Ceratosauria Marsh, 1884
Definition- (Ceratosaurus nasicornis <- Passer domesticus)
(Sereno, in press; modified from Rowe, 1989)
Other definitions- (Liliensternus liliensterni + Coelophysis
bauri + "Syntarsus" rhodesiensis + "Syntarsus"
kayentakatae + Segisaurus halli + Sarcosaurus woodi + Dilophosaurus
wetherilli + Ceratosaurus nasicornis) (Rowe and Gauthier, 1990)
(Coelophysis bauri <- Passer domesticus) (modified from Sereno,
1998)
(Ceratosaurus nasicornis <- Vultur gryphus) (
Dal Sasso, Maganuco and Cau, 2018)
= Ceratosauria sensu
Dal Sasso, Maganuco and Cau, 2018
Definition- (Ceratosaurus nasicornis <- Vultur gryphus)
Comments- Marsh (1884) erected this as a suborder including Ceratosaurus
only, though he later (1895) added Ornithomimidae. This taxon was ressurrected
in 1984 (published in 1986) by Gauthier to contain coelophysoids, Dilophosaurus
and ceratosaurs sensu stricto. This was followed by most phylogenies in the
1990's (e.g. Rowe and Gauthier, 1990; Holtz, 1994; Sereno, 1999; Holtz, 2000).
Some non-cladistic phylogenies at the time (Bakker, 1986; Paul, 1984, 1988)
advocated ceratosaurs sensu stricto as being closer to birds than coelophysoids
and Dilophosaurus, which was suggested in some more recent unpublished
analyses (Currie, 1995; Rauhut, 1998; Carrano and Sampson, 1999) and has become
the current consensus (Carrano et al., 2002; Rauhut, 2003; Wilson et al., 2003;
Carrano et al., 2005; Ezcurra and Novas, 2007; Smith et al., 2007; Ezcurra,
2012). However, a few recent studies (Tykoski and Rowe, 2004; Tykoski, 2004,
2005) have again recovered coelophysoid ceratosaurs. Tykoski (2005) found excluding
ontogenetically variable characters (mostly bone fusions) generated trees excluding
Coelophysoidea from Ceratosauria. According to Tykoski, excluding these characters
and miscoding many others have led to the current concensus. It should be noted
that only four more steps are needed to place ceratosaurs closer to tetanurines
than to coelophysoids in his trees. Thus, either topology should be considered
possible.
Not ceratosaurs- Coria et al.
(2010) initially wrote "certain anatomical features of the collected
specimens suggest abelisaur affinities in the theropods" of the
Mulichinco Formation of Argentina, but Coria et al. (2019) later
described it as the holotype of the carnosaur Lajasvenator (MLL-PV-005).
References- Marsh, 1884. The classification and affinities of dinosaurian
reptiles. Nature. 31, 68-69.
Marsh, 1895. On the affinities and classification of the dinosaurian reptiles.
American Journal of Science. 50, 483-498.
Gauthier, 1984. A cladistic analysis of the higher systematic categories of
the Diapsida. PhD thesis. University of California, Berkeley. 564 pp.
Paul, 1984. The archosaurs: A phylogenetic study. Third Symposium on Mesozoic
Terrestrial Ecosystems, Short Papers. 175-180.
Bakker, 1986. The Dinosaur Heresies. Kensington, New York. 481 pp.
Gauthier, 1986. Saurischian Monophyly and the Origin of Birds. Memoires of the
California Academy of Sciences. 8, 1-55.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
464 pp.
Rowe, 1989. A new species of the theropod dinosaur Syntarsus from the
Early Jurassic Kayenta Formation of Arizona. Journal of Vertebrate Paleontology.
9(2), 125-136.
Bonaparte, Novas and Coria, 1990. Carnotaurus sastrei Bonaparte, the
horned, lightly built carnosaur from the Middle Cretaceous of Patagonia. Natural
History Museum of Los Angeles County Contributions in Science. 416, 1-42.
Rowe and Gauthier, 1990. Ceratosauria. In Weishampel, Dodson and Osmolska (eds.).
The Dinosauria. University of California Press.
151-168.
Novas, 1992. La evolucion de los dinosaurios carnivoros. In Sanz and Buscalioni (eds.). Los Dinosaurios y Su Entorno Biotico:
Actas del Segundo Curso de Paleontologia in Cuenca. Instituto "Juan Valdez",
Cuenca, Argentina. 126-163.
Holtz, 1994. The phylogenetic position of the Tyrannosauridae: Implications
for theropod systematics. Journal of Paleontology. 68(5), 1100-1117.
Currie, 1995. Phylogeny and systematics of theropods (Dinosauria). Journal of
Vertebrate Paleontology. 15(3, 25A.
Rauhut, 1998. Elaphrosaurus bambergi and the early evolution of theropod
dinosaurs. Journal of Vertebrate Paleontology. 18(3), 71A.
Sereno, 1998. A rationale for phylogenetic definitions, with application to
the higher-level taxonomy of Dinosauria. Neues Jahrbuch für Geologie und
Paläontologie Abhandlungen. 210(1), 41-83.
Carrano and Sampson, 1999. Evidence for a paraphyletic 'Ceratosauria' and it’s
implications for theropod dinosaur evolution. Journal of Vertebrate Paleontology.
19(3), 36A.
Padian, Hutchinson and Holtz, 1999. Phylogenetic definitions and nomenclature
of the major taxonomic categories of the carnivorous Dinosauria (Theropoda).
Journal of Vertebrate Paleontology. 19(1), 69-80.
Sereno, 1999. The evolution of dinosaurs. Science. 284, 2137-2147.
Holtz, 2000 (as 1998). A new phylogeny of the carnivorous dinosaurs. Gaia. 15, 5-61.
Carrano, Sampson and Forster, 2002. The osteology of Masiakasaurus knopfleri,
a small abelisauroid (Dinosauria:Theropoda) from the Late Cretaceous of Madagascar.
Journal of Vertebrate Palaeontology. 22(3), 510-534.
Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs.
Special Papers in Palaeontology. 69, 1-213.
Wilson, Sereno, Srivastava, Bhatt, Khosla and Sahni, 2003. A new abelisaurid
(Dinosauria, Theropoda) from the Lameta Formation (Cretaceous, Maastrichtian)
of India. Contributions from the Museum of Paleontology. The University of Michigan.
31, 1-42.
Tykoski, 2004. Ontogenetic stage assessment and the position of Coelophysoidea
within basal Theropoda. Journal of Vertebrate Paleontology. 24(3), 7A-8A.
Tykoski and Rowe, 2004. Ceratosauria. In Weishampel, Dodson and Osmolska (eds.).
The Dinosauria Second Edition. University of California Press. 47-70.
Carrano, Hutchinson and Sampson, 2005. New information on Segisaurus halli,
a small theropod dinosaur from the Early Jurassic of Arizona. Journal of Vertebrate
Paleontology. 25(4), 835-849.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD
thesis. University of Texas at Austin. 553 pp.
Ezcurra and Novas, 2007 (online 2006). Phylogenetic relationships of the Triassic theropod
Zupaysaurus rougieri from NW Argentina. Historical Biology. 19(1), 35-72.
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.
Carrano and Sampson, 2008 (online 2007). The Phylogeny of Ceratosauria (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 6, 183-236.
Coria, Currie,
Koppelhus, Braun and Cerda, 2010. First record of a Valanginian (Early
Cretaceous) dinosaur association from South America. Journal of
Vertebrate Paleontology 30: 75A.
Ezcurra, 2012. Phylogenetic analysis of Late Triassic - Early Jurassic neotheropod
dinosaurs: Implications for the early theropod radiation. Journal of Vertebrate
Paleontology. Program and Abstracts 2012, 91.
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.
Coria, Currie, Ortega and Baiano, 2020 (online 2019). An Early Cretaceous,
medium-sized carcharodontosaurid theropod (Dinosauria, Saurischia) from
the Mulichinco Formation (upper Valanginian), Neuquén Province,
Patagonia, Argentina. Cretaceous Research. 111, 104319.
“Merosaurus” Welles, Powell
and Pickering vide Pickering, 1995
“M. newmani” Welles, Powell and Pickering vide Pickering, 1995
Hettangian-Early Sinemurian, Early Jurassic
Blue Lias Formation, England
Material- (GSM 109560) partial femur (~380 mm), partial tibia (lost)
Other diagnoses- Pickering (1995b) considered "Merosaurus"
to be "very similar to Sarcosaurus" but distinguished it based
on several characters. The absent trochanteric shelf is known to vary dimorphically
in ceratosaur and coelophysoid taxa. The remaining ventral portion of the femoral
head is not angled more medially, the fourth trochanter is not medially concave,
and is more medially placed not less so. Lateromedial placement of the fourth
trochanter is highly variable within theropod taxa, so cannot be used to diagnose
"Merosaurus" in any case.
Comments- Owen received the partial hindlimbs NHMUK 39496 and GSM 109560
in 1858 and used them as the basis for his dinosaur genus Scelidosaurus
in an encyclopedia entry the following year. While this is often claimed to
be an nomen nudum (e.g. Newman, 1968), genus names published before 1931 do
not require species names or illustrations to be valid (ICZN Article 12). Owen
later (1861) gave his taxon the species name harrisonii and described
it in detail, referring the ungual GSM 109561, a partial postcranium in the
Lyme Regis Museum, and the skull of NHMUK R1111. Lydekker (1888) made NHMUK 39496
the type specimen, though the basal thyreophoran NHMUK R1111 (whose postcranium
was soon found and described in 1862) formed the basis for peoples' ideas of
Scelidosaurus. Newman (1968) believed NHMUK 39496 and GSM 109560 to be
megalosaurids. However, they were only compared to Megalosaurus among
theropods, making this familial assignment in need of verification. As the name
Scelidosaurus had been associated with the thyreophoran, Charig and Newman
(1994) petitioned the ICZN to recognize NHMUK R1111 (as BMNH R1111) as the lectotype, which was
accepted in 1994 as Opinion 1788. Welles and Powell studied the theropod material
in 1974 for their unpublished European theropod paper, intending to name it
Merosaurus newmani. This was first found in publically available print
in 1995 when Pickering credited the name to Welles, Powell and Pickering in
an unpublished bibliographic manuscript. In that same year, Pickering printed
a packet with a full description of the taxon, which he credited to only Welles
and Powell (though its osteology was also credited to himself). This is a nomen
nudum however, as he didn't follow ICZN Article 8.1.3- it must have been produced
in an edition containing simultaneously obtainable copies by a method that assures
numerous identical and durable copies. Pickering intends GSM 109560 to be the
type, and referred NHMUK 39496 and GSM 109561. He considered it a probable ceratosaur
sensu lato metataxon. Pickering will describe it in his in progress work Mutanda
Dinosaurologica. Naish and Martill (2007) referred all three specimens to Tetanurae
without comment. Most recently, Benson (2009, 2010) redescribed NHMUK 39496 and
GSM 109560. He considered GSM 109560 to be an indeterminate theropod.
GSM 109560 is based on a femur lacking the head and distal end, and a tibial
shaft which was not illustrated by Owen and has been lost. Contra Owen and Pickering,
there is no reason to refer this to the same taxon as NHMUK 39496 as they share
only a short area of distal shaft, and the GSM specimen is only ~60% as large.
Also contra Pickering, the anterior trochanter is not conical but is lateromedially
narrower than anteroposteriorly, making it alariform. The specimen is not a
robust coelophysoid or ceratosaur individual based on the absence of a ridge-like
trochanteric shelf. It is most similar to Liliensternus, Dilophosaurus,
Ceratosaurus and tetanurines in having a straight shaft in anterior view.
The anteroposterior width of the anterior trochanter seems less than in tetanurines,
but greater than basal coelophysoids. It may be a gracile morph of ceratosaur.
References- Owen, 1859. Palaeontology. Encyclopaedia Britannica, Edition
8. 17, 91-176.
Owen, 1861. Monograph of the fossil Reptilia of the Liassic formations. Part
I. A monograph of the fossil dinosaur (Scelidosaurus harrisonii Owen)
of the Lower Lias. Palaeontolographical Society Monographs. 13, 1-14.
Owen, 1862. Monographs on the British Fossil Reptilia from the Oolitic Formations.
Part second, containing Scelidosaurus harrisonii and Pliosaurus grandis.
Palaeontolographical Society Monographs. 1-16.
Lydekker, 1888. Catalogue of the Fossil Reptilia and Amphibia in the British
Museum (Natural History), Cromwell Road, S.W., Part 1. Containing the Orders
Ornithosauria, Crocodilia, Dinosauria, Squamata, Rhynchocephalia, and Proterosauria.
British Museum of Natural History, London. 309 pp.
Newman, 1968. The Jurassic dinosaur Scelidosaurus harrisoni Owen. Palaeontology.
11, 40-43.
Charig and Newman, 1992. Scelidosaurus harrisonii Owen, 1861 (Reptilia,
Ornithischia): Proposed replacement in inappropriate lectotype. Bulletin of
Zoological Nomenclature. 49, 280-283.
ICZN, 1994. Opinion 1788. Scelidosaurus harrisonii Owen, 1861 (Reptilia,
Ornithischia): Lectotype replaced. Bulletin of Zoological Nomenclature. 51(3),
288.
Pickering, 1995a. Jurassic Park: Unauthorized Jewish Fractals in Philopatry.
A Fractal Scaling in Dinosaurology Project, 2nd revised printing. Capitola,
California. 478 pp.
Pickering, 1995b. An extract from: Archosauromorpha: Cladistics and osteologies.
A Fractal Scaling in Dinosaurology Project. 11 pp.
Olshevsky, DML 1999. https://web.archive.org/web/20210309141613/http://dml.cmnh.org/1999Dec/msg00193.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. 164, 493-510.
Benson, 2009. The taxonomy, systematics and evolution of the British theropod
dinosaur Megalosaurus. PhD thesis. University of Cambridge. [? pp]
Benson, 2010. The osteology of Magnosaurus nethercombensis (Dinosauria,
Theropoda) from the Bajocian (Middle Jurassic) of the United Kingdom and a re-examination
of the oldest records of tetanurans. Journal of Systematic Palaeontology. 8(1),
131-146.
Pickering, in prep. Mutanda Dinosaurologica.
"Ngexisaurus" Zhao, 1983
"N. dapukaensis" Zhao, 1985
= "Ngexisaurus changduensis" Zhao and Cheng, 1985
= "Ngexisaurus dapukanensis" 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 "coelurosaurs (Ngexisaurus
Chao)" in the Middle Jurassic. Lacking a description, illustration or species
name, it was a nomen nudum. Zhao seems to place all Jurassic coelurosaurs in
Coeluridae and states in Middle Jurassic coelurosaurs the tooth "crown
becomes thinner with serrated anterior edge only". This may indicate the
"Ngexisaurus" material includes teeth with this morphology. Zhao used
a classic concept of Coelurosauria, which only tells us "Ngexisaurus"
is probably a small theropod. 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 Ngexisaurus dapukaensis as a coelurosaur from the Middle Jurassic
Dabuka Group of Tibet, but again with no description or illustration. In the
same volume, Zhao and Cheng (1985) list the species as Ngexisaurus changduensis
instead. Zhang and Li (1997) also list it as Ngexisaurus changduensis,
from the Middle Dabuka Formation of Dabuka, Qamdo County, Xizang. Weishampel
et al. (2004) list it as Ngexisaurus dapukanensis from the Dapuka Group
of Xinjiang, Uygur Zizhiqu and refer it to Ceratosauria (sensu lato). Its assignment
to Ceratosauria by Weishampel et al. (note Zhao is a coauthor) may indicate
it is a ceratosaur sensu stricto (since coelophysoids were extinct by the Middle
Jurassic). Perhaps it is similar to small early ceratosaurs like Limusaurus
and Berberosaurus, but this is only conjecture. It is listed as Ngexisaurus
changduensis Zhao gen. et sp. nov. (MS) in Fang et al. (2006), suggesting
that Zhao's monograph was indeed never published and is still a manuscript.
They refer it to Procompsognathidae (or perhaps basal Compsognathidae). Which
species name will be used for it when it is published is uncertain.
