Tetanurae Gauthier, 1986
Definition- (Passer domesticus <- Ceratosaurus nasicornis)
(Holtz et al., 2004; modified from Padian et al., 1999; modified from Gauthier,
1986)
Other definitions- (Allosaurus fragilis + Passer domesticus)
(modified from Novas, 1992)
(Passer domesticus <- Torvosaurus tanneri) (modified from Sereno,
1998)
(Vultur gryphus <- Ceratosaurus nasicornis) (Dal Sasso, Maganuco and Cau, 2018)
(Passer domesticus <- Ceratosaurus nasicornis, Carnotaurus
sastrei) (Sereno, in press)
= "Tetanurae" Gauthier, 1984
= Intertheropoda Paul, 1988
= Avipoda Novas, 1992
= Tetanurae sensu
Dal Sasso, Maganuco and Cau, 2018
Definition- (Vultur gryphus <- Ceratosaurus nasicornis)
= Tetanurae sensu Sereno, in press
Definition- (Passer domesticus <- Ceratosaurus nasicornis,
Carnotaurus sastrei)
Comments- Most recently, Carrano et al. (2012) have published an extensive
revision and phylogenetic analysis for non-coelurosaur members of this clade.
The addition of Carnotaurus as an external specifier by Sereno (in press)
seems counter-productive. Tetanurae was designed as a stem away from Ceratosaurus,
and abelisaurids were not explicitly discussed (having been named only a year
prior). In fact, technically, Indosaurus and Indosuchus were classified
as tetanurines by Gauthier (1986), since he lists them as carnosaurs. If abelisaurids
are megalosauroids (as in Paul, 1988), it shouldn't stop megalosauroids from
being tetanurines.
References- Gauthier, 1984. A cladistic analysis of the higher systematic categories of
the Diapsida. PhD thesis. University of California. 564 pp.
Benson, 2008. A new theropod phylogeny focussing on basal
tetanurans, and its implications for European 'megalosaurs' and Middle Jurassic
dinosaur endemism. Journal of Vertebrate Paleontology. 28(3), 51A.
Benson, 2009. Middle Jurassic theropods and the early evolution of tetanurans
(Dinosauria, Theropoda). Journal of Vertebrate Paleontology. 29(3), 62A.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Dal Sasso, Maganuco and Cau, 2018. The oldest ceratosaurian
(Dinosauria: Theropoda), from the Lower Jurassic of Italy, sheds light
on the evolution of the three-fingered hand of birds. PeerJ. 6:e5976.
Chuandongocoelurus
He, 1984
C. primitivus He, 1984
Bajocian, Middle Jurassic
Laoshangou Quurry, Golden Rooster Commune,
Xiashaximiao Formation, Sichuan, China
Holotype- (CCG 20010) (2.4 m, 12 kg; juvenile) anterior and posterior
dorsal vertebrae, three sacral vertebrae, sacral vertebrae, caudal vertebrae,
partial ilium, proximal pubis, proximal ischium, femora (201 mm), tibia (231
mm), distal tibia, proximal fibula, distal fibula, astragalus, calcaneum, incomplete
metatarsal II, phalanx II-1 (30 mm), metatarsal III (122 mm), metatarsal IV
(114 mm), phalanx IV-1 (16.5 mm), phalanx IV-2 (9 mm), phalanx IV-3 (10 mm),
phalanx IV-4 (12 mm), pedal ungual IV (18 mm)
Diagnosis- base of preacetabular process less than half acetabular height;
prominent anteriorly projecting process lateral to anterior trochanter on femur;
medial condyle much larger than lateral condyle on tibia.
Other diagnoses- Though Carrano et al. (2012) state the dorsal centra
and ilium are identical to Monolophosaurus, this is untrue.
Description- Although discovered in 1970 (Kuang, 2004) and described by He Xinlu back in 1984, Chuandongocoelurus
has been largely ignored in the literature. Little known is that He based the
taxon on two specimens, one much larger than the other. While both include dorsal
and caudal vertebrae, only those of the paratype were illustrated. Thus whether
the paratype belongs to Chuandongocoelurus is unknown, and it is here
discussed separately as an elaphrosaur. Carrano et al. (2012) were the first
to note this situation in print and also thought the paratype was similar to
Elaphrosaurus and thus unlikely to be Chuandongocoelurus. Comparing
the femoral length to Elaphrosaurus, the holotype can be estimated to
have measured 2.4 meters in length. Scaling from Paul's (1988) estimated mass
of Elaphrosaurus results in a weight of 12 kilograms. The unfused sacral
centra suggest it is young, which it would have to be if it is conspecific with
the paratype.
The holotype's vertebrae were not illustrated in the paper, but are visible
in a small unpublished photograph. The sacrals are moderately elongate and gently
concave ventrally, and apparently unfused. The other vertebrae are more fragmentary,
but some are more elongate and presumably distal caudals.
The pelvis is unfused and propubic, with the pubis about 27 degrees from the
vertical. The preacetabular process is broken, but extended past the pubic peduncle
and is dorsoventrally shallow, less than half of acetabular height. The dorsal
edge of the ilium is fairly straight and there is no vertical ridge or other
ornamentation laterally. The large pubic peduncle projects anteroventrally and
is slightly expanded distally. There is an extensive supracetabular shelf that
ends halfway through the ischial peduncle. The ischial peduncle is craniocaudally
42% as long as the pubic peduncle. The postacetabular process is broken off,
but could not have been very tall.
Only the proximal section of the pubis is preserved. The dorsal margin of an
obturator fenestra can be seen, although with the ventral margin incomplete,
it could be an obturator notch. There is no room for a pubic fenestra below
it.
Only the most proximal section of the ischium is preserved. Oddly, the ilial
peduncle is much wider than the pubic peduncle, contrasting with the peduncles
they attach to. No conclusion regarding obturator processes or flanges can be
reached.
The femur is hollow and sigmoid in medial view. The head is anteromedially directed
and declined ventrally. The anterior trochanter is a bit more massive than Dilophosaurus
in proximal view and is hooked medially. There seems to be a smaller process
directly lateral to the anterior trochanter. A trochanteric shelf is apparently
absent. The fourth trochanter is smaller than Dilophosaurus, but much
larger than the reduced process in Elaphrosaurus. Distally, the femur
shows a very slight extensor groove and a deep rounded flexor groove without
a cruciate ridge.
The tibia is quite elongate (15% longer than the femur) and slightly bowed laterally.
The proximal end is craniocaudally elongate, has a single cranial cnemial crest
and the lateral condyle is smaller than the medial condyle. The fibular crest
cannot be distinguished. Distally, the astragalus backed the tibia. In distal
view, the tibia is very anteroposteriorly narrow and roughly triangular.
The tibia, astragalus and calcaneum were unfused to each other. In the astragalus,
the medial condyle is much larger, there seems to be no transverse groove extending
across the condyles and the condyles are separated from the ascending process
by a groove or excavation. The extent of the ascending process is uncertain,
as it is broken proximally. However, it was obviously more prominent than ornithopods
(contra Norman, 1990) and a bit more extensive than Dilophosaurus, possibly
making the astragalus 83-93% as tall as wide. The anterior concavity of the
astragalus in distal view is not as developed as coelurosaurs. The calcaneum
was large and rectangular.
The metatarsus, though elongate (61% of femoral length), is not arctometatarsalian.
Indeed, the third metatarsal is wider proximally than distally. I am wary of
He's pedal reconstruction however, as metatarsal IV's distal end seems to be
in lateral/medial view, as might metatarsal III's. Also, metatarsal II's broken
distal end appears to be in posterior view. All this in a reconstruction of
anterior view. Maybe they were twisted due to post-burial deformation or illustrated
inaccurately. The proximal ends of metatarsals II and IV are flared outward.
In proximal view, metatarsal II is more tapered anteriorly, metatarsal III more
narrow, with a less expanded posterior end, and metatarsal IV is wider and more
wedge-shaped than in Elaphrosaurus. A phalanx, identified as II-1, is
shown backwards as its ginglymoid articulation is facing proximally. This phalanx
probably is II-1, as it is nearly identical in comparative size and shape to
that element in Elaphrosaurus. A series of apparently articulated phalanges
are identified as digit IV. IV-1 is the largest and IV-2 is the shortest. They
are all fairly similar in morphology, with lateral ligament pits and ginglymoid
distal articulations where can be seen. The fact that IV-2 is the shortest leads
to doubt they are articulated correctly, as this is never seen in theropods.
An ungual phalanx is also preserved. It is short and gently curved, with no
obvious flexor tubercle.
Relationships- Not many authors have attempted to classify Chuandongocoelurus.
He (1984) assigned it to the Coeluridae (as did Dong, 1992). Such an assignment is obviously based on small size,
as the Coeluridae was the diminutive equivalent of the Megalosauridae at the
time. In actuality, Chuandongocoelurus is much more primitive than Coelurus.
Norman (1990) referred it to Theropoda indet., while noting the primitively
broad scapula of the paratype, low ascending process and uncompressed third
metatarsal. The low ascending process is actually broken, though it would be
of ceratosauroid level if complete (after Welles and Long, 1974). The third
metatarsal is compressed in proximal view, leading to an hourglass shape seen
in averostrans. Holtz (1992) thought the low ilium, pubic obturator foramen,
very low astragalar ascending process and gracile metatarsus were most similar
to Elaphrosaurus, but suggested retaining it as Theropoda incertae sedis.
In 2001, I independently suggested it was a ceratosaur related to Elaphrosaurus
(based partially on vertebral characters, as I was unaware the hypodigm belonged
to two individuals), but this seems true only for the paratype. The first author
to include the taxon in a published analysis, Benson (2008, 2010) found Chuandongocoelurus
to be a non-megalosaurian megalosauroid, sister to Monolophosaurus. More
recently, Carrano et al. (2012) and Rauhut et al. (2012) found it to be a tetanurine
outside Orionides+Monolophosaurus, though when Carrano et al.'s matrix
is properly ordered it is in a trichotomy with Monolophosaurus
and Orionides. Stiegler (2019) notes "examination of the holotype
specimen (CUT 20010) confirms tetanuran synapomorphies recognized by
those studies as well as a lack of ceratosaurian synapomorphies" and
that "open neurocentral sutures on all preserved vertebrae suggest that
the holotypic Chuandongocoelurus
is represented by juvenile remains." Furthermore "Some of the
holotypic Chuandongocoelurus
elements were incorporated into a full skeletal reconstruction prior to
examination of the specimens in Chengdu, so some details of its anatomy
were obscured. The remainder of CUT 20010 was nearly completely
prepared and housed at the Chengdu University of Technology
Museum. Nathan Smith examined all of the materials prior to their
display and provided additional photos of both Chuandongocoelurus (CUT 20010) and CUT 20011."
References- He, 1984. The Vertebrate Fossils of Sichuan. Sichuan Scientific
and Technological Publishing House. 168 pp.
Norman, 1990. Problematic Theropoda: "Coelurosaurs". In Weishampel,
Dodson and Osmolska (eds.). The Dinosauria. University of California Press.
280-305.
Dong, 1992. Dinosaurian Faunas of China. China Ocean Press, Beijing. 188 pp.
Holtz, 1992. An unusual structure of the metatarsus of Theropoda (Archosauria:
Dinosauria: Saurischia) of the Cretaceous. PhD thesis. Yale University. 347
pp.
Glut, 1997. Dinosaurs - The Encyclopedia. McFarland Press, Jefferson, NC. 1076
pp.
Mortimer, DML 2001. https://web.archive.org/web/20200728132015/http://dml.cmnh.org/2001Jun/msg00957.html
Kuang, 2004. A new Sauropoda from Kaljajng dinosaurs fauna inMiddle
Jurassic beds of north-eastern Sichuan. In Sun (ed.). Collection of the
90th anniversary of Tianjin Museum of Natural History. Tianjin Science
and Technology Press. 40-46.
Benson, 2008. A new theropod phylogeny focussing on basal tetanurans, and its
implications for European 'megalosaurs' and Middle Jurassic dinosaur endemism.
Journal of Vertebrate Paleontology. 28(3), 51A.
Benson, 2010. A description of Megalosaurus bucklandii (Dinosauria: Theropoda)
from the Bathonian of the UK and the relationships of Middle Jurassic theropods.
Zoological Journal of the Linnean Society. 158(4), 882-935.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Rauhut, Foth, Tischlinger and Norell, 2012. Exceptionally preserved juvenile
megalosauroid theropod dinosaur with filamentous integument from the Late Jurassic
of Germany. Proceedings of the National Academy of Sciences. 109(29), 11746-11751.
