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 für Geologie und Paläontologie, 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 Neuquén y
Chubut y sus relaciones cronologicas. Actas V Congreso Geologico
Argentino, Neuquén. 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, Neuquén,
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 Neuquén y
Chubut y sus relaciones cronologicas. Actas V Congreso Geologico
Argentino, Neuquén. 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 über die Fossilen
Reptilien Russlands. IV. Theil. Ordnung Crocodilina Oppel.
Indeterminirte Fossile Reptilien. Mémoires de l'Académie Impériale des
Sciences de Saint-Pétersbourg. VIIe série 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?
Neuquén?, 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. Natürliche Schöpfungs-Geschichte.
Druck und Verlag von Georg Reimer. 832 pp.
Bonaparte and Coria, 1993. Un neuvo y gigantesco saurópodo titanosaurio
de la Formación Rio Limay (Albanio-Cenomaniano) de la Provincia del
Neuquén, 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
für Geologie und Paläontologie 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 für Geologie und
Paläontologie Abhandlungen. 285(1), 91-96.
Clasmodosaurus
References- Huene, 1929. Los saurisquios y ornitisquios del
Cretacéo 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., á
propósito de varias, coleccionadas por la Comisión Explorada á la
laguna Iberá, bajo los auspícios 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 maëstrichtiani 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), Neuquén, 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
Española de Paleontología. 18(1), 15-21.
Royo-Torres, 2009. El saurópodo de Peñarroya de Tastavins. Monograpfías
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 Paleontologýa y Bioestratigrafýa, 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 Ciências. 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), Neuquén, Argentina.
Ameghiniana. 28(3-4), 303-310.
Bonaparte and Coria, 1993. Un neuvo y gigantesco saurópodo titanosaurio
de la Formación Rio Limay (Albanio-Cenomaniano) de la Provincia del
Neuquén, 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
Española de Paleontología. 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 Ciências. 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 filogenéticas de Pleurocoelus
Marsh (Sauropoda). Jornadas Argentinas de Paleontología 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 <- Neuquénsaurus
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.
Argentinosaurus
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
González 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 González 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- González 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 Ciências. 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,
González Riga and Kellner, 2007
Official Definition- (Futalognkosaurus dukei + Mendozasaurus
neguyelap) (Navarro, Ghilardi, Aureliano, Díez Díaz, Bandeira,
Cattaruzzi, Iori,
Martine, Carvalho, Anelli, Fernandes and Zaher, 2022; Calvo et al.,
2007; Registration Number 730)
References- Calvo, Porfiri, González Riga and Kellner, 2007. A
new Cretaceous terrestrial ecosystem from Gondwana with the description
of a new sauropod dinosaur. Anais da Academia Brasileira de Ciências.
79(3), 529-541.
Navarro, Ghilardi, Aureliano, Díez Díaz, 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 Martínez, 2008. Nuevos
materiales de Titanosauridae (Lognkosauria?) de la Formación Bajo
Barreal, Cretácico 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 Neuquén
province (Argentina). Ameghiniana. 30, 215-218.
Valieri and Calvo, 2011. Revision of MUCPv 204, a Senonian basal
titanosaur from northern Patagonia. Dinosaurios y Paleontología desde
América Latina. Anales del III Congreso Latinoamericano de
Paleontología. Editorial de la Universidad Nacional de Cuyo, Neuquén.
143-152.
Lithostrotia Upchurch, Barrett and
Dodson, 2004 vide Wilson and Upchurch, 2003
Official Definition- (Malawisaurus dixeyi + Saltasaurus
loricatus)
(Navarro, Ghilardi, Aureliano, Díez Díaz, 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, Díez Díaz, 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.