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.
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. 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.
Berberosaurus Allain, Tykoski,
Aquesbi, Jalil, Monbaron, Russell and Taquet, 2007
B. liassicus Allain, Tykoski, Aquesbi, Jalil, Monbaron, Russell
and Taquet, 2007
Toarcian, Early Jurassic
Pt locality, Douar of Tazouda, Upper bone-bed of the Toundoute continental series, Morocco
Holotype- (MHNM-Pt9) (subadult) incomplete mid cervical vertebra (~53 mm)
....(MHNM-Pt16) distal tibia
....(MHNM-Pt19) incomplete femur (~505 mm)
....(MHNM-Pt20) fibula (447 mm)
....(MHNM-Pt21) proximal tibia
....(MHNM-Pt22) metacarpal II (78 mm)
....(MHNM-Pt23) posterior synsacrum (s4 64, s5 70 mm)
Toarcian, Early Jurassic
To1 locality, Douar of Tazouda, Upper bone-bed of the Toundoute continental series, Morocco
Paratype- (MHNM-To1-218) proximal femur (~408 mm)
Toarcian, Early Jurassic
Toundoute continental series, Morocco
unknown relation to types- (MHNM coll.) postorbital, braincase, teeth, axis (Allain pers. comm. to Ibrahim et al., 2017)
Diagnosis- (from Allain et al., 2007) differs from Elaphrosaurus
in: short cervical centra; pneumatic foramina on the cervical neural arch.
from Ceratosaurus in: camerate structure of cervical vertebra; low and
short neural spine of the cervical vertebra; femoral anterior trochanter reaches
proximally to mid-point of femoral head.
from MNN TIG6 in: absence of the epipophyseal-prezygapophyseal
lamina on the cervical neural arches; short cervical neural spine.
from Abelisauria in: distal end of metacarpal with deep extensor pits; pronounced
femoral trochanteric shelf.
Comments- Discovered in 2000, Allain et al. (2004) noted that associated with the Tazoudasaurus
types "were isolated elements of medium-sized and large theropods of
uncertain affinities", the former eventually being described as Berberosaurus.
They considered the holotype a subadult based on the mix of immature
("absence of fusion between cervical ribs and their respective
vertebral centra, no co-ossification between the astragalus and tibia,
and no co-ossifcation between the astragalus and fibula") and mature
("the sacral centra exhibit full fusion to one another such that their
sutures are nearly indiscernible; the femoral anterior trochanter is a
mediolaterally compressed flange (aliform process) projecting
anteriorly from the bone; the medial side of the proximal end of the
fibula is excavated by a longitudinal groove, and the latter is
overlapped by an oblique (posteroproximally to anterodistally oriented)
ridge; and the fibula bears a medial flange that overlaps part of the
ascending process of the astragalus") characters. Ibrahim et al. (2017) report "The axis, a postorbital, the braincase and teeth of Berberosaurus
are currently being studied (Allain pers. comm.)" (translated),
although whether these belong to the holotype, paratype or new specimen
is not stated.
Allain et
al. (2007) used Tykoski's ceratosaur matrix to recover Berberosaurus as an abelisauroid sister to Xenotarsosaurus+Abelisauria.
More recetly, it has been recovered as a non-neoceratosaurian
etrigansaur (Wang et al., 2017) or a non-neoceratosaurian ceratosaur
sister to Saltriovenator using Cau's megamatrix (Dal Sasso et al., 2018).
References-
Allain, Aquesbi, Dejax, Meyer, Monbaron, Montenat, Richir, Rochdy,
Russell and Taquet, 2004. A basal sauropod dinosaur from the Early
Jurassic of Morocco. Comptes Rendus Palevol. 3, 199-208.
Allain, Tykoski, Aquesbi, Jalil, Monbaron, Russell and Taquet,
2007. An abelisauroid (Dinosauria: Theropoda) from the Early Jurassic of the
High Atlas Mountains, Morocco, and the radiation of ceratosaurs. Journal of
Vertebrate Paleontology. 27(3), 610-624.
Ibrahim, Sereno, Zouhri and Zouhri, 2017. Les dinosaures du Maroc -
aperçu historique et travaux récents. Mémoires de la Societé géologique
de France. 180, 249-284.
Wang, Stiegler, Amiot, Wang, Du, Clark and Xu, 2017 (online 2016). Extreme ontogenetic
changes in a ceratosaurian theropod. Current Biology. 27(1), 144-148.
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.
Saltriovenator Dal Sasso, Maganuco and Cau, 2018
= "Saltriosaurus" Dalla Vecchia,
2001
S. zanellai Dal Sasso, Maganuco and Cau, 2018
Early Sinemurian, Early Jurassic
Saltrio Formation, Italy
Holotype- (MSNM V3664) (~8
m; ~1.2-1.6 tons, 24 year old subadult) cranial fragment, splenial
fragment, prearticular fragment, cervical rib fragment, nine dorsal rib
fragments, scapulae (one incomplete, one fragment), partial coracoid,
incomplete furcula (~232 mm), partial sternal plate, humeri (one
proximal; 358 mm), distal carpal II, metacarpal II (129 mm), phalanx
II-1 (~65 mm), fragmentary phalanx II-2, distal manual ungual ?II,
phalanx III-1 (44 mm), phalanx III-2 (41 mm), phalanx III-3, proximal
manual ungual III, distal tarsal III, distal tarsal IV, proximal
metatarsal II, proximal metatarsal III, partial metatarsal IV, partial
metatarsal V
Paratype- (MSNM V3659) lateral tooth (43.5x18.2x~8.9 mm)
Diagnosis- (after Dal Sasso et
al., 2018) humerus with deltopectoral crest protruding anteromedially
for more than twice the shaft diameter, with distal lamina forming an
abrupt corner (about 90 degrees) with the proximodistal axis of the
humeral shaft; metacarpal II with hypertrophied semicircular extensor
lip protruding over the condylar level and bordering dorsolaterally a
very deep and wide extensor pit; manual phalanx II-1 with flexor palmar
groove which is deep and narrow, and bearing a distinct bump distal to
the dorsal extensor process; manual ungual III with prominent flexor
tubercle which is distinctly separated from articular facet by a
concave cleft.
Comments- This specimen was discovered in summer 1996 and
first announced at a press conference in 2000, mentioned in newspapers
on November 10th. The nickname 'saltriosaur' was given to it, changed
to "Saltriosaurus" in Della Vecchia's (2001) popular article, and officially described as Saltriovenator by Dal Sasso et al. (2018).
While Dal Sasso originally suggested it might be a basal tetanurine, he
later (2003, 2004) also proposed the possibility it is a carnosaur.
Benson (2010) considered Dal Sasso's (2003) evidence for tetanurine
affinity to be questionable. Dal Sasso et al. (2018) used Cau's
megamatrix to place Saltriovenator as a non-neoceratosaurian ceratosaur sister to Berberosaurus, while forcing it to be tetanurine took five more steps.
References- Dal Sasso, 2001. Update on Italian dinosaurs. 6th European
Workshop on Vertebrate Paleontology. Abstract volume, 27.
Dalla Vecchia, 2001. A new theropod dinosaur from the Lower Jurassic of Italy,
Saltriosaurus. Dino Press. 3, 81-87.
Dal Sasso, 2003. Dinosaurs of Italy. Comptes Rendus Palevol. 2(1), 45-66.
Dal Sasso, 2004. Dinosaurs of Italy. Indiana University Press. 213 pp.
Benson, 2010. The osteology of Magnosaurus nethercombensis (Dinosauria,
Theropoda) from the Bajocian (Middle Jurassic) of the United Kingdom and a re-examination
of the oldest records of tetanurans. Journal of Systematic Palaeontology. 8(1),
131-146.
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.
unnamed Ceratosauria (Galton, 1982)
Late Kimmeridgian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Colorado, US
Material- (DMNH 36284) proximal tibia (Chure, 2001)
?(USNM 8414) metatarsal III, metatarsal IV (Pickering, 1995)
(USNM 8415) humerus (201 mm) (Galton, 1982)
Middle Kimmeridgian, Late Jurassic
Salt Wash Member of the Morrison Formation, Wyoming, US
? material (Turner and Peterson, 1999)
Comments- The humerus USNM 8415 was discovered in 1883 and initially
referred to Dryosaurus, though Galton (1982) described it and referred
it to Elaphrosaurus sp. based on the straight shaft and low deltopectoral
crest. Pickering (1995b) referred it to his new taxon ?Elaphrosaurus
"philtippettensis" without justification. Neither listed any characters
to differentiate it from E. bambergi. Carrano and Sampson (2008) thought
the specimen was ceratosaurian, but could find no characters shared specifically
with Elaphrosaurus. Indeed, the straight shaft is present in all ceratosaurs,
while the low deltopectoral crest is present in Limusaurus and abelisaurians
as well. The proximal articular surface is wider than Limusaurus, but
less so than abelisaurians. The flattened distal condyles are also more derived
than Limusaurus, while Spinostropheus is intermediate. The internal
tuberosity is well developed as in Ceratosaurus and abelisaurids, but
unlike Limusaurus, Elaphrosaurus, Spinostropheus and Masiakasaurus.
The deltopectoral crest apex is placed more distally (42%) than Elaphrosaurus,
Limusaurus and especially Ceratosaurus and Masiakasaurus,
but is more proximal than abelisaurids. Based on these comparisons, I agree
the humerus cannot be assigned to Elaphrosaurus.
Rauhut and Carrano (2016) described further differences and proposed
"the lack of the abelisauroid synapomorphies of a distally placed
greater tubercle and the posterolateral tubercle might indicate that
this element represents a basal representative of the lineage leading
towards abelisauroids", while placing elaphrosaurs in the latter group.
USNM 8414 was discovered in 1883 and is assigned to Elaphrosaurus sp.
on the USNM collections website, though it has not been mentioned in the literature
to my knowledge. Pickering (1995b) referred it to his species Elaphrosaurus
"philtippettensis" without comment. Until these are illustrated or
described, their affinities remain unknown.
DMNH 36284 is a proximal tibia that was collected in 1992 and first published
in a faunal list by Carpenter (1998) as Elaphrosaurus sp.. Chure (2001)
later described it as Elaphrosaurus, though Carrano and Sampson (2008)
believed it resembled Tendaguru abelisauroid tibiae more.
Turner and Peterson (1999) listed Elaphrosaurus sp. from the Poison Creek
Quarry of the Morrison Formation in Wyoming (either from Erickson pers. comm.
1994 or Foster pers. comm. 1997), but this has yet to be described.
References- Galton, 1982. Elaphrosaurus, an ornithomimid dinosaur
from the Upper Jurassic of North America and Africa. Paläontologische Zeitschrift.
56, 265-275.
Pickering, 1995a. Jurassic Park: Unauthorized Jewish Fractals in Philopatry.
A Fractal Scaling in Dinosaurology Project, 2nd revised printing. Capitola,
California. 478 pp.
Pickering, 1995b. An extract from: Archosauromorpha: Cladistics and osteologies.
A Fractal Scaling in Dinosaurology Project. 2 pp.
Carpenter, 1998. Vertebrate biostratigraphy of the Morrison Formation near Canon
City, Colorado. Modern Geology. 23, 407-426.
Turner and Peterson, 1999. Biostratigraphy of dinosaurs in the Upper Jurassic
Morrison Formation of the Western Interior, U.S.A. In Gillette (ed.). Vertebrate
Paleontology in Utah. Utah Geological Survey Miscellaneous Publication. 99-1,
77-114.
Chure, 2001. The second record of the African theropod Elaphrosaurus
(Dinosauria, Ceratosauria) from the Western Hemisphere. Neues Jahrbuch für
Geologie und Paläontologie Monatshefte. 2001(9), 565-576.
Carrano and Sampson, 2008 (online 2007). The Phylogeny of Ceratosauria (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 6, 183-236.
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.
undescribed possible ceratosaur (Perez Garcia, Bolet, Escaso, Houssaye,
de Miguel Chaves, Mocho, Narvaez, Torices, Vidal and Ortega, 2015)
Late Campanian, Late Cretaceous
Armuna, Segovia, Spain
Material- teeth and postcrania
Comments- Said to be a "probable undetermined ceratosaur."
Reference- Perez Garcia, Bolet, Escaso, Houssaye, de Miguel Chaves, Mocho,
Narvaez, Torices, Vidal and Ortega, 2015. The vertebrate fauna from the Upper
Campanian site of Armuna (Segovia province, central Spain). Journal of Vertebrate
Paleontology. Program and Abstracts 2015, 193.
possible unnamed ceratosaur (Niedzwiedzki, Brusatte, Sulej and Butler, 2014)
Mid-Late Norian, Late Triassic
Poręba, Zbaszynek Beds, Poland
Material- (ZPAL V.39/45) distal tibia
Comments- Discovered between 2011 and 2012, Niedzwiedzki
et al. (2014) assign tibia ZPAL V.39/45 to ?Neotheropoda indet.. However, this specimen
differs from e.g. Coelophysis, Liliensternus and Dilophosaurus
in the poorly developed anterior diagonal tuberosity related to a
nearly vertical and shallow medial buttress for the astragalar
ascending process, a blunted posterolateral process in distal view, and
a more medially placed posteromedial ridge in distal view with flat
posterior and convex medial edges. Among dinosauromorphs this is most
similar to Berberosaurus,
which nonetheless differs in the nearly horizontal buttress, more
medially placed anterior diagonal tuberosity, anteriorly angled distal
edge to the posterolateral process, and more laterodistally projected
lateral malleolus. This is a potentially important specimen if
correctly identified as theropod.
Reference- Niedzwiedzki, Brusatte, Sulej and Butler, 2014. Basal dinosauriform and
theropod dinosaurs from the Mid-Late Norian (Late Triassic) of Poland:
Implications for Triassic dinosaur evolution and distribution.
Palaeontology. 57(6), 1121-1142.
unnamed possible ceratosaur (Serrano-Martinez, Ortega, Sciscio, Tent-Manclus,
Bandera and Knoll, 2015)
Bathonian-Oxfordian, Middle-Late Jurassic
Tiouraren Formation of the Irhazer Group, Niger
Material- (TP4-4) tooth (41.1x20.9x9.6 mm)
Comments- This grouped with Ceratosaurus when examined morphometrically,
and may belong to the MNN TIG6 elaphrosaur.
Reference- Serrano-Martinez, Ortega, Sciscio, Tent-Manclus, Bandera and
Knoll, 2015. New theropod remains from the Tiourarén Formation (?Middle
Jurassic, Niger) and their bearing on the dental evolution in basal tetanurans.
Proceedings of the Geologists' Association. 126(1), 107-118.
unnamed ceratosaur (Raath and McIntosh, 1987)
Tithonian, Late Jurassic
Kadze Formation, 10 km SE of Muzarabani, Zimbabwe
Material- (QG 65) two femora (~990 mm)
Comments-
Discovered between 1968 and 1972, Raath and McIntosh (1987) identified
these associated femora as ?allosaurid without rationale. Their
Table 7 gives a length of 885+ mm noting the distal end is damaged,
while the transverse proximal width is given as ~246 mm. Based on
their figure 4a, a complete length of 990 mm is reasonable.
Rauhut and Lopez-Arbarello (2008) stated they "might rather represent a
large neoceratosaur, as indicated by the low, aliform lesser trochanter
with a well-developed trochanteric shelf." Carrano et al. (2012)
stated "These femora are badly damaged but
exhibit several features that suggest affinities with Ceratosauria,
rather than Tetanurae. They are currently under study (E. Roberts, P.
O’Connor & M. Carrano in prep.)."
References- Raath and McIntosh, 1987. Sauropod dinosaurs from the central
Zambezi Valley, Zimbabwe, and the age of the Kadzi Formation. South African
Journal of Geology. 90(2), 107-119.
Rauhut and López-Arbarello, 2008. Archosaur evolution during the Jurassic:
A southern perspective. Revista de la Asociación Geológica Argentina.