Stiegler, 2019. Anatomy, systematics, and paleobiology of noasaurid
ceratosaurs from the Late Jurassic of China. PhD thesis, The George
Washington University. 693 pp.
Pandoravenator Rauhut and Pol, 2017
P. fernandezorum Rauhut and Pol, 2017
Oxfordian-Tithonian, Late Jurassic
Canadon Calcareo Formation, Chubut, Argentina
Holotype- (MPEF PV 1773-3) distal femur (~58 mm wide), proximal tibia
....(MPEF PV 1773-4) distal tibia (67 mm wide), distal fibula, astragalus (47 mm wide), calcaneum
....(MPEF PV 1773-5) tibial fragment, fibular fragment
....(MPEF PV 1773-6) proximal fibula
....(MPEF PV 1773-7) partial distal tarsal IV
....(MPEF PV 1773-8) proximal metatarsal IV
....(MPEF PV 1773-9) distal tarsal III, distal tarsal IV, proximal
metatarsal II, proximal metatarsal III, proximal metatarsal IV,
proximal metatarsal V
....(MPEF PV 1773-10) incomplete pedal ungual I, distal metatarsal II,
proximal phalanx II-1, distal metatarsal III, distal metatarsal IV,
proximal phalanx IV-1
....(MPEF PV 1773-11) distal metatarsal II
....(MPEF PV 1773-12) distal metatarsal III
....(MPEF PV 1773-13) pedal phalangeal fragment
....(MPEF PV 1773-14) pedal phalangeal fragment
....(MPEF PV 1773-15) pedal phalangeal fragment
....(MPEF PV 1773-16) pedal phalangeal fragment
....(MPEF PV 1773-17) pedal phalangeal fragment
....(MPEF PV 1773-18) proximal pedal ungual ?IV
....(MPEF PV 1773-27) ?fibular shaft
....(MPEF PV 1773-28) ?fibular shaft
Paratypes- ....(MPEF PV 1773-1) distal humerus (38 mm wide)
....(MPEF PV 1773-2) scapular fragment, coracoid fragment
....(MPEF PV 1773-19) incomplete distal caudal vertebra (~57 mm), partial distal caudal vertebra, incomplete chevron
....(MPEF PV 1773-20) two partial distal caudal vertebrae
....(MPEF PV 1773-21) fragments of two mid caudal vertebrae
....(MPEF PV 1773-36) distal tibia, partial astragalus
....(MPEF PV 1773-37) two pedal phalangeal fragments
Diagnosis- (after Rauhut and
Pol, 2017) distal caudal vertebrae postzygapophyses overhang
approximately half length of following vertebral centrum;
astragalus with anterior and distal lateral tubercles adjacent the
contact with calcaneum; anteriorly facing part of astragalar body that
is offset from the distal part by a transverse groove anteriorly;
ascending process of astragalus laminar but very low (about one third
the height of the astragalar body); ascending process triangular in
anterior view and inclined laterally; shaft but not proximal end of
metatarsal III strongly constricted between shafts of metatarsals II
and IV.
Comments- This was mentioned
early by Rauhut (2006) as "fragmentary remains of a tetanuran",
referring to Rauhut's (2002) SVP abstract. Yet theropod material
from these beds (then referred to as the upper sequence of the Ca�ad�n
Asfalto Formation) is not mentioned in that abstract. MPEF PV
1773-19 to 21 were found in 2002, MPEF PV 1773-36 and 37 were found in
2017, and everything else in January 2009. Rauhut and Pol (2017)
state "despite the disparate times of discovery, all elements were
found in the same erosional gully in an area of six to eight square
meters. Furthermore, the recovered elements are of matching size,
preservational quality, and, in the case of bones known from both left
and right side, of matching morphology (allowing for some deformation),
so that all of this material seems to represent a single
individual." They "restrict the holotype to the hindlimb elements
in order to avoid confusion in case future discoveries demonstrate that
remains of more than a single individual or taxon are present at this
locality."
Rauhut and Pol added Pandoravenator
to three phylogenetic analyses. In Carrano et al.'s tetanurine
matrix it emerged as a non-orionidan tetanurine, a basal megalosasuroid
or a basal carnosaur. In Smith et al.'s theropod matrix it
emerged as a non-orionidan tetanurine closer to birds than Chilesaurus but less than piatnitzkysaurids, and in Rauhut's theropod analysis it emerged as a tetanurine sister to Piatnityzkysaurus plus Orionides.
References- Rauhut, 2002.
Dinosaur evolution in the Jurassic: A South American perspective.
Journal of Vertebrate Paleontology. 22(3), 89A.
Rauhut, 2006. A brachiosaurid sauropod from the Late Jurassic Ca�ad�n
Calc�reo Formation of Chubut, Argentina. Fossil Record. 9, 226-237.
Rauhut and Pol, 2017. A theropod dinosaur from the Late Jurassic
Ca�ad�n Calc�reo Formation of central Patagonia, and the evolution of
the theropod tarsus. Ameghiniana. 54, 506-538.
"Monolophosauridae" Bakker, 1997
Comments- While Bakker (1997) referred to monolophosaurids in an abstract,
abstracts do not count for publication of new names in the ICZN (Article 9.10),
so this is a nomen nudum.
Reference- Bakker, 1997. Megalosaurian mid-life crisis: Diversity, co-evolution
and extinction at the Jurassic-Cretaceous boundary. Journal of Vertebrate Paleontology.
17(3), 30A.
Monolophosaurus Zhao and Currie,
1994
= "Jiangjunmiaosaurus" Anderson, 1987a
= "Monolophosaurus" Lambert, 1990
M. jiangi Zhao and Currie, 1994
= "Monolophosaurus jiangjunmiaoi" Dong, 1992
= "Monolophosaurus dongi" Grady, 1993
Late Callovian-Early Oxfordian, Middle-Late Jurassic
Jiangjunmiao, Lower Shishugou Formation, China
Holotype- (IVPP 84019) (5.71 m) skull (670 mm), mandible, atlas, axis
(61.8 mm), third cervical vertebra (67.1 mm), fourth cervical vertebra (69.3
mm), fifth cervical vertebra (70.6 mm), sixth cervical vertebra (69.6 mm), seventh
cervical vertebra (67.8 mm), eighth cervical vertebra (72.3 mm), ninth cervical
vertebra (68.4 mm), tenth cervical vertebra (67.3 mm), most cervical ribs, (dorsal
series 942.3 mm) first dorsal vertebra (60 mm), second dorsal vertebra (60.6
mm), third dorsal vertebra (60.9 mm), fourth dorsal vertebra (65.5 mm), fifth
dorsal vertebra (70 mm), sixth dorsal vertebra (71.2 mm), seventh dorsal vertebra
(74 mm), eighth dorsal vertebra (79.6 mm), ninth dorsal vertebra (79.5 mm),
tenth dorsal vertebra (78.8 mm), eleventh dorsal vertebra (80.9 mm), twelfth
dorsal vertebra (81.5 mm), thirteenth dorsal vertebra (79.8 mm), dorsal ribs
1-13, (sacrum 367.2 mm), first sacral vertebra (82.8 mm), second sacral vertebra
(75.4 mm), third sacral vertebra (67.4 mm), fourth sacral vertebra (69.5 mm),
fifth sacral vertebra (72.1 mm), first caudal vertebra (78 mm), second caudal
vertebra (76.1 mm), third caudal vertebra (74.2 mm), fourth caudal vertebra
(74.9 mm), fifth caudal vertebra (78.7 mm), sixth caudal vertebra (78.5 mm),
ilia (498 mm), pubes (495 mm), ischia (390 mm)
Diagnosis- (after Zhao and Currie, 1994) anteroposteriorly elongate premaxilla;
long, low external naris; antorbital sinuses in nasals confluent through openings
in base of crest; skull with midline crest formed by premaxillae, nasals, lacrimals
and frontals extending from above external naris to a point between orbits.
(after Brusatte et al., 2010) nasal process of premaxilla bifurcating posteriorly;
lateral surface of premaxilla with deep groove between subnarial foramen and
foramen on base of nasal process; nasal crest with straight dorsal margin nearly
parallel to alveolar margin of maxilla; two enlarged, subequal pneumatic fenestrae
in posterodorsal part of narial fossa; lacrimal with discrete, tab-like process
projecting dorsally above preorbital bar; rectangular frontals, much wider than
long (width/length=1.67).
Comments- The holotype was discovered in 1983 but not mentioned on paper
until Anderson (1987b) notes that the then just-discovered Sinraptor dongi
type "has been tentatively labelled a jiangjunmiaosaurus, a carnivore first
found in the Junggar Basin in 1983 by Zhao Xi Lin." The name "Jiangjunmaiosaurus"
was first used a month earlier in that journal, referencing the Sinraptor
type as an individual of it. Lambert (1990) is the first author to use the name
"Monolophosaurus", though its entry merely gives the etymology and
refers the reader to the "Jiangjunmiaosaurus" entry. Dong (1992; though
the latest included references are from 1990) mentions "Monolophosaurus
jiangjunmiaoi" as a new allosaurid and provides a photo of the mounted
skull and mandible. The reference given is "Zhao and Currie, in prep./press.
A new large theropod from Jurassic strata of Xinjiang.", which is an early
version of the eventual description. Currie and Zhao (1991) first report the
specimen in the scientific literature, in an SVP abstract that mentions the
cranial anatomy but does not attempt phylogenetic placement. The name "Monolophosaurus
dongi" was used by Grady in his book on the Chinese-Canadian Dinosaur Project,
which was written prior to the final description by Zhao and Currie (1994) officially
naming it Monolophosaurus jiangi. The etymology indicates Jiangjunmiao
translates to "General Jiang's Temple", so the species name still
reflects what the taxon was originally nicknamed for. Note that while volume
30(10) of the Canadian Journal of Earth Sciences lists its date as October 1993,
it was not published until February or March of 1994. Brusatte et al. (2010)
recently redescribed the skull, while Zhao et al. (2010) redescribed the postcrania.
Unfortunately, Zhao et al. state the specimen is "deeply embedded in hard
foam for travelling exhibition", so one side of it cannot be examined.
Relationships- Zhao and Currie (1994) placed Monolophosaurus as
a basal tetanurine ("megalosaurid-grade"), but also suggested it may
be a basal allosaurid (closer to Allosaurus than to Sinraptor).
Sereno et al. (1994) placed it within Allosauridae, while most authors have
found it resolves as a basal carnosaur (Holtz, 1996; Holtz, 2000; Holtz et al.,
2004; Novas et al., 2005; Yates, 2005). However, Holtz (1995) and Smith et al.
(2007) recovered Monolophosaurus as the sister taxon to Avetheropoda.
Rauhut's (2000) placement was similar, as the sister taxon of Afrovenator+Allosauroidea
within a Carnosauria containing megalosauroids. Benson (2008, 2010) found it
to be a basal megalosauroid related to Chuandongocoelurus, though his
updated analysis (Benson et al., 2010) placed the latter genus more ambiguously.
Most recently, Carrano et al. (2012) found Monolophosaurus to be a non-orionidan
tetanurine closer to Orionides than Chuandongocoelurus, though when the
matrix is properly ordered it is in a trichotomy with Chuandongocoelurus
and Orionides.
However, only one more step is needed to make it a non-megalosaurian megalosauroid,
so this placement is extremely tentative. In addition, it takes 4 more steps
to place it as a carnosaur and 5 more steps to place it sister to Avetheropoda,
so no alternative can be firmly rejected.
References- Anderson, 1987a. Chinese dinosaur dig strikes bonanza. New
Scientist. 1584, 25.
Anderson, 1987b. Chinese unearth a dinosaurs' graveyard. New Scientist. 1586,
28-29.
Lambert, 1990. The Dinosaur Data Book. New York, Avon Books. 320 pp.
Currie and Zhao, 1991. Two new theropods from the Jurassic of Xinjiang, People's
Republic of China. Journal of Vertebrate Paleontology. 11(3), 24A.
Dong, 1992. Dinosaurian Faunas of China. Ocean Press/Springer-Verlag, Beijing/Berlin.
188 pp.
Grady, 1993. The Dinosaur Project: The Story of the Greatest Dinosaur Expedition
Ever Mounted. Edmonton: Ex Terra Foundation. Toronto: Macfarlane Walter &
Ross. 61 pp.
Sereno, Wilson, Larsson, Dutheil and Sues, 1994. Early Cretaceous dinosaurs
from the Sahara. Science. 266, 267-271.
Zhao and Currie, 1994 (as 1993). A large crested theropod from the Jurassic of Xinjiang,
People's Republic of China. Canadian Journal of Earth Sciences. 30(10), 2027-2036.