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
França, Júlio, Riff, Hsiou and Langer, 2016
Official Definition- (Lirainosaurus
astibiae + Opisthocoelicaudia
skarzynskii + Saltasaurus loricatus) (Navarro, Ghilardi,
Aureliano, Díez Díaz, Bandeira, Cattaruzzi, Iori,
Martine, Carvalho, Anelli, Fernandes and Zaher, 2022; Registration
Number 719)
Other definitions- (Saltasaurus loricatus <- Nemegtosaurus mongoliensis)
(modified from França, Júlio, 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 Cretácico 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.
França, Júlio, 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, Díez Díaz, 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 Ciências. 83(1), 211-219.
Rubilar-Rogers and Gutstein, 2012. Los titanosaurios de Chile y su
contexto filogenético y biogeográfico. Boletín 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.
González-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
Española de Paleontología. 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
Díez Díaz, Garcia, Pereda Suberbiola, Jentgen-Ceschino, Stein,
Godefroit and Valentin, 2018
Official Definition- (Lirainosaurus astibiae + Ampelosaurus atacis)
(Navarro, Ghilardi, Aureliano, Díez Díaz, Bandeira, Cattaruzzi, Iori,
Martine, Carvalho, Anelli, Fernandes and Zaher, 2022; Díez Díaz,
Garcia, Pereda Suberbiola, Jentgen-Ceschino, Stein, Godefroit and
Valentin, 2018; Registration Number 727)
References- Díez Díaz, 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, Díez Díaz, 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
Laño (Northcentral Spain). Titanosaur phylogenetic relationships.
Estudios del Museo de Ciencias Naturales de Álava. 14(Número Especial
1), 235-255.
Upchurch, Barrett and Dodson, 2004. Sauropoda. In Weishampel, Dodson
and Osmólska (eds.). The Dinosauria. Second ed. University of
California Press. 259-324.
Calvo, Porfiri, González-Riga and Kellner, 2007. A new Cretaceous
terrestrial ecosystem from Gondwana with the description of a new
sauropod dinosaur. Anais da Academia Brasileira de Ciências. 79(3),
529-541.
Company, Pereda Suberbiola and Ruiz-Omeñaca, 2009. Nuevos restos
fósiles del dinosaurio Lirainosaurus (Sauropoda, Titanosauria)
en el Cretácico Superior (Campaniano-Maastrichtiano) de la Península
Ibérica. Ameghiniana. 46(2), 391-405.
González 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 Pérez García, 2009. cf. Lirainosaurus sp.
(Dinosauria: Titanosauria) en el Cretácico Superior de Sacedón
(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.
Díez Díaz, 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.
Díez Díaz, 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.
Díez Díaz, 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.
Díez Díaz, 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. Biodiversité reptilienne et
paléoenvironnement d'un gisement campanien du bassin d'Aix-en-Provence:
La Bastide Neuve (Velaux). MS thesis. Université de Poitiers. [pp?]
Tortosa, 2006. Diversité des Titanosauria du Crétacé terminal européen:
Cadre chronologique, phylogénie et paléobiogéographie. 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 Société
géologique 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 Española de Paleontología. 18(1), 15-21.
Epachthosaurus
Diamantinasauria
Poropat,
Kundrát,
Mannion, Upchurch, Tischler and Elliott, 2021
Definition- (Diamantinasaurus matildae <- Saltasaurus loricatus) (Poropat,
Kundrát,
Mannion, Upchurch, Tischler and Elliott, 2021)
Reference- Poropat, Kundrát,
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 Bonaparte, 1996
= "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. Revisión de los titanosaurios de
América del Sur. PhD Thesis. Universidad Nacional de Tucumán. 340 pp.
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.
Bonaparte, 1996. Cretaceous tetrapods of Argentina. Munchner
Geowissenschaftliche Abhandlungen. 30, 73-130.
Salgado and Calvo, 1997. Evolution of titanosaurid sauropods. II: The
cranial evidence. Ameghiniana. 34(1), 33-48.
Apesteguía, 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. Revisión de los titanosaurios de América del Sur. PhD
Thesis. Universidad Nacional de Tucumán. 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, Díez Díaz, 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 França, Júlio, 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 titanosáuridos (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 Laño (Northcentral Spain).