63(4), 557-585.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae
(Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
unnamed possible ceratosaurian (Bonaparte, 1996)
Late Aptian-Albian, Early Cretaceous
Rayoso Formation, Neuquen, Argentina
Material- (Melo coll.) distal metatarsal III (~250 mm)
Comments- Assigned to ?Ceratosauria indet. by Bonaparte (1996).
Reference- Bonaparte, 1996. Cretaceous tetrapods of Argentina. Muncher
Geowissenschaftliche Abhandlung A. 30, 73-130.
Elaphrosaurinae Rauhut and Carrano, 2016
Definition- (Elaphrosaurus bambergi <- Noasaurus leali, Abelisaurus comahuensis, Ceratosaurus nasicornis, Allosaurus fragilis) (modified from Rauhut and Carrano, 2016)
Other definitions- (Elaphrosaurus bambergi <- Noasaurus leali) (modified from Delcourt, 2018)
= Elaphrosaurinae sensu Delcourt, 2018
Definition- (Elaphrosaurus bambergi <- Noasaurus leali) (modified)
References- 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, 546-610.
Delcourt, 2018. Ceratosaur palaeobiology: New insights on evolution and
ecology of the southern rulers. Scientific Reports. 8:9730.
Deltadromeus Sereno, Dutheil,
Iarochene, Larsson, Lyon, Magwene, Sidor, Varricchio and Wilson, 1996
D. agilis Sereno, Dutheil, Iarochene, Larsson, Lyon, Magwene,
Sidor, Varricchio and Wilson, 1996
Cenomanian, Late Cretaceous
Kem Kem beds, Morocco
Holotype- (UCRC PV11; = SGM-Din 2) (~8.1 m) partial cervical rib, two anterior dorsal
neural arches, two partial dorsal ribs, two gastralia, partial caudal vertebrae
3-17 (130 mm), caudal vertebrae 20-27 (130 mm), eight chevrons, proximal scapula,
incomplete coracoid, humerus (~328 mm), proximal radius, proximal
ulna, partial ilium, pubic fragments?, partial ischia, femur (740 mm), incomplete
tibia (~700 mm), fibula, partial astragalus, calcaneum, metatarsal
II (417 mm), phalanx II-1 (140 mm), pedal ungual II (80 mm), metatarsal III
(450 mm), phalanx III-1 (140 mm), metatarsal IV (400 mm), phalanx IV-1 (98 mm),
phalanx IV-3 (52 mm), phalanx IV-4 (37 mm), metatarsal V (100 mm)
Reffered- ?(JP Cr681) distal femur (Singer, 2015 online)
?(JP Cr685) metatarsal III (Singer, 2015 online)
(Piccini coll.; cast MPCM 13573) pedal ungual IV (70 mm) (Novas, Dalla Vecchia and Pais, 2005)
Cenomanian, Late Cretaceous
Baharija Formation, Egypt
Referred- (IPHG 1911 XII 32; destroyed) distal femur
....(IPHG 1912 VIII 69; destroyed) femur
(1.22 m)
Comments- Sereno et al. (1996) described Deltadromeus as a basal
coelurosaur, but Rauhut (2003) found it to be an ornithomimosaur. More recently,
Wilson et al. (2003) and Tortosa et al. (2014) recovered it as a noasaurid,
while Carrano and Sampson (2008) found it to be a basal ceratosaur outside of
Neoceratosauria.
Sereno et al. (1996) referred the Baharija 'IPHG 1912 VIII' (described by Stromer,
1934) to Deltadromeus, specifying a coracoid, pubes, femur, proximal
tibia and fibula as the material. Yet this specimen number corresponds to numerous
specimens described by Stromer. The coracoid IPHG 1912 VIII 60 was associated
with a scapula that shares that number, the femur is IPHG 1912 VII 69 based
on the size Sereno et al. reported, and the fibula must be IPHG 1912 VIII 70
as no others are reported. Yet the pectoral girdle was found in layer m while
the femur and fibula were found in layer p. The only proximal tibia reported
is IPHG 1912 VIII 78, which is far too small to belong with the other hindlimb
elements and from a different locality. Finally, the only pubes with that number
are IPHG 1912 VIII 81, which are from yet another locality and much smaller
than even the tibia. This materials list agrees with Carrano and Sampson, though
note contrary to their statement, it is not a "partial postcranial skeleton".
Stromer used the pubes as a paratype of Bahariasaurus, questionably referred
the pectoral girdle, femur and fibula to the taxon as they cannot be compared
to the holotype, and referred the tibia to aff. Erectopus. Thus all material
was not referred to Bahariasaurus, contra Sereno et al.. The Baharija
pectoral girdle actually lacks the anteroposterior expansion considered diagnostic
for Deltadromeus by Sereno (length excluding posteroventral process 117%
of height vs. 150% in Deltadromeus), which is also found in Elaphrosaurus
and Limusaurus. Due to breakage of the posteroventral process, it's uncertain
if the coracoid's subacromial notch ('notch in anterior margin' of Sereno et
al., as it is the only notch in Deltadromeus' coracoid) is shallow as
in Deltadromeus or deeper as in Elaphrosaurus and Limusaurus.
Though again, a shallow notch might not be diagnostic of Deltadromeus
as it is also found in Ceratosaurus. The pectoral girdles also differ
in other ways if scaled to similar overall size, with Deltadromeus having
a narrower scapular shaft, a more abruptly expanded acromion, smaller glenoid,
and deeper posteroventral process. IPHG 1912 VIII 60 most closely resembles
Baryonyx, so may be spinosaurid. Sereno et al. also diagnose Deltadromeus
based on its "accessory trochanter" on the distal femoral shaft, which
it turns out is the M. adductor femoris 2 insertion scar (Ibrahim et
al., 2020) that is prominently projected in IPHG 1912 VII 69.
Carrano and Sampson (2008) incorrectly equated the "accessory
trochanter" to the M. adductor femoris 1 insertion scar on the
posteromedial distal shaft, but this region is unillustrated in the
Baharija femur. IPHG 1912 VII 69 also has an anterior process on
the lateral margin of its medial condyle, stated as diagnostic of Deltadromeus and hinted at in Sereno
et al.'s skeletal reconstruction. Carrano and Sampson equated this with the
mediodistal crest, but that projects largely laterally so is not the feature Sereno et al. had in mind. Deltadromeus differs from the Baharija femur in having a fully medially oriented head and an anterior
trochanter that extends distally to the fourth trochanter, but these are known to vary in other theropods such as Megalosaurus and Allosaurus.
While the actual accessory trochanter (on the anterior edge of the
anterior trochanter) of IPHG 1912 VII 69 is like avetheropods
(unpreserved in Deltadromeus), the related Gualicho
also has this structure. Chiarenza and Cau (2016) noted
additional differences, but the robusticity of the Bajarija femur may
be ontogenetic, the proximodistally shorter laterodistal crest could be
individual variation, and the dorsal and medial surfaces of the head
are broken off in Deltadromeus
which leads to the illusion of a mediodorsally sloped proximal edge and
lack of a ventrally hooked head. The balance of evidence is seen
here as supporting a referral of IPHG 1912 VII 69 to Deltadromeus. The tibiae
are more similar to each other than to Elaphrosaurus, Ceratosaurus, Eoabelisaurus or Erectopus in proximal view (the
only available for Deltadromeus), with Deltadromeus differing
in having a smaller, triangular posterior groove and larger lateral condyle.
The fibulae are roughly similar, though Deltadromeus has a more projected
anteroproximal corner and a proximomedial fossa that is less proximally extensive.
The supposed pubes of Deltadromeus are actually ischia (see below), so
cannot be compared to the Baharija pubes. Thus in total, the pectoral girdle is near certainly not Deltadromeus
(contrary to Sereno et al.'s claim the remains are identical), the
femur shares apomorphies though possessing a couple differences, the
tibia and fibula could be although no described apomorphies are shared,
and the pubes cannot be compared. Stromer (1934) noted the
corresponding left femur was also discovered and its distal end was
collected and catalogued as IPHG 1911 XII 32.
The supposed pubis of Deltadromeus' holotype seems to be an ischium (Longrich,
DML 2000; Carrano and Sampson, 2008). The shape of the distal boot is almost
identical to IPHG 1912 VIII 82 (a pair of ischia referred to Theropoda indet.
by Stromer and also misidentified as pubes), except that it's slightly shorter
in the latter. As Longrich describes (pers. comm. posted here),
the anterior surface is transversely convex unlike pubes, the conjoined shafts
are narrow transversely instead of having a pubic apron, the boot is completely
fused, and he identified what appeared to be pubic fragments in the Deltadromeus
holotype. Evans et al. (2015) figure the femur in posterior view,
while Ibrahim et al. (2020) figure the pes in more detail. The
latter note "All of the material of Deltadromeus agilis is now fully prepared. Previously unprepared fragments include shaft pieces that have completed the humerus and fibula."
Novas et al. (2005) described MPCM 13573 as an abelisauroid pedal
ungual, and Smyth et al. (2020) find it "is a near mirror image of the
preserved digit IV ungual of Deltadromeus (NI pers. obs.)."
References- Stromer, 1934. Ergebnisse der Forschungsreisen Prof. E. Stromers
in den Wüsten Ägyptens. II. Wirbeltierreste der Baharije-Stufe (unterstes
Cenoman). 13. Dinosauria. Abhandlungen der Bayerischen Akademie der Wissenschaften
Mathematisch-naturwissenschaftliche Abteilung, Neue Folge. 22, 1-79.
Sereno, Dutheil, Iarochene, Larsson, Lyon, Magwene, Sidor, Varricchio and Wilson,
1996. Predatory dinosaurs from the Sahara and Late Cretaceous faunal differentiation.
Science. 272(5264), 986-991.
Longrich, DML 2000. https://web.archive.org/web/20160806054506/http://dml.cmnh.org/2000Nov/msg00067.html
Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs.
Special Papers in Palaeontology. 69, 1-213.
Wilson, Sereno, Srivastava, Bhatt, Khosla and Sahni, 2003. A new abelisaurid
(Dinosauria, Theropoda) from the Lameta Formation (Cretaceous, Maastrichtian)
of India. Contributions from the Museum of Paleontology. The University of Michigan.
31, 1-42.
Novas, Dalla Vecchia and Pais, 2005. Theropod pedal unguals from the
Late Cretaceous (Cenomanian) of Morocco, Africa. Revista del Museo
Argentino de Ciencias Naturales, nuevo serie. 7, 167-175.
Carrano and Sampson, 2008 (online 2007). The Phylogeny of Ceratosauria (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 6, 183-236.
Evans, Barrett, Brink and Carrano, 2015. Osteology and bone microstructure of
new, small theropod dinosaur material from the early Late Cretaceous of Morocco.
Gondwana Research. 27(3), 1034-1041.
Singer, 2015 online. JuraPark na tropie nowych dinozaurow z Maroka. https://web.archive.org/web/20151206224352/https://jurapark.pl/jurapark-na-tropie-nowych-dinozaurow-z-maroka/
Chiarenza and Cau, 2016. A large abelisaurid (Dinosauria, Theropoda) from Morocco
and comments on the Cenomanian theropods from North Africa. PeerJ. 4:e1754.
Ibrahim, Sereno, Varricchio, Martill, Dutheil, Unwin, Baidder, Larsson,
Zouhri and Kaoukaya, 2020. Geology and paleontology of the Upper
Cretaceous Kem Kem Group of eastern Morocco. ZooKeys. 928, 1-216.
Smyth, Ibrahim, Kao and Martill, 2020 (online 2019). Abelisauroid
cervical vertebrae from the Cretaceous Kem Kem beds of southern Morocco
and a review of Kem Kem abelisauroids. Cretaceous Research. 108, 104330.
Gualicho Apesteguía,
Smith, Juárez Valieri and Makovicky, 2016
= "Nototyrannus" Anonymous, online 2011 vide Apesteguía, Smith,
Juárez Valieri and Makovicky, 2016
G. shinyae Apesteguía, Smith, Juárez Valieri and
Makovicky, 2016
= "Nototyrannus violantei" Anonymous, online 2011 vide Apesteguía,
Smith, Juárez Valieri and Makovicky, 2016
Middle Cenomanian-Early Turonian, Late Cretaceous
Huincul Formation of the Rio Limay Subgroup, Rio Negro, Argentina
Holotype- (MPCN PV 0001) four incomplete dorsal centra (92, 104, ~80 mm),
16-17 partial gastralial rows, three incomplete mid caudal vertebrae (72, 83
mm), incomplete scapulocoracoid (scapula ~439 mm), humerus (286 mm), radii (155.4,
152.8 mm), ulnae (166.8 mm), radiale?, semilunate carpal, fused metacarpal I
and II (I- 50.3, II- 81.7 mm), phalanx I-1 (64.8 mm), manual ungual I (60.6
mm), phalanx II-1 (31.1 mm), phalanx II-2 (35.9 mm), incomplete manual ungual
II, incomplete metacarpal III, incomplete pubes (~355 mm), femora (one distal;
775 mm), proximal tibia, proximal fibula, distal metatarsal II, phalanx II-1
(95.3 mm), phalanx II-2 (64.3 mm), incomplete pedal ungual II, metatarsals III
(one distal; 309 mm), phalanx III-1 (104.3 mm), phalanx III-2 (79.1 mm), phalanx
III-3 (54.8 mm), pedal ungual III (58.1 mm), phalanx IV-1 (73.4 mm), phalanx
IV-2 (55.4 mm), phalanx IV-3 (54.1 mm), phalanx IV-4 (32.4 mm), pedal ungual
IV
Diagnosis- (after Apesteguia et al., 2016) Posterior dorsal vertebrae
very elongated and with slit-like pneumatic openings; scapular blade narrow
with sinuous dorsal margin marked by shallow notch between acromion and blade;
forelimb foreshortened with reduced muscle attachments and articulations; first
and second metacarpals coossified proximally, third metacarpal reduced to splint;
pubes with little or no pubic apron; pubes with blade-like boot; femur with
mediodorsally inclined head; reduced femoral distal condyles; fibula with large
fossa and accessory flange on proximoposterior corner; ridge-like m. iliofibularis
tubercle of fibula; third metatarsal antarctometatarsal; pedal unguals with
single claw sheath grooves that define small spur or tuber near proximal end.
Comments- The holotype was discovered in 2007 and partially excavated
by Apesteguia et al., but a team from the Museo Patagónico de Ciencias
Naturales and Museum of Sao Paulo improperly collected it in 2009 (Apesteguia,
online 2011). In 2011, Museo Patagonico announced the find to the media (Anonymous
online, 2011), though whether the name made it into any paper media is unknown.
Apesteguia et al. (2016) did list Nototyrannus violantei as a nomen nudum
which Gualicho shinyae replaced. The collection and respository issues
were resolved, resulting in the publication in 2016. Near simultaneously, Motta
et al. (2016) released the online version of their paper describing similar
remains from the same locality as Aoniraptor libertatum. Their synonymy
was quickly suggested (the first outside social media being Cau, 2016 online) and followed by many,
but Aranciaga Rolando et al. (2020) noted many differences in the comparable elements (mid caudals)
The phylogenetic affinities of Gualicho are highly controversial, but Apesteguia et al. (2016) seem to be correct in that it is closely related
to the similarly problematic Deltadromeus. The initial media announcement
proclaimed the specimen to be a tyrannosauroid without
a cladistic analysis, but noting the didactyl manus which is similar to tyrannosaurids. Apesteguia et al. found both Gualicho and Deltadromeus
emerged as a clade of non-megaraptoran neovenatorids, which they recovered as
carcharodontosaur carnosaurs in a modified version of Carrano et al.'s tetanurine
analysis. However, when added to a version of Novas et al.'s tetanurine analysis,
Gualicho and Chilantaisaurus formed a clade of basal coelurosaurs
closer to birds than carcharodontosaurids or Neovenator, but outside
the clade of Megaraptora+Tyrannoraptora. Neither analysis strongly sampled ceratosaurian
characters, which Apesteguia et al. reported several of in Gualicho,
and where Deltadromeus has been generally assigned since the mid-2000s.
Only future studies with more data will determine whether Gualicho and
Deltadromeus are ceratosaurs, carnosaurs or coelurosaurs, and whether
Bahariasaurus or Chilantaisaurus are closely related as well.