Holtz, 1995. A new phylogeny of the Theropoda. Journal of Vertebrate Paleontology.
15(3), 35A.
Holtz, 1996. Phylogenetic analysis of the nonavian tetanurine dinosaurs (Saurischia:
Theropoda). Journal of Vertebrate Paleontology. 16(3), 42A.
Holtz. 2000. A new phylogeny of the carnivorous dinosaurs. GAIA. 15, 5-61.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria,
Saurischia). PhD thesis. University of Bristol. 440 pp.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria Second Edition. University of California Press. 71-110.
Novas, de Valais, Vickers-Rich and Rich, 2005. A large Cretaceous theropod from
Patagonia, Argentina, and the evolution of carcharodontosaurids. Naturwissenschaften.
92, 226-230.
Yates, 2005. A new theropod dinosaur from the Early Jurassic of South Africa
and its implications for the early evolution of theropods. Palaeontologia Africana.
41, 105-122.
Smith, Makovicky, Hammer and Currie, 2007. Osteology of Cryolophosaurus ellioti
(Dinosauria: Theropoda) from the Early Jurassic of Antarctica and implications
for early theropod evolution. Zoological Journal of the Linnean Society. 151,
377-421.
Benson, 2008. A new theropod phylogeny focussing on basal tetanurans, and its
implications for European 'megalosaurs' and Middle Jurassic dinosaur endemism.
Journal of Vertebrate Paleontology. 28(3), 51A.
Brusatte, 2008. The skull of Monolophosaurus jiangi
(Dinosauria: Theropoda) and its implications for early theropod
phylogeny and evolution. PhD thesis, University of Bristol. 94 pp.
Benson, 2010. A description of Megalosaurus bucklandii (Dinosauria: Theropoda)
from the Bathonian of the UK and the relationships of Middle Jurassic theropods.
Zoological Journal of the Linnean Society. 158(4), 882-935.
Benson, Brusatte and Carrano, 2010. A new clade of large-bodied predatory dinosaurs
(Theropoda: Allosauroidea) that survived to the latest Mesozoic. Naturwissenschaften.
97, 71-78.
Brusatte, Benson, Currie and Zhao, 2010. The skull of Monolophosaurus jiangi
(Dinosauria: Theropoda) and its implications for early theropod phylogeny and
evolution. Zoological Journal of the Linnean Society. 158(3), 573-607.
Zhao, Benson, Brusatte and Currie, 2010. The postcranial skeleton of Monolophosaurus
jiangi (Dinosauria: Theropoda) from the Middle Jurassic of Xinjiang, China,
and a review of Middle Jurassic Chinese theropods. Geological Magazine. 147(1),
13-27.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Allosaurus? tendagurensis
Janensch, 1925
= Antrodemus tendagurensis (Janensch, 1925) Huene, 1932
Late Kimmeridgian, Late Jurassic
Middle Dinosaur Member of Tendaguru Formation, Tanzania
Holotype- (MB R 3620; = 67 of Janensch) incomplete tibia (~910 mm)
Comments- Chure (2000) stated this specimen resembles abelisaurids because
it lacks a strongly curved cnemial crest or incisura tibialis and has no posterior
groove between the lateral and medial condyles, but differs from them in that
the astragalus is not fused to it and the fibular crest is more distally placed.
However, the depth of the incisura tibialis is uncertain since Rauhut (2011)
indicates the low lateral projection distally may be due to erosion. Furthermore,
the posterior groove is present and was merely not indicated by Janensch's dotted
line in the reconstruction. It also lacks the distally expanded cnemial crest
of ceratosaurs (Carrano et al., 2012). The distally placed fibular crest is
a tetanurine synapomorphy (which is fully compatible with a lack of tarsal fusion),
as noted by Rauhut. Within Tetanurae, the astragalar step excludes tendagurensis
from Coelurosauria, but its more detailed affinities are unknown. The broad
fibular crest is similar to Piatnitzkysaurus, Megalosaurus and
metriacanthosaurines
References- Janensch, 1925. Die Coelurosaurier und Theropoden der Tendaguru-Schichten
Deutsch-Ostafrikas. Palaeontographica. 1(supp. 7), 1-99.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte.
Monographien zur Geologie und Palaeontologie. 4(1), viii + 361 pp.
Chure, 2000. A new species of Allosaurus from the Morrison Formation
of Dinosaur National Monument (UT-CO) and a revision of the theropod family
Allosauridae. PhD thesis. Columbia University. 964 pp.
Rauhut, 2006. Theropod dinosaurs from the Late Jurassic of Tanzania and the
origin of Cretaceous Gondwanan theropod faunas. Journal of Vertebrate Paleontology.
26(3), 113A.
Rauhut, 2011. Theropod dinosaurs from the Late Jurassic of Tendaguru (Tanzania).
Palaeontology. 86, 195-239.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Fosterovenator Dalman, 2014
F. churei Dalman, 2014
Late Tithonian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Wyoming, US (Reed's Quarry
12)
Holotype- (YPM VP 058267A-C) partial tibiotarsus
Diagnosis- (after Dalman, 2014) proximal tibial condyles round; medial
condyle larger than lateral condyle; posterior intercondylar notch is deep and
V-shaped; lateral condyle strongly removed from medial condyle.
Other diagnoses- Dalman (2014) also includes characters from the paratype
supposed fibula (fibular shaft with uniform anteroposterior width throughout
most of length; distal fibula anteroposterior width 90% of the width of proximal
fibula), though this element is here excluded from the hypodigm.
Comments- Dalman (2014) described the theropod materials from Reed's
Quarry 12, including this tibiotarsus and a supposed fibula (YPM VP 058267D)
he made paratype of this new taxon. The latter element is unlike a theropod
fibula in that its shaft lacks significant distal taper, and the distal end
84% as wide anteroposteriorly as the proximal end. As the distal end has a triangular
outline similar to tetanurine tibiae, it may bne that element instead, in which
case the proximal end is crushed and/or incomplete. YPM VP 058267D is from a
larger individual than the holotype and as a fibula is incomparable or as a
tibia lacks shared characters, so should not be referred to the same taxon.
Dalman presents a confused and contradictory account of Fosterovenator's
relationships, officially assigning it to Ceratosauridae, then stating it "clearly
belongs to a basal tetanuran", then saying it is "more closely related
to Elaphrosaurus than to Ceratosaurus", and finally saying
it and Morrison cf. Elaphrosaurus tibia DMNH 36284 "most likely
represent the first occurrence of basal Late Jurassic abelisauroids in the Northern
Hemisphere" based on supposed similarity to Tendaguru tibiae MB.R.1750
and MB.R.1751. Of course a ceratosaurid cannot be an abelisauroid or a tetanurine,
but no evidence is presented in support of any of these positions. The tibia
seems to be tetanurine based on the widely separated fibular crest and lateral
condyle, unlike Ceratosaurus, Elaphrosaurus, DMNH 36284 or MB.R.1751.
The short and distally thick cnemial crest and posteriorly widely separated
proximal condyles are different from named Morrison genera (Ceratosaurus,
Torvosaurus, Saurophaganax, Allosaurus, Tanycolagreus)
as well as DMNH 36284 and NMMNH P-26093, so this may be a valid genus. If Dalman
is correct that the astragalar buttress is rounded and nearly vertical and that
the ascending process is likely taller than in Torvosaurus, Allosaurus
and Coelurus, this may be a coelurosaur. Interpreting morphology from
the low resolution photos with no indication of which surfaces are true vs.
broken is difficult however, and the description is less than ideal, so determining
Fosterovenator's exact relationships could be difficult.
Reference- Dalman, 2014. New data on small theropod dinosaurs from the
Upper Jurassic Morrison Formation of Como Bluff, Wyoming, USA. Volumina Jurassica.
12(2), 181-196.
Kaijiangosaurus He, 1984
K. lini He, 1984
Bajocian, Middle Jurassic
Laoshangou Quarry, Golden Rooster Commune,
Xiashaximiao Formation, Sichuan, China
Holotype- (CCG 20020) atlantal intercentrum (19 mm), third cervical vertebra
(69.5 mm), fifth cervical vertebra (91 mm), sixth cervical vertebra (78 mm),
seventh cervical vertebra (81 mm), eighth cervical vertebra (90 mm), ninth cervical
vertebra (74 mm)
Paratypes or Referred- (CCG coll.) jugal, nine teeth (35x14.7x8.2 mm;
29.7x13.3x7.4 mm; ?x16.3x7.3 mm; 21.6x9.4x5.9 mm), two dorsal centra, four proximal
caudal centra, three distal caudal vertebrae (~68 mm), incomplete scapula, coracoid (~111 mm proximodistally),
humerus (~170 mm), incomplete ulna (~108 mm), three metacarpals, three manual
unguals (~68 mm), proximal tibia, proximal fibula, metatarsal II (283 mm), partial
phalanx II-1, phalanx II-2, pedal ungual II, partial metatarsal III (315 mm),
partial phalanx III-2, phalanx III-3, pedal ungual III, incomplete metatarsal
IV (291 mm), phalanx IV-1 (~95 mm), phalanx IV-2 (~50 mm), phalanx IV-3 (~40
mm), phalanx IV-4 (~26 mm), pedal ungual IV (~56 mm) (He, 1984)
?(CCG coll.) femur (400 mm) (He, 1984)
Comments- The material was discovered in 1970 (Kuang, 2004) and comes from at least two individuals, the femur
being comparatively smaller than the other elements. The description has yet
to be translated from Chinese, leaving many details unknown to Western scientists
(such as whether the non-cervical elements are officially paratypes). Carrano
et al. (2012) suggested the cervical morphology indicated a basal tetanurine
or basal averostran. Entering it into Carrano et al.'s matrix results in a
position inside Tetanurae but excluded from Megalosauria and Avetheropoda.
References- He, 1984. [The Vertebrate Fossils of Sichuan]. Sichuan Scientific
and Technical Publishing House, Chengdu, Sichuan. 168 pp.
Kuang, 2004. A new Sauropoda from Kaljajng dinosaurs fauna inMiddle
Jurassic beds of north-eastern Sichuan. In Sun (ed.). Collection of the
90th anniversary of Tianjin Museum of Natural History. Tianjin Science
and Technology Press. 40-46.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Megalosaurus? "dapukaensis"
Zhao, 1985
?= Megalosaurus? "cachuensis" Weishampel, Barrett, Coria, Le
Loeuff, Xu, Zhao, Sahni, Gomani and Noto, 2004
Middle Jurassic
Middle Dapuka Group, Tibet, China
Material- (IVPP coll?)
Comments- Discovered in 1976 (An et al., 2021), this specimen was first reported by Zhao (1983) who while discussing
the evolution of dinosaurs in China noted "carnosaurs (Megalosaurus)"
in the Middle Jurassic. It might be surmised Zhao was referring to an undescribed
Chinese specimen of Megalosaurus, which is strengthened by the later
mention of a new Megalosaurus species from the same deposits as other
Middle Jurassic taxa Zhao mentions ("Ngexisaurus", "Lancanjiangosaurus",
"Microdontosaurus", "Changtusaurus"). As with other
new Tibetan taxa listed by Zhao (1983), it was probably supposed to be described
by Zhao in the published version of his doctoral dissertation "The Mesozoic
vertebrate remains of Xizang (Tibet), China", in the second Palaeontology
of Xizang volume. Yet this volume is only referenced by Zhao (1983; which was
submitted in September 1981) and seems never to have been printed, though the
previous volume was published by the IVPP in 1980 and the third by the NIGP
in 1981. Olshevsky (DML, 1999) notes the IVPP rejected the paper as unpublishable.
Zhao (1985) lists the new species Megalosaurus dapukaensis as being a
carnosaur from the Dabuka Group of Tibet. It is listed as an undescribed ?megalosaurid
from the Dapuka Group of Xizang Zizhiqu by Weishampel (1990). Zhang
and Li (1997) list this theropod as being from the Middle Dabuka Formation of
Dabuka, Qamdo County, Xizang. Weishampel et al. (2004) list it as ?Megalosaurus
cachuensis from the Dapuka Group of Xinjiang, Uygur Zizhiqu and questionably
refer it to Tetanurae (note this is not simply due to it being referred to Megalosaurus,
as other questionable Megalosaurus species in that section are only referred
to Theropoda). The species "cachuensis" is presumably an error, perhaps
caused by "Lancanjiangosaurus cachuensis" from the same locality.
It is listed as the megalosaurid Megalosaurus tibetensis Zhao gen. et
sp. nov. (MS) in Fang et al. (2006), suggesting that "cachuensis"
was indeed an error, and that Zhao's monograph was never published and is still
a manuscript. Of course Fang et al. are in error in attributing the genus Megalosaurus
to Zhao's manuscript. It may be referrable to Megalosaurus or Tetanurae,
but has not been described or figured so remains a nomen nudum.