Titanosaur phylogenetic relationships. Estudios del Museo de Ciencias
Naturales de Álava. 14(Número Especial 1), 235-255.
Salgado, 2003. Should we abandon the name Titanosauridae? Some comments
on the taxonomy of titanosaurian sauropods (Dinosauria). Revista
Española de Paleontología. 18(1), 15-21.
França, Júlio, 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, Díez Díaz, 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. Revisión de los titanosaurios de América del Sur. PhD
Thesis. Universidad Nacional de Tucumán. 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.
Antarctosaurinae Bonaparte, 1996
Definition- (Antarctosaurus
wichmannianus <- Saltasaurus
loricatus) (Navarro, 2021)
= "Antarctosauridae" Olshevsky, 1978
= "Antarctosaurinae" Powell, 1986
= Antarctosauridae Bonaparte, 1996 vide Chatterjee, 2020
Comments- White (1973) wrote "The family or subfamily to include
the Cretaceous
peg-toothed sauropods with procoelus caudal vertebrae should be based
upon this genus [Antarctosaurus]
because it would then be capable of
definition." and "Antarctosaurus
should be used as the root for the
family name of Cretaceous peg-toothed sauropods, and Diplodocidae
(Marsh, 1884) for the entire group." without explicitly using
Antarctosauridae or Antarctosaurinae, and thus failed ICZN Article
11.7.1.2. ("A family-group name when first published must meet all the
following criteria. It must:" ... "be clearly used as a scientific name
to denote a suprageneric taxon...").
Olshevsky (1978) first used Antarctosauridae, stating "The family
Titanosauridae Lydekker 1885, the only family in the suborder whose
name is based on a nomen dubium, has been renamed Antarctosauridae,
following the recommendation in White 1973." This would qualify
as valid under ICZN Article 13.1.3- "To be available, every new name
published after 1930 must satisfy the provisions of Article 11 and must
... be proposed expressly as a new replacement name (nomen novum) for
an available name, whether required by any provision of the Code or
not." However Olshevsky also said in a "Note added in proof"-
"John S. McIntosh (pers. comm.) indicates the following: Renaming the
Titanosauridae the Antarctosauridae (Pages 2, 27) is not necessary.
Although Titanosaurus indicus
is based on meager material, it is diagnostic and not a nomen dubium.
Therefore, references to "antarctosaurid," etc., in this document
should read "titanosaurid," etc." This has been taken to indicate
ICZN Article 8.3- "If a work contains a statement to the effect that
all or any of the names or nomenclatural acts in it are disclaimed for
nomenclatural purposes, the disclaimed names or acts are not
available", making Antarctosauridae Olshevsky, 1978 unavailable.
Breithaupt (1985) listed "Family Antarctosauridae" including "Almosaurus
sp." (sic) as part of the "Lance Formation Nonmammalian Vertebrate
Fauna", but this was not valid as it violated ICZN Article 13.1- "To be
available, every new name published after 1930
must satisfy the provisions of Article 11 and must 13.1.1. be
accompanied by a description or definition that states in words
characters that are purported to differentiate the taxon, or 13.1.2. be
accompanied by a bibliographic reference to such a published
statement".
Powell (1986) erected "subfamilia Antarctosaurinae nov." in his thesis
to include only Antarctosaurus,
to differentiate it from titanosaurine, saltasaurine and argyrosaurine
titanosaurids. He retained Titanosauridae as the family name
because (translated) "it has priority, that the genus Titanosaurus
is based on pieces with fundamental although scarce diagnostic
characters, and that Titanosauridae is a name deeply rooted in
paleontological literature." It should be
noted that a diagnostic type genus is not necessary under the ICZN, so
White's and Olshevsky's reasons were unfounded, but also that theses do
not count as published under the ICZN (Article 9) so Powell's work
cannot be used as the basis for the name.