References- Anonymous, online 2011. Presentan
al "primo" del Tyrannosaurus. RioNegro.com.ar. 8-12-2011.
Apesteguia, online 2011. https://web.archive.org/web/20160806223425/http://dml.cmnh.org/2011Aug/msg00348.html
Apesteguía, Makovicky, Smith and Juárez Valieri, 2013. A new theropod
with a didactyl manus and African affinities from the Upper Cretaceous of Patagonia,
Argentina. VIII Congreso Latinoamericano de Paleontología and XIII Congreso
Mexicano de Paleontología. [pp]
Apesteguía, Smith, Juárez Valieri and Makovicky, 2016. An unusual
new theropod with a didactyl manus from the Upper Cretaceous of Patagonia, Argentina.
PLoS ONE. 11(7), e0157793.
Cau, online 2016. http://theropoda.blogspot.com/2016/07/nuovi-resti-di-aoniraptor-ehm-benvenuto.html
Aranciaga Rolando, Marsa and Novas, 2020. Histology and pneumaticity of Aoniraptor libertatem (Dinosauria, Theropoda), an enigmatic mid-sized megaraptoran from Patagonia. Journal of Anatomy. 237(4), 741-756.
Berthasaura de Souza, Soares, Weinschutz, Wilner, Lopes, de Araujo and Kellner, 2021
B. leopoldinae de Souza, Soares, Weinschutz, Wilner, Lopes, de Araujo and Kellner, 2021
Aptian-Albian, Early Cretaceous
Cemitério dos Pterossauros Quarry, Goio Erê Formation, Brazil
Holotype- (MN 7821-V) (~1 m)
(subadult) incomplete skull, sclerotic plate, mandible, dentary, hyoids
(59 mm), atlantal intercentrum (3 mm), axial intercentrum (3 mm), axis
(9 mm), ~fourth cervical vertebra (12 mm), ~sixth cervical vertebra,
~seventh cervical neural arch, ~tenth cervical vertebra (15 mm),
cervical ribs, first dorsal vertebra (13 mm), second dorsal vertebra,
third dorsal centrum (12 mm), fourth dorsal vertebra, fifth dorsal
vertebra, ?sixth dorsal centrum (18 mm), ?seventh dorsal vertebra (19
mm), ?eighth dorsal vertebra (19.2 mm), ?ninth dorsal vertebra (19 mm),
tenth dorsal vertebra, ?eleventh dorsal vertebra, dorsal ribs (dr1
72.3, dr2 75.7, dr3 78.1, dr4 63.3, dr5 80.7 mm), gastralia, first
sacral vertebra (16 mm), second sacral centrum (15.4 mm), third sacral
centrum (16 mm), fourth sacral centrum (16 mm), fifth sacral vertebra
(17 mm), sacral neural arch, first caudal vertebra (18 mm), second
caudal vertebra (16 mm), third caudal vertebra, fourth caudal vertebra
(15 mm), fifth caudal vertebra (16 mm), sixth caudal vertebra (15 mm),
proximal caudal centrum (17 mm), two mid caudal centra (13, 12 mm),
three mid caudal neural arches, two distal caudal centra (15, 15 mm),
two distal caudal neural arches, several chevrons (38.6, 38, 27.1, 23,
16.5 mm), scapulae (84.6, 81 mm), coracoid, humerus (58 mm), radius
(31.8 mm), ulna (27.5 mm), metacarpal II, metacarpal III (5.3 mm),
manual ungual (7.5 mm), ilium (115 mm), incomplete pubes, incomplete
ischia, femora (one incomplete; 132 mm), tibiae (one partial, one
proximal), fibulae (one incomplete; 102.3 mm), phalanx III-1 (24.3 mm),
pedal ungual (15.7 mm)
Diagnosis- (after de Souza et
al., 2021) premaxilla, maxilla and dentary toothless; posteroventral
premaxillary process; lamellae on medial surface of premaxilla; short
dentaries with length anterior to external mandibular fenestra ~1.5
times height of dentary; non-bifurcated anterior end of splenial;
postzygodiapophyseal lamina in mid cervical vertebrae divided in two
parts and reduced to low ridges; metacarpus <15% of humeral length;
iliac blade strongly flattened mediolaterally; medial brevis shelf
strongly reduced; deep notch on posterior margin of ischial peduncle
producing a posteriorly oriented prong; rounded medial femoral
epicondyle.
Comments- This specimen was
collected from 2011-2015. de Souza et al. (2021) used Rauhut and
Carrano's ceratosaur matrix to recover Berthasaura as the basalmost noasaurid, outside Noasaurinae and Elaphrosaurinae.
Reference- de Souza, Soares,
Weinschutz, Wilner, Lopes, de Araujo and Kellner, 2021. The first
edentulous ceratosaur from South America. Scientific Reports. 11:22281.
unnamed elaphrosaurine (Sereno, Conrad and Wilson, 2002)
Bathonian-Oxfordian, Middle-Late Jurassic
Tiouraren Formation of the Irhazer Group, Niger
Material- (MNN TIG6; from Fako) (~3 m) posterior third cervical vertebra, fourth
cervical vertebra, fifth cervical vertebra, sixth cervical vertebra, seventh
cervical vertebra, eighth cervical vertebra, ninth cervical vertebra, tenth
cervical vertebra, cervical rib, first dorsal vertebra, second dorsal vertebra,
third dorsal vertebra, fourth dorsal vertebra, fifth dorsal vertebra, sixth
dorsal vertebra, seventh dorsal vertebra, eighth dorsal vertebra, partial ninth
dorsal vertebra, partial tenth dorsal vertebra, partial eleventh dorsal vertebra,
partial twelfth dorsal vertebra, partial thirteenth dorsal vertebra, fragmentary
dorsal ribs, first sacral neural arch, second sacral neural arch, third sacral
neural arch, ossified tendons
Diagnosis- (after Sereno et al., 2004) strongly canted anterior articular
face on mid cervical centra (30 degree angle to posterior articular face); partitioned
anterior pleurocoels in mid-cervical centra; dorsoventrally flattened epipophyseal
processes on mid cervical vertebrae; broad subrectangular neural spines on mid
cervical vertebrae.
Comments- Though originally identified as Early Cretaceous (Lapparent,
1960), the Tiouraren Formation has been reinterpreted as Bathonian-Oxfordian
(Rauhut and Lopez-Arbarello, 2009).
Discovered in 1997 or 2000, this specimen was presented at SVP 2002 by
Sereno et al. as "the first abelisaurid" based on "hypertrophied
epipophyses, a strong interzygapophyseal lamina, and paired pneumatic
openings on the centra." Sereno et al. (2004) described this
briefly, referring it to Lapparent's (1960) Elaphrosaurus gautieri from the same formation but separating it as the new genus Spinostropheus. While broadly accepted by others, Sereno et al. never explicitly state characters uniting MNN TIG6 with any of the gautieri
syntypes. They say "cervical and dorsal vertebrae overlap", but
only the angle between anterior and posterior faces is shared between
the fifth cervical of MNN TIG6 and Sereno et al.'s "holotype" of gautieri
(actually a syntype; plate XI fig. 5 of Lapparent), while "partitioned
anterior pleurocoels" cannot be determined in either from the
literature and epipophysis and neural spine shape cannot be determined
in Lapparent's specimen based on breakage. Rauhut and Carrano
(2016) were the first to state "MNN TIG6 seems to be markedly different
from the original specimens of S. gautieri",
so treat them as different taxa "pending a review of all of this
material by Ronan Allain and Paul Sereno (R. Allain, pers. comm.
2014)." Of figured specimens, only the cervicals noted above can
be compared, where MNN TIG6 differs in having a more ventrally
projected diapophysis, narrow and posterodorsally concave parapophysis,
more elongate centrum (2.6 times posterior height vs. 1.9), posterior
pair of pleurocoels (Rauhut and Carrano), and deeper
infrapostzygapophyseal recess. The diapophyseal projection and
centrum elongation are compatable with MNN TIG6 being more anteriorly
positioned, but Sereno et al.'s figure indicates posterior pleurocoels
extend to the tenth cervical. Sereno et al. do not include any
remains except MNN TIG6 in their codings for 'Spinostropheus', and find it to be the sister
taxon to Abelisauria. Carrano and Sampson (2008) found it to be a basal
ceratosaur outside Neoceratosauria but do include some scores from
Lapparent's material.
References- Lapparent, 1960. Les dinosauriens du "Continental intercalaire"
du Sahara central. Memoirs of the Geological Society of France. 88A, 1-57.
Sereno, Conrad and Wilson, 2002. Abelisaurid theropods from Africa: Phylogenetic
and biogeographic implications. Journal of Vertebrate Paleontology. 22(3), 106A.
Sereno, Wilson and Conrad, 2004. New dinosaurs link southern landmasses in the
Mid-Cretaceous. Proceedings: Biological Sciences. 271(1546), 1325-1330.
Carrano and Sampson, 2008 (online 2007). The Phylogeny of Ceratosauria (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 6, 183-236.
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.
Elaphrosaurus Janensch, 1920
Not Elaphrosaurus- Stromer (1934) referred a proximal femur (IPHG
1911 XII 29) and two incomplete tibiae (IPHG 1912 VIII 76 and 192) from the
Baharija Formation of Egypt to cf. Elaphrosaurus bambergi, but these
belong to tetanurines based on the prominent accessory trochanter and distally
placed and well separated fibular crests. Galton (1982) referred a humerus (USNM
8415) from the Morrison Formation of Colorado to Elaphrosaurus sp., but
I agree with Carrano and Sampson (2008) and Rauhut and Carrano (2016) that it is not more similar to that
taxon than to other ceratosaurs. Pickering (1995a, b) created the nomina nuda
?Elaphrosaurus "philtippettensis" and E. "philtippettorum"
with the intended holotype as distal pubes USNM 5737 (also from the Morrison
Formation of Colorado), but I agree with Carpenter et al. (2005) that these
are more probably referrable to Tanycolagreus. He referred metatarsals
II and IV (USNM 8414) from the same locality as USNM 8415 to his species, but
these have yet to be described. Similarly, Turner and Peterson (1999) listed
Elaphrosaurus sp. from the Poison Creek Quarry of the Morrison Formation
in Wyoming, but this material remains undescribed. Chure (2001) described a
proximal tibia (DMNH 36284) from the Morrison of Colorado as Elaphrosaurus,
but Carrano and Sampson (2008) considered it to resemble unnamed Tendaguru abelisauroid
tibiae more. Pol and Rauhut (2012) consider the humerus and tibia to be ceratosaurs
more basal than Elaphrosaurus.
References- Stromer, 1934. Ergebnisse der Forschungsreisen Prof. E. Stromers
in den Wüsten Ägyptens. II. Wirbeltierreste der Baharije-Stufe (unterstes
Cenoman). 13. Dinosauria. Abhandlungen der Bayerischen Akademie der Wissenschaften
Mathematisch-naturwissenschaftliche Abteilung, Neue Folge. 22, 1-79.
Galton, 1982. Elaphrosaurus, an ornithomimid dinosaur from the Upper
Jurassic of North America and Africa. Paläontologische Zeitschrift. 56,
265-275.
Pickering, 1995a. Jurassic Park: Unauthorized Jewish Fractals in Philopatry.
A Fractal Scaling in Dinosaurology Project, 2nd revised printing. Capitola,
California. 478 pp.
Pickering, 1995b. An extract from: Archosauromorpha: Cladistics and osteologies.
A Fractal Scaling in Dinosaurology Project. 2 pp.
Turner and Peterson, 1999. Biostratigraphy of dinosaurs in the Upper Jurassic
Morrison Formation of the Western Interior, U.S.A. In Gillette (ed.). Vertebrate
Paleontology in Utah. Utah Geological Survey Miscellaneous Publication. 99-1,
77-114.
Chure, 2001. The second record of the African theropod Elaphrosaurus
(Dinosauria, Ceratosauria) from the Western Hemisphere. Neues Jahrbuch für
Geologie und Paläontologie Monatshefte. 2001(9), 565-576.
Carrano and Sampson, 2008 (online 2007). The Phylogeny of Ceratosauria (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 6, 183-236.
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.
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.
E. bambergi Janensch, 1920
Late Kimmeridgian, Late Jurassic
Quarry dd, Middle Dinosaur Member of the Tendaguru Formation, Tanzania
Holotype-
(HMN MB R 4960; = HMN Gr.S. 38-44) (6.2 m, 370 kg, old adult) third
cervical vertebra (~77 mm), fifth cervical vertebra (~114 mm),
incomplete sixth cervical vertebra (~120 mm), incomplete seventh
cervical vertebra (121 mm), incomplete eighth cervical vertebra (119
mm), incomplete ninth cervical vertebra (112 mm), tenth cervical
vertebra (99 mm), incomplete first dorsal vertebra (84 mm), incomplete
second dorsal vertebra (~85 mm), incomplete third dorsal vertebra (83
mm), incomplete fourth dorsal vertebra (84 mm), incomplete fifth dorsal
vertebra (88 mm), ninth dorsal vertebra, tenth dorsal vertebra (~108
mm), eleventh dorsal vertebra (103 mm), incomplete twelfth dorsal
vertebra (96 mm), four dorsal rib fragments (three lost), synsacrum
(88, 76, 50, 42, 49, 64 mm), incomplete ~first caudal vertebra (77 mm),
incomplete ~second caudal vertebra (78 mm), incomplete ~fifth caudal
vertebra (80 mm), incomplete ~sixth caudal vertebra (76 mm), incomplete
~seventh caudal vertebra (70 mm), incomplete ~tenth caudal vertebra (73
mm), incomplete ~eleventh caudal vertebra (73 mm), incomplete
~fourteenth caudal vertebra (71 mm), ~seventeenth caudal vertebra (70
mm), ~nineteenth caudal vertebra (71 mm), ~twenty-third caudal vertebra
(82 mm), ~twenty-fifth caudal vertebra (84 mm), ~twenty-sixth caudal
vertebra (83 mm), ~twenty-eighth caudal vertebra (80 mm), ~twenty-ninth
caudal vertebra (79 mm), ~thirty-second caudal vertebra (83 mm),
~thirty-fifth caudal vertebra, ~thirty-eighth caudal vertebra,
incomplete chevron, incomplete scapulocoracoids, humerus (262 mm),
metacarpal II (51 mm), metacarpal IV (39 mm), incomplete ilia (~380
mm), proximal pubis, ischia (354 mm), femur (529 mm), tibia (608 mm),
incomplete fibula, astragalocalcaneum, metatarsal II (378 mm), phalanx
II-1 (100 mm), metatarsal III (391 mm), proximal metatarsal IV, phalanx
IV-2 (60 mm), phalanx IV-4 (36 mm)
Late Kimmeridgian, Late Jurassic
Quarry Ig, Middle Dinosaur Member of the Tendaguru Formation, Tanzania
Referred- ?(HMN coll.; lost) ninth dorsal vertebra (70 mm) (Janensch, 1925)
Diagnosis- (modified after Rauhut, 2000) pronounced ventrolateral laminae at the posterior ends of the cervical vertebrae; cervical
vertebrae strongly concave ventrally, the ventral margin arching above the mid-height
of the anterior articular facet at its highest point; brevis fossa of ilium
extremely widened, so that the brevis shelf forms an almost horizontal lateral
flange; distal end of ischium strongly expanded into a triangular boot.
(after Rauhut and Carrano, 2016) cervical pre- and postzygapophyses
narrow, more than 1.5 times longer than wide; cervical epipophyses
absent; distal end of metacarpal II offset ventrally from shaft by
distinct step; large posterior flange on the posterior side of the
ischia medially; extremely small ascending process of the astragalus,
extending for only ~3% of tibial length of the tibia(?); proximal end
of metatarsal IV almost 2.5 times deeper anteroposteriorly than wide
transversely.