References- Zhao, "1983" [unpublished]. The Mesozoic vertebrate
remains of Xizang (Tibet), China. The Series of the Scientific Expeditions to
the Qinghai-Xizang Plateau. Palaeontology of Xizang. 2, 1-200.
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.
Zhao and Cheng, 1985. The Qamdo-Simao Subregion. In Wang, Cheng and Wang (eds.).
The Jurassic System of China. Stratigraphy of China. 11, 174-179.
Weishampel, 1990. Dinosaurian distribution. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria. University of California Press. 63-139.
Zhang and Li, 1997. Mesozoic Dinosaur Localities in China and Their Stratigraphy.
In Wolberg, Sump and Rosenberg (eds.). Dinofest International, Proceedings of
a Symposium sponsered by Arizona State University. A Publication of The Academy
of Natural Sciences. 265-273.
Olshevsky, DML 1999. https://web.archive.org/web/20200720012936/http://dml.cmnh.org/1999Nov/msg00507.html
Weishampel, Barrett, Coria, Le Loeuff, Xu, Zhao, Sahni, Gomani and Noto, 2004.
Dinosaur Distribution. In Weishampel, Dodson and Osmolska (eds.). The Dinosauria
Second Edition. University of California Press. 517-606.
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.
An, Wang, Li, Wang and Wang, 2021. New discovery of Jurassic dinosaur
fossils in Chaya area, Qamdu district, Tibet. Geological Bulletin of
China. 40(1), 189-193.
"Megalosaurus" inexpectatus
del Corro, 1966
Albian-Cenomanian, Early Cretaceous-Late Cretaceous
Bayo Overo Member of the Cerro Barcino Formation, Chubut, Argentina
Holotype- (MACN 18.172) tooth (42.9 mm)
Paratypes- (MACN coll.) four teeth
Comments- del Corro (1974) notes that this taxon has wrinkled enamel
similar to Carcharodontosaurus saharicus, though Carrano et al. (2012)
specify it only has banding, as in Monolophosaurus, piatnitzkysaurids,
megalosaurines, carnosaurs and megaraptorans. Its age suggests it is an avetheropod.
References- del Corro, 1966. Un nuevo Dinosaurio carnivoro del Chubut.
Communicaciones del Museuo Argentino de Ciencias Naturales "Bernardino
Rivadavia" e Institutio Nacional de Investigacion de las Ciencias Naturales:
Paleontologia. 1(1), 1-4.
del Corro, 1974. Un nuevo megalosaurio (Carnosaurio) del Cr�tacico de
Chubut (Argentina) [A new megalosaur (Carnosauria) from the Cretaceous of Chubut
(Argentina)]. Communicaciones del Museo Argentino de Ciencias Naturales "Bernardino
Rivadavia" e Institutio Nacional de Investigacion de las Ciencias Naturales:
Paleontolog�a. 1(5), 37-44.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
"Megalosaurus" pannoniensis
Seeley, 1881
Early Campanian, Late Cretaceous
Gosau Formation, Austria
Holotype- (PIUW coll.) tooth (21x~10x6 mm), partial tooth
Santonian, Late Cretaceous
Csehbanya Formation, Hungary
Referred- ?(V.01.54, V.01.20, V.01.30, V.2003.04-08, V 2008.36.1-51)
27 teeth, 31 tooth fragments (Osi et al., 2010)
Comments- Osi et al. (2010) described numerous teeth from a slightly
earlier formation, which overlapped M. pannoniensis in a morphometric
study. They referred all of the teeth to basal Tetanurae (comparing them favorably
with Megalosaurus and Dubreuillosaurus) based on the "general
shape of the crown, the crown base length/width, the crown curvature, the morphology
of the carinae (especially the development pattern of the mesial carina)."
Carrano et al. (2012) also considered M. pannoniensis to be a tetanurine
based on slight enamel wrinkles (as in Monolophosaurus, piatnitzkysaurids,
megalosaurines, non-carcharodontosaurine carnosaurs and megaraptorans) and mesial
serrations which are absent basally, though they suggested the high DSDI (~1.75)
could indicate relationship to basal tyrannosauroids or dromaeosaurids. Its
age suggests it is an avetheropod.
Lapparent (1947) referred fragmentary cranial remains, manual phalanges and
a fibula to Megalosaurus pannoniensis, but these were later referred
to Dromaeosauridae by Allain and Taquet (2000). Three tooth fragments and three
unguals from the Late Campanian-Maastrichtian of Viso, Portugal were referred
to Megalosaurus pannoniensis by Lapparent and Zbyszewski (1957). They
are here listed as Averostra indet..
References- Seeley, 1881. On the reptile fauna of the Gosau Formation
preserved in the Geological Museum of the University of Vienna. Q. J. Geol.
Soc. London 37: 620-707.
Sauvage, 1898. Vert�br�s fossiles du Portugal: Contributions �
l'�tude des poissons et des reptiles du jurassique et du cr�tacique.
Direction des Travaux Geologiques du Portugal. Memoires. Comiss�o do
Servi�o Geol�gico de Portugal. 1-46.
Lapparent, 1947. Les dinosauriens du Cr�tac� sup�rieur
du midi de la France. M�moire de la Soci�t� g�ologique
de France. 56, 1-54.
Lapparent and Zbyszewski, 1957. Les dinosauriens du Portugal [The dinosaurs
of Portugal]. M�moires des Services G�ologiques du Portugal, nouvelle
s�rie. 2, 1-63.
Allain and Taquet, 2000. A new genus of Dromaeosauridae (Dinosauria, Theropoda)
from the Upper Cretaceous of France. Journal of Vertebrate Paleontology. 20(2),
404-407.
Osi, Apesteguia and Kowalewski, 2010. Non-avian theropod dinosaurs from the
early Late Cretaceous of Central Europe. Cretaceous Research. 31(3), 304-320.
"Mifunesaurus" Hisa, 1985
Middle Cenomanian, Late Cretaceous
Kabu Formation of the Mifune Group, Japan
Material- (YNUGI 10003; Mifune-ryu) (~6-7 m) maxillary tooth (72.7 mm)
Comments- This tooth was discovered in 1979, and described in 1984 by
Hawegawa and Murata. They referred it to Megalosauridae gen. et. sp. indet.,
and nicknamed it Mifune-ryu. Hisa (1985) later gave it the nomen nudum "Mifunesaurus"
in an illustrated booklet on dinosaurs. It was illustrated and described in
detail by Hasegawa et al. (1992), who referred it to Megalosauridae based on
similarity to Duriavenator, Gasosaurus and a Chinese specimen
of unknown stratigraphic origin probably incorrectly referred to Megalosaurus
bucklandii. They thought it was distinct from other named theropods in its
low basal width to crown length ratio (.17), but this is also seen in ceratosaurids,
for instance. It remains a nomen nudum because Hawegawa et al. did not name
the tooth, while Hisa's (1985) description was apparently deficient. While Chure
(2000) stated some of the teeth from the Kabu Formation have enamel wrinkles,
Chure et al. (1999) show he was referring to a tooth from the Jobu Formation
(MDM 341). Additional remains from the Jobu Formation of the Mifune Group (Tamara
et al., 1991- four teeth, tibia, fibula, metatarsals II and III) are sometimes
referred to "Mifunesaurus" as well, but although Chure (2000) stated
the teeth were similar, there are no reported synapomorphies which would allow
placing them in the same taxon.
The tooth crown is 72.7 mm tall, with a FABL of 22.5 mm and a basal width of
12.3 mm. It is recurved and lens shaped in section, with almost symmetrically
distributed carinae. There are 20 serrations per 5 mm on both mesial and distal
carinae. Blood grooves are present at least posteriorly, but enamel ridges are
absent. "Mifunesaurus"' tooth seems too thick to be a ceratosaurid,
and taller than in abelisaurids, so is probably from a non-maniraptoriform tetanurine,
such as a megalosauroid or carnosaur.
References- Hasegawa and Murata, 1984. First record of carnivorous dinosaur
from the Upper Cretaceous of Kyushu, Japan. Abstract of the Annual Meeting of
the Paleontological Society of Japan.
Hisa, 1985. Kyoryu zukan [Dinosaur Picture Book]. Shinchosa, Tokyo. 223 pp.
Dong, Hasegawa and Azuma, 1990. The Age of Dinosaurs in Japan and China. Fukui,
Japan: Fukui Prefectural Museum. 65 pp.
Tamara, Okazaki and Ikegami, 1991. Occurence of carnosaurian and herbivorous
dinosaurs from upper formation of Mifune Group, Japan. Memiors of the Faculty
of Education, Kumamoto University. 40, 31-45.
Hasegawa, Murata, Wasada and Manabe, 1992. The first carnosaur (Saurischia;
Theropoda) from Japan; A tooth from the Cenomanian Mifune Group of Kyushu. Science
Reports of the Yokohama National University, Series 2. 39, 41-49.
Chure, Manabe, Tanimoto and Tomida, 1999. An unusual theropod tooth from the
Mifune Group (Late Cenomanian to Early Turonian), Kumamoto, Japan. In Tomida,
Rich, and Vickers-Rich (eds.). Proceedings of the Second Gondwanan Dinosaur
Symposium. National Science Museum (Tokyo) Monographs. 15, 291-296.
Chure, 2000. A new species of Allosaurus from the Morrison Formation
of Dinosaur National Monument (Utah-Colorado) and a revision of the theropod
family Allosauridae. PhD thesis. Columbia University. 964 pp.
"Morosaurus" marchei
Sauvage, 1898
Kimmeridgian, Late Jurassic
Ourem, Alcobaca Formation?, Portugal
Holotype- (MG coll.) incomplete ~thirtieth-fortieth caudal
vertebra (100 mm)
Diagnosis- (proposed) indeterminate compared to Allosaurus and Poekilopleuron.
Comments- This is based on a caudal vertebra and a tooth (MG 16), described and illustrated by Sauvage (1898) as a new species of Morosaurus (now a junior synonym of Camarasaurus). Lapparent and Zbyszewski (1957) refer the vertebra to Megalosaurus
insignis stating "it is quite certain that it is not a Sauropod vertebra", but it cannot be compared to the type tooth of insignis. Madsen et al. (1995) stated "this specimen is now known to be a megalosaurid (JSM, personal observation)." Lapparent and Zbyszewski refer the tooth to their new taxon Apatosaurus alenquerensis, later assigned to the basal macronarian Lourinhasaurus. Mocho et al. (2017) have since referred it to ?Turiasauria indet..
The caudal has bnroken prezygapophyses and only the base of
postzygapophyses. Its centrum is 2.09 times longer than tall
posteriorly and 91% as tall as wide posteriorly. It is strongly
constricted ventrally and laterally, with a median ventral groove over
the middle 60%, prominent chevron facets and very low transverse
processes. Articular ends are amphicoelous and trapezoidally rounded
(wider dorsally). Only a low ridge is shown for the neural spine,
though it may be broken and the spine of e.g. Allosaurus
is only present anteriorly and posteriorly where the dorsal surface of
marchei may be broken away. Zygapophyseal bases are similar in height
and angle to those of Allosaurus' composite 28th caudal.
The theropod fauna in Late Jurassic Portugal is very similar to the
well sampled Morrison Formation, allowing a series of comparable taxa.
Ceratosaurus only has very distal caudals (~c33-39) as elongate which lacks transverse processes. Elaphrosaurus
has caudals with similar centrum dimensions with much larger transverse
processes, extensive neural spine and larger notch below the
postzygapophyses. Coelurus and Juratyrant lack a transverse process by that level of elongation. The most similar caudals are those of Allosaurus (28th in Madsen 1976) and Poekilopleuron (~38th in Allain and Chure), suggesting marchei is non-coelurosaur Tetanurae indet.
References- Sauvage, 1897-1898. Vertebres Fossiles du Portugual, Contributions
a l'etude des poissions et des reptiles du Jurassique et du Cretaceous. Direction
des Travaux Geologiques Portugal. 1-46.
Lapparent and Zbyszewski, 1957. Les dinosauriens du Portugal. M�moires des Services G�ologiques du Portugal, nouvelle
s�rie. 2, 1-63.
Madsen, McIntosh and Berman, 1995. Skull and atlas-axis complex of the Upper Jurassic sauropod Camarasaurus Cope (Reptilia: Saurischia). Bulletin of Carnegie Museum of Natural History. 31, 1-115.