References at least including Olshevsky (1991) and Sereno (online,
2005) have credited Bonaparte, 1987 as the published debut of Powell's
Antarctosaurinae (e.g. Olshevsky lists it as "Antarctosaurinae J.
Powell, 1986 vide Bonaparte, 1987" and Sereno states "It was first used
in broader circulation by Bonaparte (1987)"), but the only reference
including Bonaparte from 1987 that uses Antarctosaurinae is Bonaparte
and Kielan-Jaworowska (1987). They write "In addition to genera
known since the times of Lydekker and von Huene (Titanosaurus, Argyrosaurus, Laplatasaurus and Antarctosaurus), Bonaparte and
Powell (1980) described from South America Saltasaurus, and Powell (1986)
described Neuquensaurus, Eolosaurus [sic] and Epachtosaurus
[sic]. This author distinguished four subfamilies within the
Titanosauridae: Titanosaurinae, Saltasaurinae, Antarctosaurinae and
Argyrosaurinae, based on vertebral and appendicular anatomy."
However, this does not count as a valid authorship of a family-level
name due to violating Article 13.1 like Breithaupt's attempt, as it
contained no definition or diagnosis and the reference to Powell 1986
is again not a published work under the ICZN.
The same seems to be true of other references to Antarctosaurinae until
Bonaparte (1996) lists "Subfamily Antarctosaurinae" and says "POWELL
(1986) defined this subfamily as follows (abbreviated): "Large to giant
titanosaurids. Skull provided with a wide muzzle. Basioccipital toberas
separated one another. Basipterygoid processes as thin bars.
Supratemporal fenestra opening up to in between orbit and central
portion of frontal. Basioccipital complex triangular, posteriorly with
a wide and concave surface. Foramina of internal carotid posterior to
the basipterygoid processes. Cylindrical, small teeth. Scapula with
narrower blade than in Saltasaurinae and Argyrosaurinae. Ischium with
narrow distal stem, somewhat wider distally. Pubic pedicel of ischium
well developed, but short. Tibia with robust ends. Fibula with lateral
process made of two parallel tuberosities following the bone axis."",
fulfilling ICZN Article 13.1.1 and being a translation of Powell's
diagnosis. Navarro (2021) proposed a phylogenetic definition in
his thesis- "All titanosaurians more closely related to Antarctosaurus wichmannianus Huene
1929 than to Saltasaurus loricatus
Bonaparte & Powell 1980", and also included Baalsaurus, Brasilotitan and Pellegrinisaurus although none of
these were used in his phylogenetic analysis. A diagnosis was
provided, however.
Chatterjee's (2020) usage may be the first valid one for
Antarctosauridae, although the family was implicitly erected along with
Antarctosaurinae in 1996. It reads- "Isisaurus
shows similarities to Antarctosaurus
and Argentinosaurus of
Argentina, Opisthocoelicoda
[sic] of Mongolia,
and Alamosaurus
of North America; these taxa are accommodated in the family
Antarctosauridae." This is based on the supposed absence of
osteoderms compared to titanosaurids, including Jainosaurus, Malawisaurus, Mendozasaurus, Vahiny and presumably Titanosaurus. Not only has
this division never been supported by a phylogenetic analysis, Alamosaurus osteoderms have been
reported (e.g. Carrano and D'Emic, 2015), Argentinosaurus and Antarctosaurus
are too fragmentary to allow the absence of evidence to be evidence of
absence, and Opisthocoelicaudiidae would have priority from 1990.
References- White, 1973. Catalogue of the genera of dinosaurs.
Annals of Carnegie Museum. 44, 117-155.
Olshevsky, 1978. The Archosaurian Taxa (excluding the Crocodylia).
Mesozoic Meanderings. 1, 50 pp.
Breithaupt, 1985. Nonmammalian vertebrate faunas from the Late
Cretaceous of Wyoming. Thirty-Sixth Annual Field Conference - Wyoming
Geological Association Guidebook. 159-175.