Comments- The Elaphrosaurus
holotype was excavated from 1909 to 1913 and initially described by
Janensch (1920) before being described in more detail by him in
1925. Janensch (1929) reported that "after the skeleton had been
set up, a few bones came to light during further processing of the very
large material from the excavation site dd, which ... probably belong
to that skeleton to which they certainly fit in size"
(translated). This consisted of two dorsal rib fragments and
incomplete scapulocoracoids. After being mounted and thus
difficult to study for several decades, Rauhut began restudying the
holotype (Rauhut, 1998), which was redescribed in detail by Rauhut and
Carrano (2016). Rauhut and Carrano thought cervical four was
missing instead of cervical eight unlike Janensch, reidentified
Janensch's metacarpal I as metacarpal II, and his pedal phalanx II-2 as
IV-2.
Janensch (1920) referred about 150 teeth to Elaphrosaurus
from the Middle and Upper Dinosaur Members, narrowing that sample to
those he termed types a, b and c in 1925. These are typical small
theropod teeth 20-40 mm long with serrated mesial and distal carinae,
with type B being labiolingually ticker and thus more anterior, and
type c being J-shaped and thus premaxillary. While adult Limusaurus
are toothless, younger individuals have teeth although they are as of
yet poorly figured and undescribed. Most Tendaguru theropod teeth
have not been reevaluated in a modern context, and these are referred
to Averostra indet. here. Janensch (1925) referred a posterior
dorsal from contemporaneous Quarry Ig to Elaphrosaurus,
saying it "corresponds in all aspects with the ninth vertebra of the
skeleton; but it is only 3/4 the size" (translated). Rauhut and
Carrano (2016) "could not locate the isolated vertebra in the
collections in
Berlin, and it seems to be lost." Janensch also refers a manual
phalanx (HMN M.B.R. 1762) from the same quarry to the taxon, but Rauhut
and Carrano state "as there is no overlap with the type specimen, its
referral to Elaphrosaurus
cannot be tested, and therefore we do not consider it further
here." Based on its elongation it is here placed in
Maniraptoriformes indet.. Janensch (1929) referred a radius HMN
M.B.R. 1755 from the later Upper Dinosaur Member to Elaphrosaurus,
but Rauhut and Carrano rejected this as there is no overlap with the
holotype and ceratosaurs would be expected to have shorter radii.
Carrano and Sampson (2008) stated "HMN MB.R.1756, a distal left
ischium, may also pertain to this taxon." This element, also from
the Upper Dinosaur Member, was figured by Rauhut (2005) who referred it
to Abelisauroidea and suggested it may belong to the same individual as
tibia HMN MB.R.1751. It is roughly similar to Elaphrosaurus but has a much deeper shaft relative to ischial foot depth.
Janensch (1920) initially referred Elaphrosaurus
to Coelurosauria sensu Huene, stating (translated) "in the formation of
the metatarsus there is to a certain extent an analogy with the younger
genera Ornithomimus and Struthiomimus,
insofar as in these the proximal ends of Mt 2 and Mt 4 also come closer
together", but that "a direct or even more distant direct genetic
connection between the African form and Ornithomimus
cannot be proven due to the lack of intermediate forms, although it is
by no means excluded." Janensch (1925) later said (translated)
"in the build of the metatarsus of Elaphrosaurus one could see the beginning of a development that would lead to Struthiomimus. But ... it could be a product of convergence through similar function" and that "Elaphrosaurus shows ... the lamella-like, wing-shaped form of the transverse processes that are already developed on" Pterospondylus and Podokesaurus." While this is prescient of our current ideas, despite saying "the investigation of any relationship to Coelurus, Ornitholestes, and Compsognathus would need to rely heavily on the skull, which is not known in Elaphrosaurus",
Janensch ends up tentatively referring his taxon to Coeluridae.
Nopcsa (1928) first referred it to his Ornithomiminae, and while often
placed in Coeluridae it was generally seen as an intermediate between
that family and ornithomimids in precladistic studies (e.g. Russell,
1972). Paul (1988) was the first to argue for a position outside
Tetanurae (his Avetheropoda) based on "the low ankle process and a very
robust central cannon bone", instead placing it in Coelophysoidea (his
Coelophysidae). Holtz (1994) was the first to include Elaphrosaurus
in a phylogenetic analysis, recovering it as a ceratosaur closer to
abelisaurids than ceratosaurids. This is still a popular position
(e.g. Rauhut and Carrano, 2016), although some studies find it outside
the ceratosaurid+abelisaurid clade instead (e.g. Wang et al., 2017).
References- Janensch, 1920. Ueber Elaphrosaurus bambergi und die
Megalosaurier aus den Tendaguru-Schichten Deutsch-Ostafrikas. Sitzungsberichte
der Gesellschaft Naturforschender Freunde zu Berlin. 1920, 225-235.
Janensch, 1925. Die Coelurosaurier und Theropoden der Tendaguru-Schichten Deutsch-Ostafrikas.
Palaeontographica. (Supp. 7)1, 1-99.
Nopcsa, 1928. The genera of reptiles. Palaeobiologica. 1, 163-188.
Janensch, 1929. Ein aufgestelltes und rekonstruiertes Skelett von Elaphrosaurus
bambergi mit einem Nachtrag zur Osteologie dieses Coelurosauriers. Palaeontographica
(Supp. 7)1, 279-286.
Russell, 1972. Ostrich dinosaurs from the Late Cretaceous of western Canada. Canadian
Journal of Earth Sciences. 9(4), 375-402.
Galton, 1982. Elaphrosaurus, an ornithomimid dinosaur from the Upper
Jurassic of North America and Africa. Paläontologische Zeitschrift. 56,
265-275.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster.
464 pp.
Rauhut, 1998. Elaphrosaurus bambergi and the early evolution of theropod
dinosaurs. Journal of Vertebrate Paleontology. 18(3), 71A.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria,
Saurischia). PhD thesis. University of Bristol. 440 pp.
Rauhut, 2005. Post-cranial remains of 'coelurosaurs' (Dinosauria, Theropoda)
from the Late Jurassic of Tanzania. Geological Magazine. 142(1), 97-107.
Carrano and Sampson, 2008 (online 2007). The Phylogeny of Ceratosauria (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 6, 183-236.
Carrano and Rauhut, 2016. The theropod dinosaur Elaphrosaurus bambergi Janensch, 1920, from the Late Jurassic of Tendaguru, Tanzania. Journal of Vertebrate Paleontology. Program and Abstracts, 110.
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.
Wang, Stiegler, Amiot, Wang, Du, Clark and Xu, 2017 (online 2016). Extreme ontogenetic
changes in a ceratosaurian theropod. Current Biology. 27(1), 144-148.
Huinculsaurus Baiano, Coria and Cau, 2020
H. montesi Baiano, Coria and Cau, 2020
Middle Cenomanian-Early Turonian, Late Cretaceous
Huincul Formation of Rio Limay Subgroup, Rio Negro, Argentina
Holotype- (juvenile to subadult) partial eleventh dorsal
vertebra, twelfth dorsal vertebra (47.5 mm), thirteenth dorsal vertebra
(52.2 mm), fused first (50.6 mm) and fragmentary second sacral
vertebrae
Diagnosis- (after Baiano et
al., 2020) prezygapophyseal articular facets in posterior dorsal
vertebrae twice longer than wide; anterior centroparapophyseal lamina
strongly developed as an extensive lateral lamina in posterior dorsal
vertebrae; pneumatic foramina located ventrally to the
postzygodiapophyseal lamina in posterior dorsal vertebrae; posteriorly
tapering postzygapophysis pointed posteriorly; posterior dorsal neural
arches with the parapophyseal centrodiapophyseal fossa divoded in two
by an accessory lamina (also in Carnotaurus).
Comments- This was discovered in 1992 (Baiano pers. comm. to Cau, 2020 online).
Baiano et al. (2020) add this to Wang et al.'s basal theropod analysis and Cau's megamatrix, where it emerged sister to Elaphrosaurus in both.
References- Baiano, Coria and Cau, 2020. A new abelisauroid
(Dinosauria: Theropoda) from the Huincul Formation (lower Upper
Cretaceous, Neuquén Basin) of Patagonia, Argentina. Cretaceous
Research. 110, 104408.
Cau, 2020 online. http://theropoda.blogspot.com/2020/02/huinculsaurus-montesi-baiano-coria-e.html
unnamed elaphrosaurine (He, 1984)
Bajocian, Middle Jurassic
Laoshangou Quarry, Golden Rooster Commune, Xiashaximiao Formation, Sichuan, China
Material- (CUT 20011; = CCG 20011; paratype of Chuandongocoelurus primitivus)
(~5 meters, ~175 kg, subadult) ~third-fourth cervical vertebra (63 mm),
tenth cervical vertebra (74 mm), third dorsal centrum (65 mm), ~sixth
dorsal centrum (69 mm), mid dorsal neural arch, first sacral centrum
(60 mm), ~first caudal centrum (63 mm), ~third caudal centrum (55 mm),
incomplete ~sixth caudal vertebra (53 mm), four distal caudal vertebrae
(66, 54, 54, 54 mm), partial scapula (282 mm; lost), proximal pedal
ungual
Diagnosis- lateral depression of third cervical centrum extends over
60% of central length; scapula less than 5.5 times longer than wide.
Description- Of the two cervical vertebrae preserved in the paratype, one is clearly from
a very anterior position due to its slender centrum (ventral length 2.9 times
posterior height). Glut (1997) refers to an axis, but the strong parapophyses
suggest it was a third cervical instead, as basal theropods have very reduced
axial parapophyses. The centrum is platycoelous or amphiplatyan, with an anterior
face 38% wider than tall. There is a deep lateral fossa extending from near
the posterior border of the centrum to beneath the diapophyses, though whether
this contained foramina is uncertain. The diapophyses extend ventrolaterally
to almost contact the parapophyses, thus the ribs were not fused to the vertebra.
A dorsal fossa on the diapophyses as is present in basal ceratosaurs seems to
be present. A posterodorsally projecting posterior infradiapophyseal lamina
is present, and there may be a low angled bump on the posterior neural arch
edge. The latter may also be due to breakage though. The prezygopophyses are
broken off, but thir bases show they were massive. The posyzygopophyses were
more slender, and are broken so that epipophyseal morphology is unknown. The
neural spine is not well preserved, but was low and rounded.
The other cervical is from the posterior portion of the series as seen by the
postzygopophyses extending past the centrum and the craniocaudally short neural
spine. Comparison with Elaphrosaurus suggests it is the tenth. The centrum
is again elongate (2.6 times posterior height), with a slight lateral depression.
The anterior face is perhaps slightly convex and 38% wider than tall, while
the posterior face is flat or concave. The parapophyses are massive and positioned
on the anteroventral corners. There is a circular neural canal, a bit less than
40% as tall as the central face. The prezygopophyses are quite massive compared
to Elaphrosaurus, though broken at their tips. The ventrolaterally projecting
diapophyses end much further from the parapophyses than in the third cervical.
The cervical ribs were apparently unfused to the vertebrae. The neural spine
is low and short craniocaudally, although only its base remains. There is a
large postzygopophyseal-central choana again, but no step this time. There is
no evidence of epipophyses.
A centrum is probably from the third dorsal, as the parapophyses are partially
on the centrum and partially on the neural arch, as seen in the third dorsals
of Elaphrosaurus and Dilophosaurus. There is a slight lateral
depression, the anterior face is 14% wider than tall and it looks slightly opisthocoelous.
The other dorsal has a parapophysis placed on the neural arch, but not at the
dorsal edge of the prezygopophysis. This is seen in the fourth dorsal of Elaphrosaurus.
The centrum is again rather elongate, with a small lateral depression and indeterminate
face convexity. The ventral edge is more strongly concave than Elaphrosaurus.
The diapophyses project laterally and appear backswept. The prezygopophyses
are very short, but still more massive than Elaphrosaurus. A large postzygopophyseal-central
choana is still present, with a step like the third cervical. There are large
postzygopophyses and a moderate sized rectangular neural spine, with ventral
margins sloping towards the zygopophyses, especially the postzygopophyses.
The caudal vertebra is probably from around the fifth position, judging by elongation.
The centrum has no lateral depression and a moderately concave ventral surface.
The anterior face is perhaps concave while the posterior is slightly convex.
There are prominent transverse processes and the bases of large prezygopophyses.
The neural spine is craniocaudally expansive and there looks to be a small anterior
spine medial to the prezygopophyses. Postzygopophyses are broken off.
The scapula is very broad for a theropod (~5.1 times longer than broad), which
is more than even abelisaurs and megalosaurs (~5.9 times). However, it lacks
the expanded distal end of coelophysoids, Dilophosaurus and basal non-theropod
dinosaurs and is thus still strap-like. Most of the anterior edge is lacking,
but it is generally similar to Carnotaurus, differing in the slightly
concave posterior margin. An extensive posteriorly facing glenoid is present.
Comments- Discovered in 1970 (Kuang, 2004), He (1984) based his
new taxon Chuandongocoelurus on two specimens, the holotype (CCG 20010)
considered a juvenile and the much larger paratype considered an adult
or subadult. Stiegler (2019) notes "there is no overlap in preserved
elements of the holotype and paratype (CUT 20011) with the possible
exceptions of a sacral centrum and a pedal ungual, depending on
positional identity." Thus whether the paratype belongs to Chuandongocoelurus cannot be determined directly. CUT 20011 can be estimated to be
~5 meters long based on comparing presacral lengths with Elaphrosaurus.
The resulting weight estimate is ~175 kilograms. The unfused
neurocentral sutures in the dorsal vertebrae show He was correct it was
immature, so this was not its maximum size. Unfortunately, Stiegler
states "neither the scapula nor any other forelimb material (which was
mentioned by He, but not described or figured) could be located during
this study." While Norman (1990) claims "This material is
insufficient to provide diagnostic characters that justify a binomial",
Stiegler (2019) finds it "likely represents a new taxon, but is not
named in this dissertation." A new genus name is used in Figure
5.13 and 5.15, but this is omitted here as it has not been published.
He (1984) referred Chuandongocoelurus to Coeluridae, but viewed it as a
primitive member of that family sharing characters with the supposedly
ancestral podokesaurids, with the amphicoelous cervicals noted as more
similar to Coelophysis than Coelurus (based on YPM 1996-1997). Of
the paratype material, Norman (1990) merely said "The scapula appears
to be unusually broad in lateral aspect, instead of the straplike form
of theropods generally." In 2001, I noted the resemblence between
the Chuandongocoelurus material
and Elaphrosaurus,
(Mortimer, DML 2001) though I was unaware of the fact it belonged to
two individuals. I suggested "Compared to Ceratosaurus and Carnotaurus,
Elaphrosaurus and Chuandongocoelurus share the following
synapomorphies- elongate anterior dorsal centra (posterior central face
height <65% of central length); anterior cervicals with low rounded
neural spines; proximal caudal neural spines elongate
anteroposteriorly, extending over ~2/3 of central length" and thus
"Chuandongocoelurus is a neoceratosaur most closely related to
Elaphrosaurus, although how closely related other noasaurids are to
either of them is difficult to determine." Rauhut and
Lopez-Arbarello (2009) wrote "Chuandongocoelurus also includes material
that shows ceratosaurian or abelisauroid affinities (O. Rauhut,
personal observation, 2006)", while Carrano et al. (2012) stated "CCG
20011 possesses anteroposteriorly long cervical centra with
anteroposteriorly elongate pleurocoelous fossa, similar to those seen
in the ceratosaur Elaphrosaurus." Rauhut and Carrano (2016) agree
"The cervical vertebrae are remarkably similar to those of Elaphrosaurus"
"on the basis of the personal observations made by of one of us (O.R.)
on the actual specimen" and recover it as an elaphrosaurine noasaurid
closer to Elaphrosaurus than to Limusaurus. Stiegl;er (2019) used Wang's ceratosaur matrix to recover it as a basal noasaurine noasaurid.
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.
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.
Rauhut and Lopez-Arbarello, 2009 (online 2008). Considerations on
the age of the Tiouaren Formation (Iullemmeden Basin, Niger, Africa):
Implications for Gondwanan Mesozoic terrestrial vertebrate faunas.
Palaeogeography, Palaeoclimatology, Palaeoecology. 271 (3-4), 259-267.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
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.