Mocho, Royo-Torres,
Escaso, Malafaia, de Miguel Chaves, Narv�ez, P�rez-Garc�a, Pimentel, Silva and
Ortega, 2017. Upper Jurassic sauropod record in the Lusitanian Basin
(Portugal): Geographical and lithostratigraphical distribution.
Palaeontologia Electronica. 20.2.27A: 1-50.
Phaedrolosaurus Dong,
1973
P. ilikensis Dong, 1973
Valanginian-Albian, Early Cretaceous
Lianmugin Formation of Tugulu Group, Xinjiang, China
Lectotype- (IVPP V 4024-1) (~7 m) tooth (31 mm)
Paralectotype- ?(IVPP V 4024-3) proximal femur
Diagnosis- Provisionally indeterminate.
Comments- Dong originally based this taxon on a tooth, partial hindlimb
and proximal femur, all from different localities. Sues (1977) noted since the
diagnosis is based on dental characters, the tooth should be the lectotype.
Rauhut and Xu (2005) later made the hindlimb the holotype of a new species,
Xinjiangovenator parvus, which they found to be a relative of Bagaraatan.
Unfortunately, they neither mention the referred femur nor describe the tooth.
The tooth is about twice the size of Deinonychus, which would indicate
a theropod about seven meters long.
According to Dong's figure, the tooth's outline is near identical to Deinonychus.
It is compressed, recurved and serrated like most theropods. Serrations extend
from the base to the tip of the distal carina, with eighteen serrations per
five mm. The base of the mesial carina is smooth, but serrations are present
starting halfway up. It is said to be thicker than Deinonychus teeth.
The proximal femur is briefly mentioned, but not described or illustrated. It
was found at a separate site, so should not be regarded as Phaedrolosaurus.
Barsbold and Osmolska (1999) note that this femur has a wing-like anterior trochanter.
This excludes it from Maniraptora, which have either finger-like or fused anterior
trochanters. The lack of further information makes a more exact placement within Averostra impossible.
Both Molnar (pers. comm. to Glut 1989; in Glut, 1997) and Sues (1977) state
that Phaedrolosaurus appears dromaeosaurid. Barsbold and Osmolska (1999)
say the wing-like lesser trochantor is distinctly non-dromaeosaurid. Besides
these opinions, authors have generally just placed this genus in the Dromaeosauridae
without question. The tooth is said to be thicker than Deinonychus and
Velociraptor, but thickness varies with position in the tooth row, and
Bambiraptor and Atrociraptor also have generally thicker teeth
than the former two genera. Carrano et al.'s tetanurine analysis suggests only
tetanurine taxa have apically restricted mesial serrations in the Cretaceous,
but further studies of ceratosaurs may find that the state is more widely distributed.
Among tetanurines, only spinosaurids are definitely known in the Cretaceous,
suggesting Phaedrolosaurus is most likely an avetheropod.
References- Dong, 1973. Dinosaurs from Wuerho. In Reports of paleontological expedition to Sinkiang
(II), pterosaurian fauna from Wuerho, Sinkiang. Memoirs of the Institute of
Vertebrate Paleontology and Paleoanthropology Academia Sinica. 11, 45-52.
Sues, 1977. The skull of Velociraptor mongoliensis, a small Cretaceous
theropod dinosaur from Mongolia. Pal�ontologische Zeitschrift. 51, 173-184.
Glut, 1997. Dinosaurs - The Encyclopedia. McFarland Press, Jefferson, NC. 1076pp.
Barsbold and Osm�lska, 1999. The skull of Velociraptor (Theropoda)
from the Late Cretaceous of Mongolia. Acta Palaeontologica Polonica. 44, 189-219.
Rauhut and Xu, 2005. The small theropod dinosaurs Tugulusaurus and Phaedrolosaurus
from the Early Cretaceous of Xinjiang, China. Journal of Vertebrate Paleontology.
25(1), 107-118.
Prodeinodon? "tibetensis"
Zhang and Li, 1997
Early Cretaceous
Lura Formation, Tibet, China
Material- (IVPP coll.)
Comments- Discovered in 1976 (An et al., 2021), this specimen was first reported by Zhao (1983) who while discussing
the evolution of dinosaurs in China noted "the carnosaurs (Prodeinodon
Osborn) began to specialize" in the Early Cretaceous. It might be surmised
Zhao was referring to an undescribed Tibetan specimen of Prodeinodon,
which is strengthened by the later mention of a new Prodeinodon species
from the same deposits as other Early Cretaceous taxa Zhao mentions (?Microvenator,
Monkonosaurus, ?Asiatosaurus). As with other new Tibetan taxa listed
by Zhao (1983), it was probably supposed to be described by Zhao in the published
version of his doctoral dissertation "The Mesozoic vertebrate remains of
Xizang (Tibet), China", in the second Palaeontology of Xizang volume. Yet
this volume is only referenced by Zhao (1983; which was submitted in September
1981) and seems never to have been printed, though the previous volume was published
by the IVPP in 1980 and the third by the NIGP in 1981. Olshevsky (DML, 1999)
notes the IVPP rejected the paper as unpublishable. The species was first named
by Zhang and Li (1997) as Prodeinodon tibetensis from the Laoran Formation
of Laoran, Markam County, Tibet. It is near certainly the same specimen listed
as Tetanurae indet. by Weishampel et al. (2004) from the Lura Formation of Xizang
Zizhiqu. As the specimen has never been described or illustrated, it is a nomen
nudum. Given the age and occurance in China, it is likely to be a tetanurine
and as other Prodeinodon species are based on teeth this is likely for
this species as well. However, there is still no published evidence for this
or its generic referral.
References- Zhao, "1983" [unpublished]. The Mesozoic vertebrate
remains of Xizang (Tibet), China. The Series of the Scientific Expeditions to
the Qinghai-Xizang Plateau. Palaeontology of Xizang. 2, 1-200.
Zhao, 1983. Phylogeny and evolutionary stages of Dinosauria. Acta Palaeontologica
Polonica. 28(1-2), 295-306.
Zhang and Li, 1997. Mesozoic Dinosaur Localities in China and Their Stratigraphy.
In Wolberg, Sump and Rosenberg (eds.). Dinofest International, Proceedings of
a Symposium sponsered by Arizona State University. A Publication of The Academy
of Natural Sciences. 265-273.
Olshevsky, DML 1999. https://web.archive.org/web/20200720012936/http://dml.cmnh.org/1999Nov/msg00507.html
Weishampel, Barrett, Coria, Le Loeuff, Xu, Zhao, Sahni, Gomani and Noto, 2004.
Dinosaur Distribution. In Weishampel, Dodson and Osmolska (eds.). The Dinosauria
Second Edition. University of California Press. 517-606.
An, Wang, Li, Wang and Wang, 2021. New discovery of Jurassic dinosaur
fossils in Chaya area, Qamdu district, Tibet. Geological Bulletin of
China. 40(1), 189-193.
Spinostropheus Sereno, Wilson
and Conrad, 2004
S. gautieri (Lapparent, 1960) Sereno, Wilson and Conrad, 2004
= Elaphrosaurus gautieri Lapparent, 1960
Bathonian-Oxfordian, Middle-Late Jurassic
Tiouraren Formation of the Irhazer Group, Niger
Syntypes-
(MNHN 1961-28 in part; from In Tedrift) cervical neural arch, two dorsal vertebrae,
two sacral vertebrae (140 mm), partial caudal vertebra, three manual unguals
(40, 45, 60 mm), tibiae (700 mm), distal fibula, proximal metatarsal, four pedal
phalangeal fragments
?(MNHN 1961-28 in part; from In Tedrift) (many individuals) cervical vertebra (80 mm), two
anterior dorsal vertebrae (70, 80 mm), posterior dorsal vertebra (50 mm), four
dorsal fragments, three sacral fragments, three caudal vertebrae (80-85 mm),
two caudal fragments, partial humerus (200 mm), ulna (300 mm), distal pubis,
distal femur, incomplete tibia, incomplete fibula, proximal metatarsal, four
metatarsal fragments, partial pedal phalanx
Albian-Early Cenomanian, Early Cretaceous-Late Cretaceous
Tegama Group, Niger
Syntype- ?(MNHN 1961-28 in part; from In Abangarit) proximal metatarsal
Comments- Though originally identified as Early Cretaceous (Lapparent,
1960), the Tiouraren Formation has been reinterpreted as Bathonian-Oxfordian
(Rauhut and Lopez-Arbarello, 2009).
Lapparent referred this species to Elaphrosaurus without reason, distinguishing
it from his Elaphrosaurus iguidiensis by its larger size and "the
form of the vertebrae" and from E. bambergi
based on the supposedly stouter humerus. Lapparent's syntype series for
the species consists of numerous unassociated remains from In Tedrift
(not mentioned in the list of associated specimens), a proximal
metatarsal from In Abangarit, and an associated individual from In
Tedrift, all catalogued as MNHN 1961-28 (Rauhut and Carrano, 2016).
Sereno et al. (2004) listed only a cervical from the unassociated In
Tedrift remains as the holotype, but this would actually make it a
lectotype since it is a restricted portion of the syntypes. However,
Rauhut and Carrano point out that the ICZN states (Article 74.7.1) "to
be valid, a lectotype designation made after 1999 must ... employ the
term "lectotype" or an exact translation (e.g. "lectotypus", but not
"the type")", which Sereno et al. obviously did not do. Thus there is
no valid lectotype, and Rauhut and Carrano note that "the syntype
material includes at least the proximal ends of two right tibiae, ...
and thus clearly represents more than a single animal." Sereno et al.
described a new partial axial series MNN TIG6 from the Fako locality of
the Tiouraren Formation as a referred specimen of gautieri, which they place in a new genus Spinostropheus. While Sereno et
al. state the first bones of Spinostropheus
were "disarticulated vertebrae and two partial limb bones" described by
Lapparent, no justification for referring material of Lapparent's to
the taxon were given. Rauhut and Carrano conclude that "MNN TIG6 seems
to be markedly different from the original specimens of S. gautieri",
so that "the referral of this specimen to this taxon is not accepted
here (see below), pending a review of all of this material by Ronan
Allain and Paul Sereno (R. Allain, pers. comm. 2014)."
As for what Spinostropheus gautieri
is without Sereno et al.'s material, Rauhut and Carrano say
"examination of the lectotype [sic] and originally referred material
(Lapparent, 1960) of this taxon by one of us (O.R.) indicates that it
lacks ceratosaurian synapomorphies, and might represent a basal
tetanuran." This matches their differential diagnosis from Elaphrosaurus-
very slightly opisthocoelous cervical; single pair of anteriorly placed
pleurocentral foramina; shorter cervical centrum with length <2
times posterior height; humerus without bulbous head; humerus with
rounded distal condyles. It should be noted this is based on two
elements of the unassociated In Tedrift material, so that the ultimate
identity will depend on which element(s) are designated the
lectotype(s).
References- Lapparent, 1960. Les dinosauriens du "Continental intercalaire"
du Sahara central. Memoirs of the Geological Society of France. 88A, 1-57.
Sereno, Wilson and Conrad, 2004. New dinosaurs link southern landmasses in the
Mid-Cretaceous. Proceedings: Biological Sciences. 271(1546), 1325-1330.
Rauhut and Lopez-Arbarello, 2009. Considerations on the age of the Tiouaren
Formation (Iullemmeden Basin, Niger, Africa): Implications for Gondwanan Mesozoic
terrestrial vertebrate faunas. Palaeogeography, Palaeoclimatology, Palaeoecology.
271, 259-267.
Rauhut and Carrano, 2016. The theropod dinosaur Elaphrosaurus bambergi Janensch, 1920, from the Late Jurassic of Tendaguru, Tanzania. Zoological Journal of the Linnean Society. 178(3), 546-610.
Vectaerovenator Barker, Naish, Clarkin, Farrell, Hullmann, Lockyer, Schneider, Ward and Gostling, 2020
V. inopinatus Barker, Naish, Clarkin, Farrell, Hullmann, Lockyer, Schneider, Ward and Gostling, 2020
Late Aptian, Early Cretaceous
Member XIII, Ferruginous Sands Formation, Lower Greensand, England
Holotype- (IWCMS 2019.84) (subadult) incomplete ~first dorsal vertebra (52 mm), incomplete mid caudal vertebra (77.7 mm)
....(IWCMS 2020.400) incomplete ~fourth dorsal vertebra (52.3 mm)
....(IWCMS 2020.407) incomplete ~third cervical vertebra (54.7 mm), ~fourth cervical prezygapophysis, partial cervical rib
Diagnosis- (after Barker et
al., 2020) conspicuous lateral furrow between lateral margin of
postzygapophysis and prezygoepipophyseal lamina in cervical vertebrae;
convex spinoprezygapophyseal laminae in anterior cervical vertebra;
curved lamina bisecting postzygocentrodiapophyseal fossa; external
extension of air sac ventral to parapophysis of anterior dorsal
vertebra; complex partitioning of anteriormost dorsal pneumatic foramen
into several internal pneumatic features; anterior articular facet of
anterior dorsal vertebrae wider than centrum such that vertebra is
T-shaped in dorsal or ventral view; shallow fossae lateral to base of
postzygapophyses in mid caudal vertebrae.