Powell, 1986. Revision de los titanosaurios de America del Sur. PhD
Thesis. Universidad Nacional de Tucuman. 340 pp.
Bonaparte
and Kielan-Jaworowska, 1987. Late Cretaceous dinosaur and mammal faunas
of Laurasia and Gondwana. In Currie and Koster (eds.). Fourth Symposium
on Mesozoic Terrestrial Ecosystems, Short Papers. 24-29.
Olshevsky, 1991. A revision of the parainfraclass Archosauria Cope,
1869, excluding the advanced Crocodylia. Mesozoic Meanderings. 2, 196
pp.
Bonaparte, 1996. Cretaceous tetrapods of Argentina. Munchner
Geowissenschaftliche Abhandlungen. 30, 73-130.
Sereno, online 2005. Stem Archosauria - TaxonSearch. http://www.taxonsearch.org/dev/file_home.php
[version 1.0, 2005 November 7]
Carrano and D'Emic, 2015. Osteoderms of the titanosaur sauropod
dinosaur Alamosaurus sanjuanensis
Gilmore, 1922. Journal of Vertebrate Paleontology. e901334.
Chatterjee, 2020. The age of dinosaurs in the Land of Gonds. In Prasad
and Patnaik (eds.). Biological Consequences of Plate Tectonics: New
Perspectives on Post-Gondwana Break-up - A Tribute to Ashok Sahni.
181-226.
Navarro, 2021. Postcranial osteology and phylogenetic relationships of
the Early Cretaceous titanosaur Tapuiasaurus
macedoi Zaher et al. 2011. Volume 1 - Main text. MS thesis.
Universidade de São Paulo. 123 pp.
Antarctosaurus
Narambuenatitan
Filippi, Garcia and Garrido, 2011
= "Narambuenatitan" Filippi, Garcia and Garrido, 2010 online
N. palomoi Filippi, Garcia and Garrido, 2011
= "Narambuenatitan palomoi" Filippi, Garcia and Garrido, 2010 online
Early-Middle Campanian, Late Cretaceous
Anacleto Formation of the Rio Colorado Subgroup, Neuquen, Argentina
Holotype- (MAU-Pv-N-425) (subadult) premaxilla, maxilla, braincase,
quadrates, incomplete anterior cervical vertebra, cervical rib
fragments, incomplete posterior dorsal vertebra (194 mm), three dorsal
rib fragments, eleven proximal caudal vertebrae (155, 140, 120 mm), six
mid caudal vertebrae (118, 108 mm), two caudal transverse processes,
two chevron fragments, partial coracoid, sternal plate (538 mm),
humerus (920 mm), ulna (605 mm), ilial fragment, pubes (770 mm),
proximal ischium, incomplete femur
Diagnosis- (after Filippi et al., 2011) great participation of
frontal in anterior border of supratemporal fossa; absence of
ornamentation on lateral borders of frontal; lamina joining each
parietal crest with anterior surface of supratemporal; dorsal
pleurocoel border defined by prominent convex bony edge in posterior
dorsal vertebrae; distal enlargement of posterior dorsal prespinal
lamina, matching with lateral enlargement of neural spine;
kidney-shaped posterior dorsal prezygapophysis; presence of numerous
pneumatic cavities located posterior to centroparapophyseal lamina and
centrodiapophyseal lamina in posterior dorsal vertebrae; mid caudal
vertebrae with anterodorsal border of neural spine anteriorly inclined;
middle caudal vertebra with slender prezygapophysis and neural arch.
Comments- The holotype was found in 2005-2006. The article
describing this taxon was released online in December 2010, but not
officially published until 2011. Found to be a lithostrotian more
derived than Malawisaurus, in a trichotomy with Epachthosaurus
and eutitanosaurs in Filippi et al.'s analysis.
Reference- Filippi, García and Garrido, 2011. A new titanosaur
sauropod dinosaur from the Upper Cretaceous of North Patagonia,
Argentina. Acta Palaeontologica Polonica. 56(3), 505-520.