Stiegler, 2019. Anatomy, systematics, and paleobiology of noasaurid
ceratosaurs from the Late Jurassic of China. PhD thesis, The George
Washington University. 693 pp.
Limusaurus Xu, Clark,
Mo, Choiniere, Forster, Erickson, Hone, Sullivan, Eberth, Nesbitt, Zhao, Hernandez,
Jia, Han and Guo, 2009
L. inextricabilis Xu, Clark, Mo, Choiniere, Forster, Erickson,
Hone, Sullivan, Eberth, Nesbitt, Zhao, Hernandez, Jia, Han and Guo, 2009
Late Oxfordian, Late Jurassic
TBB 2005, Wucaiwan, Upper Shishugou Formation, Xinjiang, China
Holotype-
(IVPP V15923) (~1.7 m; 2-3 year old subadult) incomplete skull,
sclerotic ring, mandible (105.44 mm), hyoid, axis, third-tenth cervical
vertebrae (c7- ~44.60 mm), cervical ribs, first-fifth dorsal centra
(d4- 26 mm), dorsal ribs, gastralia, sacrum, first-eleventh caudal
vertebrae (c1- 23.94, c2- 27, c12- 26 mm), eleven chevrons, scapula
(108.60 mm), coracoid, furcula, humeri (83.20 mm), radii (45.20 mm),
ulnae (44.00 mm), metacarpals II (13.12 mm), phalanx II-1, phalanx
II-2, proximal manual ungual II, metacarpals III (13.10 mm), phalanx
III-1, metacarpal IV, ilia (142.64 mm), pubis (~185.00 mm), ischium
(133.95 mm), femur (210.62 mm), tibiae (249.35 mm), fibula,
astragalocalcaneum, distal tarsals III, distal tarsals IV, metatarsals
I, phalanges I-1, pedal unguals I, metatarsals II, phalanges II-1,
phalanges II-2, pedal unguals II, metatarsals III (158.96 mm),
phalanges III-1 (36 mm), phalanges III-2 (26 mm), phalanges III-3 (20
mm), pedal unguals III (21 mm), metatarsals IV, phalanges IV-1,
phalanges IV-2, phalanges IV-3, phalanges IV-4, pedal unguals IV,
metatarsal V, gastroliths
Referred- (IVPP V20093) (<1
year old juvenile) skull, mandible, hyoid(?), axis, ~third-seventh cervical vertebrae, dorsal
vertebrae, sacrum, ~first-thirtieth caudal vertebrae, few chevrons, pectoral girdle, ilium, pubis, ischium,
femur, tibia (52.90 mm), three metatarsals, few pedal phalanges (Wang, Stiegler, Amiot,
Wang, Du, Clark and Xu, 2017)
(IVPP V20094) (<1 year old juvenile) skull, mandible (Wang, Stiegler, Amiot, Wang, Du, Clark and Xu, 2017)
(IVPP
V20095) (<1 year old juvenile) skull, mandible, several anterior
cervical
vertebrae, partial ?ilium, proximal ?pubes, femur, tibiae, proximal
fibula, fragments (Wang, Stiegler, Amiot, Wang, Du, Clark and Xu,
2017)
(IVPP V20096) (4 year old subadult) premaxillae, maxillae, nasals,
lacrimals, jugal, quadrates, parabasisphenoid, partial prootic,
palatine, pterygoids, sclerotic ring, dentary, fragmentary postdentary
mandibular elements, postcranial skeleton including axis, third-tenth
cervical vertebrae, cervical ribs, dorsal ribs, first sacral centrum,
incomplete synsacrum, first-thirty-second caudal vertebrae (three
distal caudals lost), scapula, ilia (164.90 mm), ischium (264.78 mm),
femur, tibia (289.07 mm), metatarsal III (~181.81 mm) and pedal unguals
(Wang, Stiegler, Amiot, Wang, Du, Clark and Xu, 2017)
(IVPP V20097) (subadult or adult) seven dorsal vertebrae, partial dorsal ribs,
scapula (Wang, Stiegler, Amiot, Wang, Du, Clark and Xu, 2017)
(IVPP V20098) (1 year old juvenile) posterior dorsal vertebrae,
gastralia, most caudal vertebrae, pelvis including ilium (70.60 mm),
hindlimbs including femur (100.88 mm), tibia (123.49 mm) and metatarsal
III (76.58 mm), gastroliths (Wang, Stiegler, Amiot, Wang, Du, Clark and
Xu, 2017)
(IVPP V20099) (>6 year old adult) squamosal(?), third-tenth cervical
vertebrae, posterior dorsal vertebrae, partial dorsal ribs, gastralia,
second sacral centrum, incomplete synsacrum, sternum, five sternal
ribs, partial forelimb including partial or fragmentary manus, partial
ilia, incomplete pubes, femur, tibia (300.00 mm), fibula, metatarsal
II, phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III,
phalanx III-1, phal;anx III-2, phalanx III-3, pedal ungual III,
metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4,
gastroliths (Wang, Stiegler, Amiot, Wang, Du, Clark and Xu, 2017)
(IVPP V20100) (<1 year old juvenile) anterior skull, frontals,
fourth-tenth
cervical vertebrae (c7 8.91 mm), cervical ribs, first dorsal vertebra,
third-twelfth dorsal vertebrae, partial dorsal ribs, fragmentary
sacrum, first-fifth caudal vertebra (c1 8.02 mm),
scapulae (30.79 mm), coracoids, humeri (24.32 mm), radii (~15.13 mm),
ulnae (15.44 mm), metacarpal III (3.52 mm), manual fragments,
fragmentary ilium (Wang, Stiegler, Amiot, Wang, Du, Clark and Xu, 2017)
(IVPP V20103) (subadult) incomplete cervical series, two posterior dorsal centra,
dorsal ribs, gastralia, first-eleventh caudal vertebrae, ten chevrons, scapula,
humerus, partial ilium, femur, tibiae, fibula, two metatarsals, pedal phalanges, gastroliths
(Wang, Stiegler, Amiot, Wang, Du, Clark and Xu, 2017)
(IVPP V20104) (subadult) two posterior dorsal vertebrae, dorsal rib
fragment, about
thirty-three caudal vertebrae, chevrons, incomplete ilia, femur, tibia,
astragalus, distal tarsals, metatarsal I, metatarsal II, phalanx II-1,
phalanx II-2, pedal ungual II, metatarsal III, phalanx III-1, phalanx
III-2, phalanx III-3, pedal ungual III (Wang, Stiegler, Amiot,
Wang, Du, Clark and Xu, 2017)
(IVPP V20105) (subadult) axis, third-tenth cervical vertebrae, cervical ribs (sixth-eighth
fused with vertebrae), first-eighth dorsal vertebrae, dorsal ribs, scapulae,
humerus, radius, ulna, phalanx III-1, phalanx III-2 and
manual ungual III (Wang, Stiegler, Amiot, Wang, Du, Clark and Xu, 2017)
(IVPP uncatalogued) pubis, femur (Stiegler, 2019)
Late Oxfordian, Late Jurassic
TBB 2002, Wucaiwan, Upper Shishugou Formation, Xinjiang, China
Paratype-
(IVPP V15304; = IVPP V15924 of Xu et al., 2009) (~2 m; 4-5 year old
subadult) frontals, parietal, laterosphenoids, two sclerotic ossicles?,
six cervical vertebrae, cervical ribs, several dorsal vertebrae, dorsal
ribs, gastralia, first-sixth caudal vertebrae (c1 30.80 mm), five
chevrons, furcula, partial coracoid, humeri (108.02 mm), radius (53.06
mm), ulna (51.43 mm), metacarpal I, metacarpal II (16.24 mm), phalanx
II-1, phalanx II-2, manual ungual II, metacarpal III (16.15 mm),
phalanx III-1, phalanx III-2, manual ungual III, proximal metacarpal
IV, ilia, pubes (238.35 mm), ischia (157.62 mm), femora,
tibiofibulotarsi (283.86 mm), distal tarsals III, distal tarsals IV,
metatarsal I, metatarsal II, phalanges II-=1, phalanges II-2, pedal
unguals II, metatarsal III, phalanges III-1, phalanges III-2, phalanges
III-3, pedal unguals III, metatarsal IV, phalanges IV-1, phalanges
IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, metatarsals V,
gastroliths
Referred- (IVPP V15303) (4 year
old subadult) fifth-tenth cervical vertebrae, dorsal vertebrae
including first, proximal dorsal ribs, sacrum, proximal caudal
vertebrae, forelimbs (lost), ilia, pubis, femora, tibia (245.60 mm),
fibula, astragalocalcaneum, distal tarsal III, distal tarsal IV,
metatarsus, pedal digit II, pedal digit III, pedal digit IV (Eberth, Xu
and Clark, 2010)
Late Oxfordian, Late Jurassic
TBB 2001, Wucaiwan, Upper Shishugou Formation, Xinjiang, China
Paratype-
(IVPP V15297; = IVPP V16134 of Xu et al., 2009) (~1.7 m, 6 year old
subadult) anterior skull, fifth-tenth cervical vertebrae, cervical
ribs, tenth-twelfth dorsal vertebrae, few dorsal ribs, synsacrum,
first-twenty-fourth caudal vertebrae, three chevrons, partial coracoid,
some forelimb elements (lost), radius, ulna, radiale(?),
intermedium(?), metacarpal I, metacarpal II (15.22 mm), metacarpal III
(15.22 mm), pelves including ilia, femora (233.71 mm), tibia (295.72
mm), fibula, astragalus, calcaneum, metatarsi, pedal digit II, pedal
digits III, pedal digit IV, gastroliths
Referred-
(IVPP V15298) (adult) distal tarsals, metatarsal I, phalanx I-1, pedal
ungual I, metatarsal II, phalanx II-1, phalanx II-2, pedal ungual II,
metatarsal III, phalanx III-1, phalanx III-2, phalanx III-3, pedal
ungual III, metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-3,
phalanx IV-4, incomplete pedal ungual IV (Eberth, Xu and Clark, 2010)
(IVPP V15299) (5 year old subadult) ninth-twelfth dorsal vertebrae,
sacrum, first-~twenty-fourth caudal vertebrae, >fourteen chevrons,
pectoral girdle, ilia, pubis, ischium, femur, tibiae (270.2 mm),
fibulae, astragalus, calcaneum, distal tarsal III, distal tarsal IV,
metatarsal I, metatarsal II, phalanx II-1, phalanx II-2, metatarsal
III, phalanges III-1, phalanges III-2, phalanx III-3, metatarsals IV,
phalanges IV-1, phalanges IV-2, phalanx IV-3, phlanax IV-4, pedal
ungual IV, metatarsal V (Eberth, Xu and Clark, 2010)
(IVPP V15300) (adult) ~17 distal caudal vertebrae (Eberth, Xu and Clark, 2010)
(IVPP V15301) (<1 year old juvenile) skull (50.26 mm), sclerotic
ring, mandibles (46.69 mm), hyoids, several cervical vertebrae (c7 9.58
mm), several dorsal vertebrae, dorsal ribs, sacrum,
~seventh-~twenty-eighth caudal vertebrae, chevron, scapula (36.54 mm),
partial coracoid, humerus (27.80 mm), metacarpal I, phalanx I-1, manual
ungual I, metacarpal II (3.44 mm), phalanx II-1, incomplete phalanx
II-2, metacarpal III (4.19 mm), phalanx III-1, phalanx III-2,
metacarpal IV, incomplete ilium (~46.07 mm), pubis (57.73 mm), partial
ischium, femur, tibiae (86.38 mm), fibula, astragalus, calcaneum,
distal tarsal III, distal tarsal IV, metatarsal III (55.78 mm),
metatarsals, pedal phalanges, pedal unguals (Eberth, Xu and Clark, 2010)
Diagnosis- (after Xu et al., 2009) skull half as long as the femur; premaxilla
toothless; premaxilla with a convex ventral edge; maxilla toothless; nasal with
a lateral shelf dorsal to antorbital fossa; short and wide nasal less than one-third
of skull roof length and only twice as long as wide; ventral process of lacrimal
strongly inclined anteriorly; slender jugal with rod-like suborbital and subtemporal
rami; dentary toothless; large external mandibular fenestra about 40% of mandibular
length; flange on anterior margin of scapular blade; radius tightly adhering
to ulna; radius longer than ulna; olecranon process absent; metacarpal I highly
reduced and carrying no phalanges; phalanx II-1 with distinct lateral process
proximodorsally; metacarpal II much more robust than other metacarpals; metacarpal
III with sub-triangular proximal articular surface; metacarpal III with non-ginglymoidal
distal end; pubis with laterally ridged, prominent posterior boot; pedal digit
I small, only 17% as long as metatarsal III; metatarsus forming a strong transverse
arch; robust ventral process at medial margin of proximal end of metatarsal
III; metatarsal IV nearly straight, appressed against lateral surface of metatarsal
III for nearly its whole length.
Comments-
Clark et al. (2002) first announced a "theropod, represented by the
posterior half of an articulated skeleton, is tentatively identified as
the oldest definitive record of an ornithomimosaur", referring to
either IVPP V15297 or V15299 discovered in August-September 2001.
Their SVP presentation identified three specimens found in 2001- IVPP
V15297, V15299 and V15300. Stiegler (2019) notes it's possible
the skull, cervicals and forelimb elements of IVPP V15297 "belong to
the other Limusaurus skeletons from TBB2001 (IVPP V15298, V15299) or a
separate individual" and that "It is possible that these caudals [IVPP
V15300] belong to V15298, but they are preserved at the bottom and top
of the bone bed [respectively], making this scenario unlikely."
Also that in IVPP V15297 "Some forelimb elements were lost during
collection" and "Two discrete areas with different texture than
adjacent sediment were present distal to the radius in IVPP V15297 and
proximal to metacarpals I and II at the time of this study, but it is
not clear whether these represent fragments of the distal end of the
radius, partially ossified carpals, the former locations of carpal
cartilages (radiale and intermedium?), or whether they are an artifact
of preparation and/or consolidants." The following year IVPP
V15303 and V15304 were found, and in 2005 the remaining specimens were
excavated- IVPP V15923 and V20093-V20105. Xu and Clark (2006)
stated "Four specimens in this collection are here identified as two
new species of ceratosaurian theropod (=neoceratosaurian theropod)"
which "display an unusual combination of character states. While
similar to other ceratosaurs in many derived character states, they
share a number of derived cranial and limb characters with tetanurans
and also a few salient cranial and pelvic features with
coelophysoids." Clark (pers. comm., 6-2009) confirms the
hypothesis of two taxa being present was based on some small specimens
being toothed, which turned out to be ontogenetic instead of taxonomic
with individuals losing their teeth as they grew. Xu and Clark
(2008) also mention this taxon in an abstract as "a new basal
ceratosaur with a strongly reduced digit I and a distally asymmetrical
metacarpal II", contributing to their II-III-IV theory of tetanurine
manual digit homology. Xu et al. (2009) described the taxon as
Limusaurus using IVPP V15923 from 2005 as the holotype, 'IVPP V15924'
from 2002 as the paratype, and mentioned 'IVPP V16134' from 2001 in the
supplementary information. Stiegler notes "IVPP V15924 (Xu et al.
2009) is the same specimen as IVPP V15304 (Eberth, Xing, and Clark
2010), and IVPP V16134 (Xu et al. 2009) is the same as IVPP V15297
(Eberth, Xing, and Clark 2010). IVPP V15304 and IVPP V15297 are the
valid specimen numbers based on the records of the IVPP
collections." Eberth et al. (2010) confirmed five specimens from
2001, two from 2002 and "at least nine Limusaurus inextricabilis" from
2005, although "specimen numbers [were] mostly unavailable at the time
of writing." Note however that IVPP V15301 is the only juvenile
from the first two years and conspicuously not figured in the
publication. Stiegler describes Limusaurus fully in his 2019 thesis.
Limusaurus was found by Xu et al. (2009) to be the sister taxon of Elaphrosaurus
within Ceratosauria, though it also shares some characters with
neoceratosaurs, abelisauroids, noasaurids and tetanurines. Wang
et al.'s (2017) analysis recovered it as a noasaurid, either
elaphrosaurine or noasaurine depending on how OTUs were formed (each
specimen separately, most mature specimen preserved scored for each
character, separate juvenile and adult OTUs, etc.).