Comments- The type material was discovered in March, May and September of 2019.
Barker et al. (2020) added Vectaerovenator to Cau's theropod analysis
and recovered it as a megaraptoran, but it could be constrained to
Megalosauroidea in two steps and Carnosauria in three steps. They
recommended it be placed as Tetanurae incertae sedis.
Reference- Barker, Naish,
Clarkin, Farrell, Hullmann, Lockyer, Schneider, Ward and Gostling, 2020. A highly pneumatic middle Cretaceous theropod from the
British Lower Greensand. Papers in Palaeontology. 6(4), 661-679.
undescribed Tetanurae (Ostrom, 1970)
Mid-Late Albian, Early Cretaceous
Himes Member, Cloverly Formation, Montana, US
Material- (MOR coll.) tooth (Maxwell, 1993)
(YPM 5397) two partial teeth (?x~29x? mm) (Ostrom, 1970)
Aptian-Early Albian, Early Cretaceous
Little Sheep Mudstone Member, Cloverly Formation, Wyoming, US
(YPM 4885) distal metatarsal III fragment (~50 mm wide) (Ostrom, 1970)
Mid-Late Albian, Early Cretaceous
Himes Member, Cloverly Formation, Wyoming, US
(YPM 5366) tooth (~7x~3x? mm), fragment (Ostrom, 1970)
(YPM 5369) tooth (27x~14x? mm) (Ostrom, 1970)
(YPM 5378) tooth (29x~12x? mm) (Ostrom, 1970)
?(YPM 5408) incomplete dorsal neural arch (Ostrom, 1970)
(YPM 5538) partial angular (Ostrom, 1970)
Comments- Ostrom (1970)
referred YPM 4885, 5369, 5378, 5397 and 5408 to Megalosauridae in his
systematic section, but only referred 4885 there in his locality
section, with the others only listed as Theropoda. Tooth YPM 5377
and supposed dorsal vertebra YPM 5285 were also listed as megalosaurid,
but have since been referred to Acrocanthosaurus by D'Emic et al. (2012). Note Ostrom (p. 69) mistakenly lists the largest tooth as YPM 5379, which is a small Deinonychus
tooth, while the actual number is YPM 5397 (as listed on page 218;
confirmed via the YPM online database). The teeth are said to
differ from Deinonychus in
having a low DSDI and are probably tetanurine
(carcharodontosaurid, megaraptoran, tyrannosauroid, etc.)
based on stratigraphy. The dorsal neural arch YPM 5408 is not
described by Ostrom, and that specimen number was accidentally used as
a Sauropelta specimen in the systematic section, but a photo on the YPM online database shows it does not resemble a Sauropelta
dorsal (postzygapophyses not connected and at low angle, neural spine
not backswept). However, it does seem to lack a hyposphene so may
be ornithischian (Tenontosaurus?).
The distal metatarsal III YPM 4885 was described as "slightly grooved
and not strongly ginglymoid", comparing closely to "Allosaurus, but not Deinonychus or Ornithomimus." This would also differ from the Acrocanthosaurus
holotype, which it is half the size of. Carrano et al. (2012)
said it "bears features common to many theropods, although it does seem
to differ from most coelurosaurs." The YPM online database shows
three elements are catalogued under YPM 5538, a proximal Deinonychus
fibula from locality 64-75 in Montana, a fibular shaft fragment labeled
"Therapoda indet. (deinonychus?)" without locality information but with
color and specimen number writing matching the other fibula, and the
partial angular listed by Ostrom under Megalosauridae. The
fragment is from the anterior or middle third, about 1.9 times larger
than Deinonychus YPM 5210, without the mediodorsal lip of that taxon or the dorsoventrally limited prearticular contact. Acrocanthosaurus has an even more limited prearticular contact and lacks the anterior expansion of YPM 5538 and Deinonychus. Allosaurus also lacks anterior expansion, which is present in Zuolong, Aorun, Erlikosaurus and Tyrannosaurus. Only the latter has extensive prearticular contact like YPM 5538, which is also missing in Murusraptor,
suggesting it could be from a tyrannosauroid. The latter clade is
known from the Cloverly Formation based on premaxillary tooth FMNH PR
2750.
YPM 5366 was collected in 1964 and was questionably referred to Microvenator
by Ostrom (1970), though Makovicky and Sues (1998) stated "this identification cannot be confirmed
because the holotype does not include any unambiguously associated teeth." It is ziphodont and recurved, so unlikely to
belong to that oviraptorosaurian taxon, but is probably tetanurine
(juvenile carcharodontosaurid or megaraptoran, tyrannosauroid, etc.)
based on stratigraphy.
Maxwell (1993) reported "one Microvenator tooth" from a microsite in Unit VII east of Bridger, Montana, found in 1992.
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, 234 pp.
Maxwell, 1993. Neonate dinosaur remains and dinosaur eggshell from the Lower
Cretaceous Cloverly Formation of Montana. Journal of Vertebrate Paleontology.
13(3), 48A.
Makovicky and Sues. 1998. Anatomy and phylogenetic relationships of the theropod
dinosaur Microvenator celer from the Lower Cretaceous of Montana. American
Museum Novitates. 3240, 1-27.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
D'Emic, Melstrom and Eddy, 2012. Paleobiology and geographic range of the large-bodied Cretaceous theropod dinosaur Acrocanthosaurus atokensis. Palaeogeography, Palaeoclimatology, Palaeoecology. 333-334, 13-23.
undescribed Tetanurae (Carrano, 1998)
Late Cretaceous?
Alberta, Canada?
Material- (RTMP 81.10.2) femur (317 mm)
(RTMP 81.10.2?) femur (180.7 mm)
(RTMP 91.40.16) femur (195.1 mm)
Comments- These are listed as Tetanurae in Carrano's (1998) table without
locality information, but if they are from Late Cretaceous Alberta, they are
likely maniraptoriforms or juvenile tyrannosaurids.
Reference- Carrano, 1998. The evolution of dinosaur locomotion: Functional
morphology, biomechanics, and modern analogs. PhD Thesis, The University of
Chicago. 424 pp.
unnamed tetanurine (Owen, 1842)
Early Toarcian, Early Jurassic
Jet Rock Formation, England
Material- (Ripley coll.; lost) incomplete anterior dorsal vertebra (87 mm)
Comments- This was originally referred to Streptospondylus cuvieri
by Owen (1842), though that taxon lived much later and no unique characters
are shared by the two now that more basal tetanurine anterior dorsals are known.
The parapophysis is smaller and more ventrally placed than in S. cuvieri,
though this may be positional variation. The opisthocoelous centrum is tetanurine,
and while Carrano et al. (2012) stated the single pleurocoel is, this is also
present in Cryolophosaurus and "Dilophosaurus" sinensis.
References- Owen, 1842. Report on British fossil reptiles. Report of
the British Association for the Advancement of Science. 11, 60-204.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
undescribed Tetanurae (Reynolds, 1939)
Early Bathonian, Middle Jurassic
Chipping Norton Limestone Formation (= Charlbury Formation), England
Material- (GSM 37523) dorsal vertebra (Welles and Pickering, 1999)
(SDM 44.17) proximal scapula (Reynolds, 1939)
(SDM 44.22) humerus (Reynolds, 1939)
Comments- Reynolds (1939) misidentified SDM 44.22 as an ischium. He referred
the SDM material to Megalosaurus sp., while Welles and Pickering (1999)
referred it to "Metriacanthosaurus" "reynoldsi".
However, I view their hypodigm of the latter species as mixing Megalosaurus
bucklandii material and a unique ilium from Oakham Quarry. Which of these
taxa, if either, this material belongs to is unknown.
References- Reynolds, 1939. A collection of reptile bones from the Oolite
near Stow-on-the-Wold, Gloucestershire. Geological Magazine. 76, 193-214.
Welles and Pickering, 1999. Megalosaurus bucklandii. Private publication
of Stephen Pickering, An extract from Archosauromorpha: Cladistics & Osteologies.
A Fractal Scaling in Dinosaurology Project. 119 pp.
undescribed Tetanurae (Welles and Pickering, 1999)
Early-Middle Bathonian, Middle Jurassic
Sharp's Hill Formation, England
Material- (OUM J13720) proximal caudal vertebra (Welles and Pickering,
1999)
(OUM J29799) proximal caudal vertebra (Welles and Pickering, 1999)
Comments- This material was referred to "Metriacanthosaurus"
"reynoldsi" by Welles and Pickering (1999), but I view their hypodigm
of the latter species as mixing Megalosaurus bucklandii material and
a unique ilium from Oakham Quarry. Which of these taxa, if either, this material
belongs to is unknown.
Reference- Welles and Pickering, 1999. Megalosaurus bucklandii.
Private publication of Stephen Pickering, An extract from Archosauromorpha:
Cladistics & Osteologies. A Fractal Scaling in Dinosaurology Project. 119
pp.
undescribed tetanurine (Naish and Martill, 2007)
Kimmeridgian, Late Jurassic
Kimmeridge Clay, England
Material- cervical vertebrae, dorsal vertebrae, sacral vertebrae, caudal
vertebrae, pelvic elements, hindlimb elements
Comments- This specimen was mentioned as a "peculiar, gracile tetanurine"
by Naish (online, 2006) and is due to be studied. Naish and Martill (2007) mention
it in print as pers. comm. from Powell.
References- Naish, online 2006. http://darrennaish.blogspot.com/2006/12/obscure-dinosaurs-of-kimmeridge-clay.html
Naish and Martill, 2007. Dinosaurs of Great Britain and the role of the Geological
Society of London in their discovery: Basal Dinosauria and Saurischia. Journal
of the Geological Society, London. 164, 493-510.
unnamed tetanurine (Naish, 1999)
Berriasian-Valanginian, Early Cretaceous
Hastings Beds, England
Material- (NHMUK R9385) partial tibia (~250 mm)
Comments- Naish (1999) described this as a non-coelurosaurian tetanurine.
Reference- Naish, 1999. Theropod dinosaur diversity and palaeobiology
in the Wealden Group (Early Cretaceous) of England: Evidence from a previously
undescribed tibia. Geologie en Mijnbouw. 78, 367-373.
unnamed Tetanurae (Malafaia, Ortega, Silva, Escaso and Dantas, 2008)
Late Kimmeridgian, Late Jurassic
Praia de Valmit�o, Praia da Amoreira-Porto Novo Formation, Portugal
Material- (SHN.036/30) lateral tooth (18.1x11x4.4 mm)
(SHN.036/31) lateral tooth (23.7x10x5.2 mm)
(SHN.036/32) lateral tooth (12.5x5.7x3.9 mm)
Comments-
These were found by a private collector associated with the Lusovenatorholotype
and donated to the SHN in
2008. Although initially referred to the skeleton by Malfaia et
al. (2008), Malafaia et al. (2017) later stated that although "they
have a morphology and size
compatible with the postcranial elements", they "do not preserve
any parts of the root, suggesting that they correspond to shed
teeth."
They thus excluded the teeth (SHN.03630-32) from the hypodigm and
described them as "compatible with lateral teeth of basal tetanurans."
References- Malafaia, Ortega, Silva, Escaso and Dantas, 2008. Un nuevo
ejemplar de Allosauroidea (Dinosauria: Tetanurae) del Jur�sico Superior
de Valmit�o (Lourinh�, Portugal). XXIV Jornadas de la Sociedad
Espa�ola de Paleontolog�a, abstracts. 148-149.