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.
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 Cretácico Superior
de Mato Grosso, Brasil. 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 Neuquén. Ameghiniana. 40(30), 34-60.
Calvo and Porfiri, 2010. Panamericansaurus schroederi gen. nov.
sp. nov. Un nuevo Sauropoda (Titanosauridae-Aeolosaurini) de la
Provincia del Neuquén, Cretácico Superior de Patagonia, Argentina.
Brazilian Geographical Journal: Geosciences and Humanities. 1, 100-115.
Gondwanatitan
Aeolosaurus
Rinconsauria Calvo, González 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, González Riga and Porfiri, 2007. A new
titanosaur sauropod from the Late Cretaceous of Neuquén, 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
Bustingorrytitan
Simón and Salgado, 2023
= "Sauropodus" Simón vide Anonymous, 2001
B. shiva
Simón and Salgado, 2023
Late Cenomanian, Late Cretaceous
Bustingorry II site, Huincul Formation of the Rio Limay Subgroup of the
Neuquen Group, Neuquen, Argentina
Holotype- (MMCH-Pv 59/1-40)
Paratype- (MMCH-Pv 60/1-6)
Referred- (MMCH-Pv 61/1)
(MMCH-Pv 62/1-3)
Comments- Discovered in 1999, this was initially called
Sauropodus in an anonymous (2001) news article. The name may be a
mistranslation of "sauropod" however, as it does not appear in the
Spanish article (Chavez, 2001) and Simón only said it may be a new
taxon. It was later described officially as a new taxon of
lithostrotian Bustingorrytitan shiva
by Simón and Salgado (2023). This entry is very basic at the
moment and mostly meant as a placeholder as "Sauropodus" had a prior
entry.
References- Anonymous, 2001. Argentine find may be previously
unknown herbivore. Reuters. 2-8-2001.
Chavez, 2001. Rescatan los restos fósiles de un gran dinosaurio
herbívoro. La Nacion. 2-9-2001.
Simón, 2001. A giant sauropod from the Upper Cretaceous of El Chocon.
Ameghiniana. 38(4S), 19R.
Simón and Salgado, 2023. A new gigantic titanosaurian sauropod from the
early Late Cretaceous of Patagonia (Neuquén Province, Argentina). Acta
Palaeontologica Polonica. 68(4), 719-735.
Saltasauridae Powell, 1992 vide
Sereno, 1998
Official Definition- (Opisthocoelicaudia skarzynskii + Saltasaurus
loricatus)
(Navarro, Ghilardi, Aureliano, Díez Díaz, 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 Cretácico 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
für Geologie und Paläontologie 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, Díez Díaz, 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, Díez Díaz, 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
für Geologie und Paläontologie 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, Díez Díaz, 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, Díez Díaz, 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. Revisión de los titanosaurios de
América del Sur. PhD Thesis. Universidad Nacional de Tucumán. 340 pp.
Powell,
1987. Hallazgo de un dinosaurio hadrosaurido (Ornithischia,
Ornithopoda) en la Formación Allen (Cretácico Superior) de Salitral
Moreno, Provincia de Río Negro, Argentina. Congreso Geológico Argentino
10(3), 149-152.
Powell, 1992. Osteologia de Saltasaurus loricatus (Sauropoda -
Titanosauridae) del Cretácico 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
für Geologie und Paläontologie 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, Díez Díaz, 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. Présence d'un dinosaurien sauropode
dans l'Albien du Pays de Bray. Annales de la Société géologique du
Nord. 66, 236-243.
Buffetaut, 1984. Une vertèbre de dinosaurien sauropode dans le Crétacé
du Cap de la Hève (Normandie). Actes du Muséum 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, Díez Díaz, 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 Cretácico 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 Neuquén Basin. In Veiga,
Spalletti, Howell and Schwarz (eds.). The Neuquén 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, Díez Díaz, 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