Is metacarpal I is tetanurines actually metacarpal II? Xu et al. (2009)
hypothesized Limusaurus may indicate metacarpals I-II-III-IV of tetanurines
are homologous to metacarpals II-III-IV-V in other amniotes, based on several
characters. Digit I in Limusaurus and Aucasaurus are highly reduced,
with no phalanges. Yet Ceratosaurus shows an articular surface for phalanx
I-1, showing the condition in Limusaurus may be derived within ceratosaurs
as opposed to a basal ceratosauroid (ceratosaur+tetanurine) state. Metacarpal
II is medially twisted in Limusaurus, Dilophosaurus and some Coelophysis
bauri specimens, similar to metacarpal I in other saurischians. Unfortunately,
this is not easily determinable from most figures, so the condition in basal
tetanurines is unknown. While Xu et al. note metacarpal III lies ventral to
metacarpal II in tetanurines, as metacarpal IV does to III in Limusaurus
and coelophysoids, this is also true of metacarpal IV in tetanurines (e.g. Guanlong,
as seen in its supplementary information; Tanycolagreus). Similarly,
while metacarpal I does not overlap II in non-tetanurines, this is seemingly
also true in Xuanhanosaurus and Acrocanthosaurus (overlap is present
in Yangchuanosaurus, Megaraptor, Torvosaurus, Allosaurus
and Guanlong however). There is a dorsolateral flange on metacarpal II
of Dilophosaurus and Limusaurus which is similar to one on metacarpal
I of some tetanurines (e.g. Yangchuanosaurus, Allosaurus, Guanlong).
But Xuanhanosaurus and Megaraptor also have such a flange on metacarpal
II, but not I, like basal theropods. Both flanges seem to exist in Acrocanthosaurus,
while none exist in Aucasaurus and Torvosaurus. Xu et al. state
metacarpal II is more robust than I in non-tetanurine theropods, homologizing
it to the robust metacarpal I in tetanurines, but the situation is more complex.
It's clearly the size of the base which is important, since even Coelophysis
and Dilophosaurus have metacarpal I shafts more robust than those of
II. Yet basal tetanurines (e.g. Xuanhanasaurus, Yangchuanosaurus,
Torvosaurus, Megaraptor, Acrocanthosaurus, Allosaurus)
have metacarpal II more robust than I, while Aucasaurus and Herrerasaurus
have the opposite condition. This is true in Xu et al.'s tetanurine example
of Guanlong too, while even their example of Deinonychus has more
proximal area and depth on metacarpal II, just less width. Phalanx I-1 in tetanurines
is said to be longer than phalanx I-1 in Herrerasaurus, Dilophosaurus
and ceratosaurs, but phalanx I-1 is not preserved in any ceratosaur except for
two questionably identified elements in Masiakasaurus. Furthermore, it
is not as if any phalanx on digit II in Dilophosaurus or Herrerasaurus
is notably more elongate than their phalanx I-1, and some basal tetanurines
like Torvosaurus actually have an extremely short phalanx I-1. Metacarpal
II is longest in tetanurines, while III is longest in more basal theropods.
Yet Limusaurus and Ceratosaurus resemble tetanurines in this (contra
Xu et al.'s statements and measurements about the former), and Dilophosaurus
and Coelophysis rhodesiensis are polymorphic (e.g. II longer in the paratype
of Dilophosaurus, III longer in the holotype). There is a proximal dorsolateral
process on metacarpal III in coelophysoids and Limusaurus, similar to
one on metacarpal II in some basal tetanurines like Guanlong. Yet Guanlong
also has a process on metacarpal III, which partly covers metacarpal IV, even
though the latter is not illustrated in Xu et al.'s paper. Acrocanthosaurus
and Allosaurus also have a processes on metacarpal III, while Yangchuanosaurus
lacks processes on metacarpals II or III. This process on metacarpal III of
tetanurines could be homologous to the process on III in basal theropods as
easily as it could the process on II. Finally, Xu et al. state metacarpal III
in tetanurines is short, slender and proximally triangular like metacarpal IV
in basal theropods. Of course, metacarpal IV in tetanurines is also short and
slender (moreso than III), with those of Xuanhanosaurus and Yangchuanosaurus
resembling metacarpal IV in basal theropods more than their metacarpal III.
This is a case where the more reduced metacarpal III in derived tetanurines
like Guanlong and Deinonychus (illustrated by Xu et al.) resembles
basal theropod metacarpal IV more than metacarpal III in basal tetanurines (e.g.
Xuanhanosaurus, Yangchuanosaurus, Torvosaurus, carnosaurs)
do, with the thicker shaft and robust distal articulation in the latter taxa.
As for their triangular proximal outline, metacarpal IV in Dilophosaurus
and Limusaurus are more round than triangular, but Yangchuanosaurus
shows basal tetanurines have triangular metacarpal IV too in any case. Xu et
al. ran a phylogenetic analysis which determined that when characters states
are ordered, the resulting tree assuming tetanurines have digits II-III-IV-V
is six steps longer than if they are assumed to have digits I-II-III-IV. The
length of the unordered trees is equal, but leaving characters unordered potentially
leads to ridiculous "intermediate synapomorphies" like coelophysoids
and Herrerasaurus being diagnosed by having a single phalanx on digit IV (not
more or less) or taxon being diagnosed by having an intermediate ratio, as opposed
to relatives with low and high ratios. Furthermore, neither Xuanhanosaurus,
Yangchuanosaurus dongi or Megaraptor were included in their matrix,
though these taxa show high homoplasy if II-III-IV-V is assumed, as noted above.
I conclude that there is little evidence tetanurine hands lacked digit I.
References- Clark, Xu, Forster, Wang and Andres, 2002. New small dinosaurs
from the Upper Jurassic Shishugou Formation at Wucaiwan, Xinjiang, China. Journal
of Vertebrate Paleontology. 22(3), 44A.
Xu and Clark, 2006. New ceratosaurs from the Jurassic Shishugou Formation of
Western China. Journal of Vertebrate Paleontology. 26(3), 142A.
Xu and Clark, 2008. Homologies in the hand of theropods. Journal of Vertebrate
Paleontology. 28(3), 163A.
Vargas, Wagner and Gauthier, 2009. Limusaurus and bird digit identity. Nature Precedings. DOI: 10.1038/npre.2009.3828.1
Xu, Clark, Mo, Choiniere, Forster, Erickson, Hone, Sullivan, Eberth, Nesbitt,
Zhao, Hernandez, Jia, Han and Guo, 2009. A Jurassic ceratosaur from China helps
clarify avian digital homologies. Nature. 459, 940-944.
Eberth, Xu and Clark, 2010. Dinosaur death pits from the Jurassic of China. Palaios. 25(2), 112-125.
Xu, Clark, Choiniere, Hone and Sullivan, 2011. Reply to "Limusaurus and bird digit identity". Nature Precedings. DOI: 10.1038/npre.2011.6375.1
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cases aid in resolving the relationships of basal Neotheropoda, gauge topological
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Langer, Martins, Manzig, Ferreira, Marsola, Fortes, Lima, Sant'ana,
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Sinocoelurus Young, 1942
S. fragilis Young, 1942
Tithonian?, Late Jurassic
IVPP locality 47, upper Guangyuan Group, Sichuan, China
Syntypes- (IVPP V232-234) incomplete tooth, two partial teeth, tooth
(? x 9 x 4.5; ? x 8.5 x 6; ? x 7.5 x 5, ? x 8 x 5.2 mm)
Referred- ?(IVPP V196) distal ischia (~65 mm deep distally) (Young, 1942)
?(IVPP coll.) proximal tibia (Young, 1942)
Other diagnoses- Young (1942)
diagnosed the taxon with- "Teeth long and slender. Compression and
curvature moderate. Both anterior and posterior sides ridged but no
trace of any serrations, enamel very thin."
Comments-
These isolated teeth were discovered "in three successive days" in late
Spring 1941 and described by Young (1942) as a new taxon of coelurosaur
(sensu Huene) based on "the general features of the teeth". Though none
are complete, the crowns would have been ~40 mm long. Dong et al.
(1983) incorrectly included IVPP V235 as a syntype, but this is a Szechuanosaurus
syntype instead. The identity of the incomplete tooth and two partial
teeth which are illustrated, which number includes two teeth, and which
specimen each set of FABL and BW measurements is from are unknown,
although Dong et al. give V232 as the number of the incomplete tooth.
The teeth are highly elongate, strongly compressed (BW/FABL = 50-70%)
and slightly curved lingually and distally. There are both mesial and
distal carinae, with the distal carina being stronger. The carinae lack
serrations, which Young used to justify creation of a new genus of
uncertain relationship to other coelurosaurs. Wu et al. (2009) further
described them as striated. Within Coelurosauria, Steel (1970) referred
Sinocoelurus to Coeluridae, which was followed by most
traditional authors. This was due to Coeluridae being a wastebasket taxon for
small theropods, in addition to Sinocoelurus' name no doubt. More recently,
Sinocoelurus has usually been referred to as Theropoda indet. (e.g. Norman,
1990), though Holtz et al. (2004) placed it in Tetanurae indet. without comment.
Young (1942) placed locality 47 at "the top part of the Kuangyuan Series and
immediately below the Chentsianyen conglomerate", now known as the
Guangyuan Group and the Chengqiangyan Group, with the former
corresponding to the Xiashaximiao Formation through the Penglaizhen
Formation. As it was found "immediately below" the boundary, Sinocoelurus
may be from the Penglaizhen Formation or slightly lower
Shuining Formation. Young et al. (1943) list the genus as being
found in layers 9 and 8b. The age is listed as Tithonian on
fossilworks and in Weishampel (1990), the latter cited as from "Dong
(pers. comm.)". Geographically, locality 47 is "the hill slopes under the so-called
Chentsianyen escarpment S. of the Kuangyuan city", now known as
Guangyuan. Dong et al. (1983) incorrectly said "Young's
original description stated that because Sinocoelurus
was derived from the ostracod-rich beds at the top of the Guangyuan
System, the specimens should be recognized as being derived from the
Qianfoya Fm." while Young never mentions the latter formation (more
recently called the Qianfuyan Formation and placed near the EJ-MJ
boundary).
Wu et al. (2009) examined the material and considered it to "most probably belong to
a plesiosaur." However, plesiosaur teeth are circular in section (except for
the tip in some taxa), including the contemporaneous Yuzhoupliosaurus,
and their carinae are perpendicular to the plane of curvature. Rozhdestvensky (1977) said that Sinocoelurus "proved to be a crocodile" without explanation.
A placement in Crocodyliformes seems possible, as taxa such as Kaprosuchus,
Shantungosuchus, Torvoneustes and Pehuenchesaurus have
compressed, recurved teeth without serrations. The recent discovery of toothed juveniles of Limusaurus
provides a plausible identification, as these are recurved, compressed
and serrationless, of similar size and close geographically and
stratigraphically. Confirmation awaits detailed description
of Limusaurus teeth, but even if true the lack of other preserved elaphrosaurine teeth means Sinocoelurus is not likely to be diagnostic enough to be a senior synonym of Limusaurus.
Referred material- Young (1942) also briefly describes a proximal tibia as cf. Sinocoelurus fragilis (stated to be "possible that it belongs to Sinocoelurus fragilis") without providing its catalogue number. As in Limusaurus, Berthasaura, Gualicho and tetanurines the fibular crest is separated from the proximal end, but unlike Limusaurus (although like Elaphrosaurus) the medial condyle reaches furthest posteriorly. Similarly, a pair of distal ischia (IVPP V196) are listed as cf. Sinocoelurus fragilis ("Its belonging to Sinocoelurus is equally probable"), but the expanded distal ends are about as similar to Limusaurus as they are to Yangchuanosaurus in the illustrated lateral views, making these Averostra indet. pending further description.
References- Young, 1942. Fossil vertebrates from Kuangyuan, N. Szechuan,
China. Bulletin of the Geological Society of China. 22(34), 293-309.
Young, Bien and Mi, 1943. Some geologic problems of the Tsinling. Bulletin
of the Geological Society of China. 23(1-2), 15-34.
Steel, 1970. Part 14. Saurischia. Handbuch der Paläoherpetologie/Encyclopedia
of Paleoherpetology. Gustav Fischer Verlag, Stuttgart. 87 pp.
Rozhdestvensky, 1977. The study of dinosaurs in Asia. Journal of the Palaeontological
Society of India. 20, 102-119.
Dong, Zhou and Zhang, 1983. Dinosaurs from the Jurassic of Sichuan. Palaeontologica
Sinica, New Series C. 162(23), 1-136.
Norman, 1990. Problematic Theropoda: "Coelurosaurs". In Weishampel,
Dodson and Osmolska (eds). The Dinosauria. University of California Press.
280-305.
Weishampel, 1990. Dinosaurian distribution. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria. University of California Press. 63-139.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel,
Dodson and Osmolska (eds.). The Dinosauria (second edition).
University of California Press. 71-110.
Wu, Currie, Dong, Pan and Wang, 2009. A new theropod dinosaur from the Middle
Jurassic of Lufeng, Yunnan, China. Acta Geologica Sinica. 83(1), 9-24.
undescribed elaphrosaurine (Allain, Vullo, Leprince, Neraudeau and
Tournepiche, 2011)
Hauterivian-Barremian, Early Cretaceous
Angeac-Charente lignite, France
Material- (ANG 10-07) partial metatarsal IV (Allain et al., 2014)
(ANG 10-24) tibia (320 mm) (Allain et al., 2014)
(ANG 10-158) tibia (385 mm) (Allain et al., 2014)
(ANG 10-175) posterior cervical vertebra (66 mm) (Neraudeau et al., 2012)
(ANG 10-237) partial metatarsal III (Allain et al., 2014)
(ANG 10-240) partial metatarsal II (Allain et al., 2014)
(ANG 11-1000) tibia (402 mm) (Allain et al., 2014)
(ANG 11-1357) tibia (349 mm) (Allain et al., 2014)
(ANG 12-1808) tibia (279 mm) (Allain et al., 2014)
(ANG 12-1893) tibia (320 mm) (Allain et al., 2014)
(ANG coll.) (at least 20 individuals, juvenile, subadult and adult)
1370 elements including postorbital, dentary, twenty-one cranial and
mandibular elements, fifteen cervical vertebrae, five cervical ribs,
thirty-seven dorsal vertebrae, twenty partial dorsal ribs, twenty
sacral vertebrae, thirty-six caudal vertebrae, fourteen chevrons, four
proximal scapulae, two coracoids, furcuila, humerus, radius, ulna,
fifteen partial pelvic elements including ilium, pubis and ischium,
femora, forty-eight partial tibiae, eight metatarsals, eleven pedal
phalanges and eleven pedal unguals (Allain et al., 2011)
Comments- Though mentioned by Allain et al. (2011) and Neraudeau et al.
(2012), this was basically undescribed until Allain et al. (2014) noted several
features and viewed the taxon as a Rosetta Stone showing Thecocoelurus,
Wessex femur MIWG 6214, Wessex tibiae NHMUK R186 and UOP-C002-2004, and Valdoraptor
were all related ornithomimosaurs. Of these, Thecocoelurus is more similar
to Falcarius
so is listed as a therizinosaur here, although the other specimens are
provisionally retained close to the Angeac taxon pending study.
Unpublished photos of preserved elements suggest this is not an
ornithomimosaur, but rather a ceratosaur similar to Elaphrosaurus and Limusaurus.
Supporting evidence includes the posterior cervical pleurocoel, short
distal caudals, unexpanded chevrons, broad scapula with prominent
dorsal flange, radius and ulna less than half humeral length, manus
reduced so that to date only a single element has been found, barely
ventrally extended preacetabular process, distal obturator notch, and
reduced side metatarsals. While it was recovered as an
ornithomimosaur in Hartman et al. (2019) using the published fragments,
adding unpublished data moves it into Ceratosauria by Deltadromeus, Afromimus and Limusaurus.