Malafaia, Mocho, Escaso and Ortega, 2017 (online 2016). A juvenile
allosauroid theropod (Dinosauria, Saurischia) from the Upper Jurassic
of Portugal. Historical Biology. 29, 654-676.
unnamed Tetanurae (Nessov, 1995)
Callovian, Middle Jurassic
Balabansai Svita, Kyrgyzstan
Material- (ZIN PH 7/42) tooth (31.5x17x11.5 mm)
(ZIN PH 8/42) tooth (31.2x11.9x10 mm)
(ZIN PH 9/42) tooth (26.8x12x6 mm)
(ZIN PH 10/42) tooth (30.1x12.3x11.9 mm)
(ZIN PH 11/42) tooth (24.2x10.9x6 mm)
(ZIN PH 12/42) tooth (21.1x8.7x6.7 mm)
(ZIN PH 13/42) tooth
(ZIN PH 14/42) tooth (18.7x7x5.1 mm)
(ZIN PH 15/42) tooth (13x6.6x3.7 mm)
(ZIN PH 16/42) tooth (13.2x6.7x4.3 mm)
(ZIN PH 17/42) tooth (?x6x3 mm)
(ZIN PH 18/42) tooth (9.2x3.3x3 mm)
(ZIN PH 19/42) tooth (17x8x4.3 mm)
(ZIN PH 20/42) tooth (10.2x6.8x3.6 mm)
(ZIN PH 26/42) tooth (44.4x17.5x8.9 mm)
(ZIN PH 27/42) tooth (?x21.8x13.4 mm)
(ZIN PH 28/42) tooth (17x9.7x3.8 mm)
(ZIN PH 29/42) tooth (4.9x3.5x2.2 mm)
Comments- Nessov (1995) identified these as cf. coelurid and megalosaurid,
while Averianov et al. (2005) stated all variation could be ontogenetic and
positional. They believed the teeth resembled dromaeosaurids primarily in the
basally unserrated mesial carina, but Carrano et al. (2012) noted this and other
characters were consistent with basal tetanurines too.
References- Nessov, 1995. Dinosaurs of nothern Eurasia: New data about
assemblages, ecology, and paleobiogeography. Institute for Scientific Research
on the Earth's Crust, St. Petersburg State University, St. Petersburg. 1-156.
Martin and Averianov, 2004. Middle Jurassic vertebrates from Kyrgyzstan (central
Asia). Journal of Vertebrate Paleontology. 24(3), 277A.
Averianov, Martin and Bakirov, 2005. Pterosaur and dinosaur remains from the
Middle Jurassic Balabansai Svita in the northern Fergana depression, Kyrgyzstan
(central Asia). Palaeontology. 48(1), 135-155.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
undescribed tetanurine (Hao, Fei, Hao and Liu, 2017)
Bajocian, Middle Jurassic
Guling Dinosaur Deposit, Xiashaximiao Formation, Chongqing, China
Material- (CLGPR coll.) few dorsal ribs, pubes, femur, tibia, fibula, several appendicular elements
Comments- Discovered in 2016,
Hao et al. (2017) reported this assemblage in Pu'an Township covers
three formations spanning most of the Jurassic. The Dinosaur
Fossil Wall providing most of the material has since been identified as
the Xiashaximiao Formation (the source of Omeisaurus puxiensis CLGPR V00005, Bashanosaurus primitivus CLGPR V00006 and Shunosaurus lii
referred specimen CLGPR V00007), making this the likely horizon of this
skeleton. Stiegler (2019) called this the "Guling Dinosaur Deposit" in
his Figure 5.2 showing Jurassic theropod localities of the Sichuan
basin. Hao et al.'s Figure 2 ("Photograph showing a dinosaur
skeleton in situ" shows a theropod skeleton including pubes with a
large interpubic fenestra exposed anteriorly, a tetanurine character.
References- Hao, Fei, Hao and
Liu, 2017. A large dinosaur assemblage has been discovered in Pu'an,
Yunyang County, Chongqing, China. Acta Geologica Sinica (English
Edition). 91(4), 1487-1488.
Stiegler, 2019. Anatomy, systematics, and paleobiology of noasaurid
ceratosaurs from the Late Jurassic of China. PhD thesis, The George
Washington University. 693 pp.
unnamed Tetanurae (He, 1984)
Bathonian-Callovian, Middle Jurassic
Hexi Commune, Shangshaximiao Formation, Sichuan, China
Material-
(CUT coll; = CCG coll) (multiple individuals) many teeth (~63 mm),
anterior cervical centrum (~68 mm; immature; Fig. 6-16, Pl. X Fig. 3),
mid cervical vertebra (~69 mm; Fig. 6-17), tenth cervical centrum
(immature; Fig. 6-19a), ~second dorsal centrum (~55 mm; immature?; Pl.
X Fig. 4), incomplete ~fourth dorsal vertebra (Fig. 6-19b), mid dorsal
centrum (immature; Fig. 6-19c), more than forty caudal vertebrae
including proximal caudal vertebra (Fig. 6-19d) and distal caudal
vertebra (~70 mm; Pl. X Fig. 5), incomplete coracoid (~98 mm
proximodistally), humerus (265 mm), ischium (~356 mm), femur, tibia
(~730 mm), fibula (~709 mm) and unguals
Comments-
He (1984) states that in 1964, 1979 and 1980 in Chengdu the institute
(= CUT) "conducted systematic collections in Hexi Commune (near Huomu
Station) in the suburbs of Qingyuan City, including many carnosaur
specimens, including many teeth, cervical vertebrae, dorsal vertebrae,
more than forty caudal vertebrae, complete ischium, femur, tibia and
fibula, as well as relatively complete humerus, coracoid and claws."
(translated) He referred these to Szechuanosaurus campi
because the syntypes were also found in the suburbs of Guangyuan and
believed to be from the Shangshaximiao Formation based on faunal
similarities and fossil abundance, "there is no significant difference
in shape and size" between S. campi
and the Hexi teeth, and "there is no evidence of the existence of two
or more carnosaurs" from that horizon. However, the teeth of S. campihave
not been shown to be diagnostic within e.g. Metriacanthosauridae,
multiple taxa with megalosaur-grade teeth are now known from the
Shangshaximiao (Leshansaurus, Yangchuanosaurus shangyouensis, Sinraptor hepingensis), and S. campi itself may be from the Penglaizhen Formation or slightly lower Shuining
Formation instead. Furthermore, He notes "that the tooth size in this batch of Szechuanosaurus campi
material we collected is quite varied, which means that in addition to
the differences in individual size, there may also be immature
specimens, because some vertebral centra and neural arches are unfused.
The largest individual is comparable to the type of Yanchuanosaurus shangyouensis
[sic], and the smallest individual is estimated to be only 4-5 meters
in length." Thus multiple individuals and perhaps multiple taxa
are involved, with only the tibia and fibula in Plate X Figures 8-10
being claimed to be from one individual. Note while Chure (2000)
mentioned a metatarsal as being in this material, He does not indicate
as such and Chure might have mistaken Plate X Figure 6 which is a
humerus. Indeed, Chure seems not to have translated the text so
understates the preserved vertebral number and misses the reference to
unguals. Yang et al. (2021) later describe the humeral histology, noting their Szechuanosaurus specimen is from Hexi and citing He's paper. This paper confirms the material "contains several incomplete individuals with large differences in size" (translated), that "The
specimen is currently preserved in the Museum of Chengdu University of
Technology" and that it was recovered from the Shangshaximiao Formation
at the same locality as Mamenchisaurus "guangyuanensis". "One
of the medium-sized individuals was recovered, mounted and exhibited",
which is photographed in their Figure 2, although it cannot be
determined what material is real and what is plaster.
This material was originally referred to Megalosauridae by He (1984)
based on tetanurine plesiomorphies (large teeth, short presacral
vertebrae, distally expanded ischium), while Chure (2000) placed it in
non-avetheropod Tetanurae based on the supposed lack of a posterodistal
coracoid process and subglenoid fossa, although Figure 6-18 of He
clearly shows both. Although Chure believes the information
available in the literature "make(s) it impossible to refer this
material to any family" and considered it indeterminate, the figures
and plates suggest otherwise. Among Late Jurassic theropods, the
slightly opisthocoelous cervicals are only known in piatnitzkysaurids
and coelurosaurs, with the long and low neural spines being unlike most
contemporary non-coelurosaur theropods, meaning the mid cervical
vertebra at least is not megalosauroid or carnosaurian.
Similarly, the large coracoid tubercle is unlike basal tetanurines and
more similar to ceratosaurs or coelurosaurs, although lacking the
hypertrophied size of the former. As suggested by Chure, the
ischium does resemble Megalosaurus in the ventral kink of the shaft and boot morphology, although it is much more robust, thus a referral to Leshansaurus is plausible. The tibia on the other hand is more similar to Sinraptor
in the anteroposteriorly short proximal end and anteroposterior
compression distally, so may be metriacanthosaurid. Based on this
brief comparison, the material deserves restudy and probably represents
multiple tetanurine taxa.
References- He, 1984. The Vertebrate Fossils of Sichuan. Sichuan Scientific
and Technical Publishing House, Chengdu, Sichuan. 168 pp.
Chure, 2000. A new species of Allosaurus from the Morrison Formation
of Dinosaur National Monument (Utah-Colorado) and a revision of the theropod
family Allosauridae. PhD thesis. Columbia University. 964 pp.
Yang, Liu and Zhang, 2021. The humeral diapophyseal histology and its biometric significance of Jurassic Szechuanosaurus campi (Theropoda, Megalosauridae) in Guangyuan City, Sichuan Province. Acta Geologica Sinica. 95(8), 2318-2332.
unnamed tetanurine (Han, Clark, Xu, Sullivan, Choiniere and Hone, 2011)
Late Callovian-Early Oxfordian, Middle-Late Jurassic
Wucaiwan, Lower Shishugou Formation, Xinjiang, China
Material- (IVPP V15848) tooth (~20x~10x5.4 mm)
Comments- This is called
Morphotype 2 by Han et al. (2011), who refer it to a basal tetanurine
or basal tyrannosauroid. It is placed in Tetanurae here based
on the enamel wrinkles.
Reference- Han, Clark, Xu,
Sullivan, Choiniere and Hone, 2011. Theropod teeth from the
Middle-Upper Jurassic Shishugou Formation of northwest Xinjiang, China.
Journal of Vertebrate Paleontology. 31(1), 111-126.
unnamed Tetanurae (Buffetaut and Ingavat, 1984)
Late Barremian, Early Cretaceous
Phu Wiang 1, Sao Khua Formation, Thailand
Material- (SM-TF 1739-1) proximal tibia (~14 mm deep proximally)
?(SM-TF 1739-2) proximal humerus (~16 mm broad)
Comments- These were discovered
in 1982 and described by Buffetaut and Ingavat (1984) as compsognathid,
based on size and general morphology. Samathi et al. (2019)
stated the tibia "represents a small-sized tetanuran based on the
following synapomorphy: the fibular crest of the tibia does not extend
to the proximal end."
References- Buffetaut and
Ingavat, 1984. A very small theropod dinosaur from the Upper Jurassic
of Thailand. Comptes rendus de l’Acad�mie des Sciences Paris II. 298(20),
915-918.
Samathi, Chanthasit and Sander, 2019. A review of theropod dinosaurs
from the Late Jurassic to mid-Cretaceous of southeast Asia. Annales de
Pal�ontologie. 105(3), 201-215.
undescribed Tetanurae (Manabe and Barrett, 2000)
Valanginian-Hauterivian, Early Cretaceous
Kuwajima Formation of the Tetori Group, Japan
Material- (SBEI-008) tooth (Matsuoka et al., 2002)
(SBEI coll.) teeth (Matsuoka et al., 2002)
Comments- These large teeth were announced as tyrannosaurid in Barrett
and Manabe's (2000) abstract, but Matsuoka et al. (2002) merely calls them Theropod
Type A. The enamel wrinkles are tetanurine and the high labiolingual compression
is like Fukuiraptor, but not tyrannosaurids. Given their age and location,
it would not be surprising if they were megaraptoran.
References- Barrett and Manabe, 2000. The dinosaur fauna from the Earliest
Cretaceous Tetori Group of Central Honshu, Japan. Journal of Vertebrate Paleontology.
20(3), 28A-29A.
Matsuoka, Kusuhashi, Takada and Setoguchi, 2002. A clue to the Neocomian vertebrate
fauna: Initial results from the Kuwajima 'Kaseki-kabe' (Tetori Group) in Shiramine,
Ishikawa, central Japan. Memoirs of the Faculty of Science, Kyoto University,
Series of Geology and Mineralogy. 59(1), 33-45.
unnamed tetanurine (Goodwin et al., 1999)
Tithonian, Late Jurassic
Mugher Mudstone, Ethiopia
Material- (UCMP 170802) partial tooth
(UCMP 172477) tooth fragment
(UCMP 172478) fragmentary tooth
Comments- Referred to cf. Acrocanthosaurus sp. by Goodwin et al.
(1999), but this is unlikely given the provenance. The referral was only due
to rectangular serrations, which are plesiomorphic for theropods. Carrano et
al. (2012) notes the presence of enamel wrinkles means the specimens are tetanurine.
