyposphene-hypantrum articulation; middle caudal vertebrae amphicoelous;
ilium with anterolaterally expanded blade) or classic characters diagnosing
larger clades within it (internal texture of presacral vertebrae camellate;
proximal caudal vertebrae deeply procoelous; middle caudal vertebrae with anteriorly
placed neural arch). Whether the few remaining characters will prove diagnostic
once further comparisons are made is uncertain, especially as only one caudal
from each section of the tail is known, features (especially pneumatic ones)
are known to vary between individuals and along the axial column, and the authors
note their antepostzygapophyseal foramen is at least positionally homologous
to the interzygapophyseal fossa present in several other titanosaurs.
Comments- The holotype was discovered in 1990, initially mentioned in
Buffetaut (1995), then described officially by Le Loeuff et al. (2013) as a
new taxon. The authors believed Normanniasaurus was a non-eutitanosaurian
titanosaur. When added to Mannion et al.'s macronarian matrix, it emerges as
closer to Saltasaurus than Opisthocoelicaudia, though this has
yet to be tested in a matrix designed for lithostrotians.
Note the caudals originally described by Buffetaut (1984) and Lapparent (1946)
(said by Le Loeuff et al. to be "very probably" and "tentatively"
referrable to Normanniasaurus, respectively) are only referred based
on having anteriorly displaced neural arches and prominent chevron facets. As
the former is standard for titanosaurs and the latter isn't even proposed to
be diagnostic for the genus, correct referral is doubtful.
References- Lapparent, 1946. Pr�sence d'un dinosaurien sauropode
dans l'Albien du Pays de Bray. Annales de la Soci�t� g�ologique
du Nord. 66, 236-243.
Buffetaut, 1984. Une vert�bre de dinosaurien sauropode dans le Cr�tac�
du Cap de la H�ve (Normandie). Actes du Mus�um d’Histoire
naturelle de Rouen. 7, 215-221.
Buffetaut, 1995. Dinosaures de France. Editions du BRGM. 144 pp.
Le Loeuff, Suteethorn and Buffetaut, 2013. A new sauropod dinosaur from the
Albian of Le Havre (Normandy, France). Oryctos. 10, 23-30.
Bonatitan
Rocasaurus
Saltasaurini Powell, 1992 vide Coria and Salgado, 2005
Official Definition- (Saltasaurus loricatus + Neuquensaurus australis)
(Navarro, Ghilardi, Aureliano, D�ez D�az, Bandeira, Cattaruzzi, Iori,
Martine, Carvalho, Anelli, Fernandes and Zaher, 2022; Salgado and
Bonaparte, 2007; modified from Coria and Salgado, 2005; Registration
Number 725)
= Saltasaurinae sensu
Salgado, Coria and Calvo, 1997
Definition- (Saltasaurus loricatus + Neuquensaurus australis)
(modified)
References-
Powell, 1992. Osteologia de Saltasaurus loricatus (Sauropoda - Titanosauridae)
del Cret�cico Superior del noroeste Argentino. In Sanz and Buscalioni (eds.). Los Dinosaurios y Su Entorno Biotico:
Actas del Segundo Curso de Paleontologia in Cuenca. Institutio "Juan de
Valdes", Cuenca, Argentina. 165-230.
Salgado, Coria and Calvo, 1997. Evolution of titanosaurid sauropods. I: Phylogenetic
analysis based on the postcranial evidence. Ameghiniana. 34(1), 3-32.
Coria and Salgado, 2005. Mid-Cretaceous turnover of saurischian dinosaur communities:
Evidence from the Neuqu�n Basin. In Veiga, Spalletti, Howell and Schwarz
(eds.). The Neuquen Basin, Argentina: A Case Study in Sequence Stratigraphy
and Basin Dynamics. Geological Society, London: Special Publications. 252, 317-327.
Salgado and Bonaparte, 2007. Sauropodomorpha. In Gasparini, Salgado and Coria
(eds.). Patagonian Mesozoic Reptiles. Indiana University Press. 188-228.
Navarro, Ghilardi, Aureliano, D�ez D�az, Bandeira, Cattaruzzi, Iori,
Martine, Carvalho, Anelli, Fernandes and Zaher, 2022. A new nanoid
titanosaur (Dinosauria: Sauropoda) from the Upper Cretaceous of Brazil.
Ameghiniana. 59(5), 317-354.
Neuquensaurus
Saltasaurus