References- Goodwin, Clemens, Hutchison, Wood, Zavada, Kemp, Duffin and
Schaff, 1999. Mesozoic continental vertebrates with associated palynostratigraphic
dates from the northwestern Ethiopian plateau. Journal of Vertebrate Paleontology.
19(4), 728-741.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
unnamed basal tetanurine (Janensch, 1925)
Late Kimmeridgian, Late Jurassic
Middle Dinosaur Member of the Tendaguru Formation, Tanzania
Material- (HMN MB R 1763) tibia (163 mm)
Comments- Originally described as 'coelurosaurier A', Rauhut (2005) has
identified it as a basal tetanurine.
References- Janensch, 1925. Die Coelurosaurier und Theropoden der Tendaguru-Schichten
Deutsch-Ostafrikas. Palaeontographica. (Supp. 7)1, 1-99.
Rauhut, 2005. Post-cranial remains of 'coelurosaurs' (Dinosauria, Theropoda)
from the Late Jurassic of Tanzania. Geological Magazine. 142(1), 97-107.
unnamed Tetanurae (Janensch, 1920)
Late Tithonian, Late Jurassic
Upper Dinosaur Member of the Tendaguru Formation, Tanzania
Material- (HMN MB R. 1755) radius (198 mm) (Janensch, 1929)
(HMN MB R 1936; = TL 43) (juvenile) posterior dorsal centrum (86
mm) (Janensch, 1920)
....(HMN MB R 2163; = TL 44) (juvenile) mid dorsal centrum (89 mm) (Janensch, 1920)
....(HMN coll.; = TL 8; lost) (juvenile) first dorsal centrum (73 mm) (Janensch, 1920)
(HMN MB R 2161; = TL 46c and 46d) incomplete distal caudal vertebra, partial distal
caudal vertebra (Janensch, 1925)
....(HMN MB R 2165; = TL 46a and 46b) distal caudal vertebra (105 mm), partial distal
caudal vertebra (Janensch, 1925)
(HMN MB R 3622; = TL 30) femur (825 mm) (Janensch, 1925)
....(HMN MB R 3623; = TL 16; = HMN TL 96 of Carrano, 1998) femur (822 mm) (Janensch, 1925)
....(HMN MB R 3624; = TL 42) tibia (807 mm) (Janensch, 1925)
Comments- Janensch (1920) initially referred the dorsal centra to Ceratosaurus
(followed by Madsen and Welles, 2000), though he later (1925) was more cautious.
Rauhut (2011) found them to be a tetanurine based on the strong ventral keel,
though further assignment is difficult (besides exclusion from clades with pleurocoelous
posterior dorsals). Rauhut could only identify the hindlimb elements to non-coelurosaurian
Tetanurae, and the distal caudals to Tetanurae.
Janensch (1929) referred a radius HMN M.B.R. 1755 excavated in 1912 from the later Upper
Dinosaur Member to Elaphrosaurus, but Rauhut and Carrano (2016) rejected this
as there is no overlap with the holotype and ceratosaurs would be
expected to have shorter radii.
References- Janensch, 1920. Uber Elaphrosaurus bambergi und die
Megalosaurier aus den Tendaguru-Schichten Deutsch-Ostafricas. Sitzungsberichte
Gesellschaft Naturforschender Freunde Berlin. 8, 225-235.
Janensch, 1925. Die Coelurosaurier und Theropoden der Tendaguru-Schichten Deutsch-Ostafrikas.
Palaeontographica. 1(supp. 7), 1-99.
Janensch, 1929. Ein aufgestelltes und rekonstruiertes Skelett von Elaphrosaurus
bambergi mit einem Nachtrag zur Osteologie dieses Coelurosauriers. Palaeontographica
(Supp. 7)1, 279-286.
Carrano, 1998. The evolution of dinosaur locomotion: Functional morphology,
biomechanics, and modern analogs. PhD Thesis, The University of Chicago. 424
pp.
Madsen and Welles, 2000. Ceratosaurus (Dinosauria, Theropoda) a revised
osteology. Miscellaneous Publication 00-2 Utah Geological Survey. 80 pp.
Rauhut, 2011. Theropod dinosaurs from the Late Jurassic of Tendaguru (Tanzania).
Palaeontology. 86, 195-239.
Rauhut and Carrano, 2016. The theropod dinosaur Elaphrosaurus bambergi Janensch, 1920, from the Late Jurassic of Tendaguru, Tanzania. Zoological Journal of the Linnean Society. 178(3), 546-610.
unnamed tetanurine (Ricqles, 1967)
Aptian-Albian, Early Cretaceous
Elrhaz Formation of the Tegama Group, Niger
Material- partial manual ungual I
Comments- This specimen was described by Ricqles (1967) as dinosaurian,
and Rozhdestveksky (1970) placed it in Theropoda. Nessov (1995) referred it
to Therizinosauria or "groups most closely related to them" (which
in his opinion consisted of spinosaurids and dryptosaurids). The ungual closely
resembles both Baryonyx and Alxasaurus, though it tapers more
than the former and is less curved than the latter.
References- Ricqles, 1967. La paleontologie de terrain: Un bilan international.
Atomes. 243, 337-341.
Rozhdestvensky, 1970. Giant claws of enigmatic Mesozoic reptiles. Paleontological
Journal. 1970(1), 131-141.
Nessov, 1995. Dinosaurs of nothern Eurasia: new data about assemblages, ecology,
and paleobiogeography. Institute for Scientific Research on the Earth's Crust,
St. Petersburg State University, St. Petersburg. 1-156.
unnamed Tetanurae (Stromer, 1914)
Cenomanian, Late Cretaceous
Baharija Formation, Egypt
Material- (IPHG 1911 XII 29; destroyed) proximal femur (Stromer, 1914)
(IPHG 1912 VIII 76; destroyed) tibia (575 mm) (Stromer, 1934)
(IPHG 1912 VIII 192; destroyed) incomplete tibia (Stromer, 1934)
Comments- Stromer (1934) described the tibiae IPHG 1912 VIII 76 and 1912 VIII 192 from the Baharija Formation
of Egypt and assigned them to cf. Elaphrosaurus bambergi. He stated they were (translated) "most similar to that of Elaphrosaurus bambergi,
but still differences are evident. As has been expected by the
geological age difference, impossibly the same species can be present,
but a close relative." The distally placed fibular crest suggest
these are tetanurines, as noted by Rauhut and Carrano (2016), and the
lack of an ascending process buttress further constrains their
relationships. They are roughly similar accounting for distortion, and
share the same small cnemial crest which leads to an unusually broad
proximal outline (76-81% of anteroposterior length), so may be
conspecific. Stromer further referred proximal femur IPHG 1911 XII 29
which he had previously (Stromer, 1914) placed in Ornithopoda, although
this was only loosely defended based on the size similarity to the
Baharija tibiae and that it couldn't be compared well enough to Elaphrosaurus
to discount being closely related. In fact this femur has a
prominent pointed accessory trochanter showing it is an avetheropod and
not an elaphrosaur.
References- Stromer, 1914.
Ergebnisse der Forschungreisen Prof. E. Stromers in den Wusten
Agyptens. I: Die Topographie und Geologie der Strecke Gharqu-Baharije
nebst Ausfuhrungen uber de geologische Geschichte Agyptens.
Abhandlungen der Koniglichen Bayerischen Akademie der Wissenschaften,
Mathematisch-physikalischen Classe, Munchen. 26, 1-78.
Stromer, 1934. Ergebnisse der Forschungsreisen Prof. E. Stromers
in den W�sten �gyptens. II. Wirbeltierreste der Baharije-Stufe (unterstes
Cenoman). 13. Dinosauria. Abhandlungen der Bayerischen Akademie der Wissenschaften
Mathematisch-naturwissenschaftliche Abteilung, Neue Folge. 22, 1-79.
Rauhut and Carrano, 2016. The theropod dinosaur Elaphrosaurus bambergi Janensch, 1920, from the Late Jurassic of Tendaguru, Tanzania. Zoological Journal of the Linnean Society. 178(3), 546-610.
unnamed Tetanurae (Rauhut, 2002)
Late Toarcian-Bajocian, Early-Middle Jurassic
Ca�ad�n Asfalto Formation, Chubut, Argentina
Material- (MPEF PV 1717) partial maxillae, nasal fragments, partial palatines (Rauhut, 2002)
distal caudal vertebra, metatarsal IV (Pol and Rauhut, 2012)
Comments- Rauhut (2002) initially considered MPEF PV 1717 a ceratosaur, but described
it in 2007 as a basal tetanurine. While it differs from Piatnitzkysaurus,
it cannot be compared to Condorraptor.
Pol and Rauhut (2012) note a distal caudal and metatarsal as a basal tetanurine much
larger than Piatnitzkysaurus or Condorraptor.
References- Rauhut, 2002. Dinosaur evolution in the Jurassic: A South
American perspective. Journal of Vertebrate Paleontology. 22(3), 89A.
Rauhut, 2007. A fragmentary theropod skull from the Middle Jurassic of Patagonia.
Ameghiniana. 44(2), 479-483.
Pol and Rauhut, 2012. A Middle Jurassic abelisaurid from Patagonia
and the early diversification of theropod dinosaurs. Proceedings of the Royal
Society B. 279(1741), 3170-3175.
undescribed tetanurine (Apesteguia and Bonaparte, 2004)
Late Berriasian-Valanginian, Early Cretaceous
Bajada Colorada Formation, Neuquen, Argentina
Material- (MMCH-PV-68-6) (large) tooth (Gallina, Apesteguia, Haluza and
Canale, 2014)
fragmentary femur (Apesteguia and Bonaparte, 2004)
Comments- The distal femur noted by Apesteguia and Bonaparte (2004) is
said to have a promiment fourth trochanter, weak extensor groove, and moderately
developed mediodistal crest. The authors state it resembles Streptospondylus
in having a medially projected tibial condyle and wide intercondylar groove.
References- Apesteguia and Bonaparte, 2004. Bajada Colorada (Valanginian)
dinosaurs from Neuquen: Note on the oldest Cretaceous dinosaurs from the Neuquen
basin. XX Jornadas Argentinas de Paleontologia de Vertebrados, resumenes. 34R.
Canale, Apesteguia, Gallina, Haluza, Gianechini and Pazo, 2014. Theropod remains
from the Bajada Colorada Formation (Berriasian-Valanginian) from Neuquen Province,
Argentina. Reunion de Comunicaciones de la Asociacion Paleontologica Argentina,
abstracts. Ameghiniana. 52(1) suplemento, 5.
unnamed Tetanurae (Benson, Rich, Vickers-Rich and Hall, 2012)
Late Aptian-Early Albian, Early Cretaceous
Eumeralla Formation of the Otway Group, Victoria, Australia
Material- (NMV P208234) (adult) dorsal vertebra (18 mm) (Benson, Rich, Vickers-Rich and Hall, 2012)
(NMV P252405) manual phalanx II-1 (96 mm) (Poropat, White, Vickers-Rich and Rich, 2019)
(NMV P252700) cervical rib (Poropat, White, Vickers-Rich and Rich, 2019)
(NMV P252704) mid caudal vertebra (53 mm) (Poropat, White, Vickers-Rich and Rich, 2019)
Comments- Poropat et al. (2019)
stated "based on our comparisons of NMV P252700 with the cervical ribs
of several megalosauroid, allosauroid, and tyrannosauroid taxa, we
contend that it is most similar to megaraptorids" but ended up
interpreting it more generally "as a non-maniraptoriform tetanuran
cervical rib." They said NMV P252704 was similar to Gualicho (as Aoniraptor)
in that "the transverse processes are restricted to the posterior
two-thirds of the centrum, the postzygapophyses extend only slightly
posterodorsal to the posterior margin of the centrum, the
prezygapophyses are relatively small and project anterodorsally rather
than mostly anteriorly, and a prespinal fossa is present between the
prezygapophyses." However, "assigning NMV P252704 to
Megaraptoridae on the basis of these superficial similarities is not
possible, because several of these features are more widely distributed
among Theropoda." Regarding tha manual phalanx, they say "there
are similarities between NMV P252405 and the Australovenator
phalanx originally identified as right manual III-1" but they "regard
NMV P252405 as Tetanurae indet., because it clearly does not pertain to
Ceratosauria."
References- Benson, Rich, Vickers-Rich and Hall, 2012. Theropod fauna
from southern Australia indicates high polor diversity and climate-driven dinosaur
provinciality. PLOS One. 7(5), e37122.
Poropat, White, Vickers-Rich and Rich, 2019. New megaraptorid
(Dinosauria: Theropoda) remains from the Lower Cretaceous Eumeralla
Formation of Cape Otway, Victoria, Australia. Journal of Vertebrate
Paleontology. 39, e1666273.