References- Allain, Vullo, Leprince, Neraudeau and Tournepiche, 2011.
An ornithomimosaur-dominated bonebed from the Early Cretaceous of Southwestern
France. Journal of Vertebrate Paleontology. Program and Abstracts 2011, 61.
Néraudeau, Allain, Ballèvre, Batten, Buffetaut, Colin, Dabard,
Daviero-Gomez, Albani, Gomez, Grosheny, Le Loeuff, Leprince, Martín-Closas,
Masure, Mazin, Philippe, Pouech, Tong, Tournepiche and Vullo, 2012. The Hauterivian-Barremian
lignitic bone bed of Angeac (Charente, south-west France): Stratigraphical,
palaeobiological and palaeogeographical implications. Cretaceous Research. 37,
1-14.
Allain, Vullo, Le Loeuff and Tournepiche, 2014. European ornithomimosaurs (Dinosauria,
Theropoda): An undetected record. Geologica Acta. 12(2), 127-135.
Rozada, Allain, Vullo, Leprince and Tournepiche, 2014. Taphonomy of the ornithomimosaur
dinosaur herd from the Early Cretaceous lignitic bone bed of Angeac-Charente
(France). Journal of Vertebrate Paleontology. Program and Abstracts 2014, 217-218.
Valdoraptor Olshevsky, 1991
V. oweni (Lydekker, 1889) Olshevsky, 1991
= Megalosaurus oweni Lydekker, 1889
= Altispinax oweni (Lydekker, 1889) Huene, 1923
Valanginian, Early Cretaceous
Grinstead Clay Formation, England
Holotype- (NHMUK 2559; intended paratype of Altispinax "lydekkerhueneorum")
incomplete metatarsal II (~195 mm), incomplete metatarsal III (~240 mm), incomplete
metatarsal IV
Referred- (NHMUK 2661) metatarsal II (170 mm) (Lydekker, 1888)
Diagnosis- (after Naish and Martill, 2007) metatarsal II with prominent
dorsolateral ridge.
(after Naish, 2011) metatarsal II ~81% of metatarsal III length; suboval interosseous
space that separates distal ends of metatarsals II and III.
Other diagnoses- Olshevsky (1991) originally diagnosed Valdoraptor
by its more slender metatarsus than Megalosaurus with "larger and
more robust metatarsal III", which is more accurately the m,ore slender
metatarsals II and IV.
Naish and Martill (2007) included the mediolaterally compressed metatarsal II
in their diagnosis, but this is true of all ornithomimosaurs.
Comments- This was originally referred to Hylaeosaurus by Owen
(1856), but was identified as theropod by Hulke (1881). Lydekker (1888) referred
both NHMUK 2559 and 2661 to Megalosaurus dunkeri. After it was named as
a new Megalosaurus species by Lydekker (1889), Olshevsky (1991) proposed
the genus Valdoraptor as a possible allosaurid. Holtz et al. (2004) suggested
it might prove synonymous with Neovenator or Eotyrannus, but Naish
and Martill (2007) confirm it is distinct. Carrano et al. (2012) refer it to
Avetheropoda based on the trapezoidal cross section of metatarsal III. Most
recently, Allain et al. (2014) proposed it was almost identical to their undescribed
supposed ornithomimosaur from the Angeac-Charente lignite of France but also somehow
a nomen dubium. If Allain et al. were correct that the genus was almost identical
to and shared a synapomorphy with their specimens, it would make their unnamed
taxon Valdoraptor, it wouldn't make Valdoraptor indeterminate.
However, they fail to mention that the Angeac taxon lacks the dorsolateral ridge
on metatarsal II Naish and Martill used to diagnose Valdoraptor, or the
suboval interosseous space between metatarsals II and III that Naish used to
diagnose the genus in 2011. They also misunderstand Naish's short metatarsal
II length as only meaning it was shorter than III (which is near universal in
theropods), whereas he was referring to the extreme shortness (81%) that is
shorter than other ornithomimosaurs. The length of metatarsal II in the Angeac
taxon is unpublished still, as only partial metatarsals have been figured. In
any case, the ridge and interosseous space validate Valdoraptor
and distinguish it from the Angeac taxon, though they do appear to be
related. Unpublished photos strongly support a ceratosaurian
identity for the Angeac material, so Valdoraptor may belong in that clade as well.
Lydekker (1890) referred NHMUK R604a, 604c-d and 1525 from the earlier Wadhurst
Clay and NHMUK 2574, 2661 and 2680 from the type horizon to the species, but
most have not been compared in detail (see Altispinax? sp. indet. entry).
Pickering (1995) intended to use the oweni holotype and the 604 specimens
as paratypes of his proposed species Altispinax "lydekkerhueneorum",
but this is a nomen nudum and objective junior synonym of Altispinax dunkeri
(see entry). Naish (2007; 2011) did note that NHMUK 2661 was properly referred
to oweni however.
References- Owen, 1856. Monograph on the fossil Reptilia of the Wealden
Formation. Part IV. Palaeontographical Society Monographs. 10, 1-26.
Hulke, 1881. Polacanthus foxii, a large undescribed dinosaur from the
Wealden Formation in the Isle of Wight. Philosophical Transactions of the Royal
Society of London. 172, 653-662.
Lydekker, 1888. Catalogue of the Fossil Reptilia and Amphibia in the British
Museum (Natural History), Cromwell Road, S.W., Part 1. Containing the Orders
Ornithosauria, Crocodilia, Dinosauria, Squamata, Rhynchocephalia, and Proterosauria.
British Museum of Natural History, London. 309 pp.
Lydekker, 1889. On the remains and affinities of five genera of Mesozoic reptiles.
Quarterly Journal of the Geological Society of London. 45, 41-59.
Lydekker, 1890. Contributions to our knowledge of the dinosaurs of the Wealden
and the sauropterygians of the Purbeck and Oxford Clay. Quarterly Journal of
the Geological Society of London. 46, 36-53.
Huene, 1923. Carnivorous Saurischia in Europe since the Triassic. Bulletin of
the Geological Society of America. 34, 449-458.
Huene, 1926. The carnivorous Saurischia in the Jura and Cretaceous formations,
principally in Europe. Revista del Museo de La Plata. 29, 1-167.
Olshevsky, 1991. A revision of the parainfraclass Archosauria Cope, 1869, excluding
the advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.
Olshevsky, unpublished. Valdoraptor oweni (Theropoda: Allosauridae) from
the Upper Wealden of West Sussex, England. Mesozoic Vertebrate Life #2.
Pickering, 1995. Jurassic Park: Unauthorized Jewish Fractals in Philopatry,
A Fractal Scaling in Dinosaurology Project, 2nd revised printing. Capitola,
California. 478 pp.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria Second Edition. University of California Press. 71-110.
Naish, 2007. The large theropods Becklespinax and Valdoraptor
from the Lower Cretaceous of England. SVPCA 2007, Abstracts Programme. 24-25.
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.
Naish, 2011. Theropod dinosaurs. In Batten (ed.). English Wealden Fossils. The
Palaeontological Association. 526-559.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Allain, Vullo, Le Loeuff and Tournepiche, 2014. European ornithomimosaurs (Dinosauria,
Theropoda): An undetected record. Geologica Acta. 12(2), 127-135.
unnamed possible Elaphrosaurinae (Lydekker, 1888)
Barremian, Early Cretaceous
Wessex Formation, England
Material- (NHMUK R186) tibia (158.0 mm) (Lydekker, 1888)
(Dinosaur Expeditions Centre coll.; cast UOP-C002-2004) dorsal vertebra, partial
tibia (~214 mm), proximal metatarsal (Naish, 2011)
(Dinosaur Expeditions Centre coll.) partial sacrum, ilial fragment, distal
tibia (~260 mm), fibular fragment, proximal metatarsals, four pedal phalanges
(one partial) (Mattsson, pers. comm. 2015)
(MIWG 6214) (juvenile) incomplete femur (123 mm) (Naish, 1998; described in
Naish, 2000)
Comments- NHMUK R186 was originally listed as belonging to Hypsilophodon
foxii by Lydekker (1888), then referred to Calamosaurus foxi by him
in 1891. Since then it has been regarded as a basal coelurosaur, close to Coelurus
or Compsognathus.
Note that Galton (1973) was incorrect in claiming the astragalus was
attached. Naish (1999) redescribed it in his thesis and proposed in
Naish et al. (2001) that earlier authors had gotten its orientation
backwards. This was shown to be incorrect by Galton and Molnar (2005).
More recently, Allain et al. (2014) argue that its morphology is very
similar to their undescribed supposed ornithomimosaur (probably a
ceratosaur) from the Angeac-Charente lignite of France. Longrich et al.
(2021) suggested it resembled dromaeosaurids "in having a triangular
fibular crest, a broad distal end of the tibia an astragalar suture
that is straight in posterior view, and a relatively low ascending
process of the astragalus" and may belong to Ornithodesmus.
Naish (2000) described a femur he thought was most similar to Microvenator,
but referred to Theropoda indet.. Allain et al. (2014) tentatively referred
it to Ornithomimosauria based on preliminary study of the Angeac taxon.
A specimen referred to Calamosaurus was discovered in 2002 which is displayed
at the Dinosaur Expeditions Centre (Mattsson, pers. comm. 2015), though it remains
undescribed. Naish (2011) did figure the partial tibia though and states it
is "morphologically identical" to NHMUK R186. While it cannot be compared
to Calamosaurus, the dorsal vertebra should make comparison to Aristosuchus
and the Angeac taxon possible. A second undescribed specimen was discovered in 2014 and is also on
display at the Dinosaur Expeditions Centre (Mattsson, pers. comm. 2015), similarly
referred to Calamosaurus although it cannot be compared to that genus
either. In this case, the sacrum should allow comparison with Aristosuchus and the Angeac taxon.
Mattsson reports its tibia "appears to be a perfect match of the 2002 specimen"
and that Siebert is researching these materials.
References- Lydekker, 1888. Catalogue of the fossil Reptilia and Amphibia
in the British Museum (Natural History). Part I. Containing the orders Ornithosauria,
Crocodilia, Dinosauria, Squamata, Rhynchocephalia, and Proterosauria. British
Museum (Natural History), London. 309 pp.
Lydekker, 1891. On certain ornithosaurian and dinosaurian remains. Quarterly
Journal of the Geological Society of London. 47, 41-44.
Galton, 1973. A femur of a small theropod dinosaur from the Lower Cretaceous
of England. Journal of Paleontology. 47, 996-1001.
Naish, 1998. Predatory dinosaurs in England's Lower Cretaceous: Crocodile heads,
puffin snouts and the odd Lazarus taxon. Newsletter of the European Palaeontological
Association. 13, 15.
Naish, 1999. Studies on Wealden Group theropods - an investigation into the
historical taxonomy and phylogenetic affinities of new and previously neglected
specimens. Masters thesis. University of Portsmouth. 184 pp.
Naish, 2000. A small, unusual theropod (Dinosauria) femur from the Wealden Group
(Lower Cretaceous) of the Isle of Wight, England. Neues Jahrbuch für Geologie
und Paläontologie Monatshefte. 2000, 217-234.
Naish, Hutt and Martill, 2001. Saurichian dinosaurs 2: Theropods. in Martill
and Naish (eds). Dinosaurs of the Isle of Wight. The Palaeontological Association.
242-309.
Galton, and Molnar, 2005. Tibiae of small theropod dinosaurs from southern England:
From the Middle Jurassic of Stonesfield near Oxford and the Lower Cretaceous
of the Isle of Wight. In Carpenter (ed.). The Carnivorous Dinosaurs. Indiana
University Press. 3-22.
Naish, 2011. Theropod dinosaurs. In Batten (ed.). English Wealden Fossils. The
Palaeontological Association. 526-559.
Allain, Vullo, Le Loeuff and Tournepiche, 2014. European ornithomimosaurs (Dinosauria,
Theropoda): An undetected record. Geologica Acta. 12(2), 127-135.
Longrich, Martill and Jacobs, 2021. A new dromaeosaurid dinosaur from
the Wessex Formation (Lower Cretaceous, Barremian) of the Isle of
Wight, and implications for European palaeobiogeography. Cretaceous
Research. Journal Pre-proof. DOI: 10.1016/j.cretres.2021.105123
Abelisauridae sensu Rowe et al., 1997
Definition- (Carnotaurus sastrei <- Elaphrosaurus bambergi)
(modified)
= Noasaurinae sensu Delcourt 2018
Definition- (Noasaurus leali <- Elaphrosaurus bambergi)
(modified)
References-
Delcourt, 2018. Ceratosaur palaeobiology: New insights on evolution and
ecology of the southern rulers. Scientific Reports. 8:9730.
Neoceratosauria Novas, 1991
Definition- (Ceratosaurus nasicornis + Carnotaurus sastrei)
(Hendrickx, Hartman and Mateus, 2015; modified from Holtz, 1994)
Other definitions- (Ceratosaurus nasicornis <- Coelophysis
bauri) (modified from Padian et al., 1999)
(Ceratosaurus nasicornis + Abelisaurus comahuensis) (Dal Sasso, Maganuco and Cau, 2018)
= "Neoceratosauria" Novas, 1989
= Ceratosauroidea Marsh, 1884 vide Bonaparte, Novas and Coria, 1990
Other definitions- (Carnotaurus sastrei + Noasaurus leali) (Delcourt, 2018)
= Neoceratosauria sensu Dal Sasso, Maganuco and Cau, 2018
Definition- (Ceratosaurus nasicornis + Abelisaurus comahuensis)
*= Etrigansauria Delcourt, 2018
Definition- (Ceratosaurus nasicornis, Carnotaurus sastrei <- Noasaurus leali) (Delcourt, 2018)
Comments- This clade was first proposed by Novas (1989) in his thesis,
and later published by him in 1991. It included ceratosaurids, noasaurids and
abelisaurids in both, distinguishing them from coelophysoid ceratosaurs. It
has been used since in a similar manner, though lately it also serves to separate
the former taxa from basal ceratosaurs such as Elaphrosaurus.
Bonaparte et al. (1990) named Ceratosauroidea for the same clade,
though distinguished from allosauroid carnosaurs this time.
Ceratosauroidea has been used much less often and has usually been
given definitions congruent with ceratosaurian coelophysoids, so
Neoceratosauria is preferred here. An exception is Delcourt
(2018), who redefined Ceratosauroidea via a series of errors. He
first used Wilson et al.'s (2003) poor definition for Abelisauroidea (Carnotaurus sastrei + Noasaurus leali),
and noted that in Wang et al.'s (2017) topology this includes
Ceratosaurus. Because the family-level name Ceratosauroidea has
priority over Abelisauroidea according to the ICZN, Delcourt decided to
give Ceratosauroidea Wilson et al.'s definition for
Abelisauroidea. The first error was using Wilson et al.'s
definition for Abelisauroidea, which obviously does not function in
Wang et al.'s topology as Ceratosaurus
can never be an abelisauroid under the ICZN's rules or in any concept
of Abelisauroidea used by any other author. The second error was
believing superfamily phylogenetic definitions should be transferrable,
which is an area of overlap between the ICZN and Phylocode that has
never been instituted. Finally, (Carnotaurus sastrei + Noasaurus leali) cannot be a definition for a Ceratosaur- clade as it does not contain Ceratosaurus nasicornis in its definition (Phylocode Article 11.7).
Etrigansauria was named by Delcourt (2018) for a clade including
ceratosaurids and abelisaurids but not noasaurids, which self destructs
in this website's phylogeny. If noasaurids are outside a
ceratosaurid plus abelisaurid clade, Neoceratosauria would have
priority for the latter clade. Technically they would not be
synonyms as Etriganosauria is defined as stem-based to include taxa
closer to ceratosaurids and abelisaurids, while Neoceratosauria is
node-based to include only descendents of the most recent common
ancestor of ceratosaurids and abelisaurids. Yet Delcourt (2018:
Fig. 1) illustrates Etrigansauria as node-based, excluding Berberosaurus
despite the stem-based definition. The need for such a
distinction is also questionable, as one of the few analyses that
allows Etrigansauria to exist, Wang et al. (2017), places no taxon
definitively as a non-neoceratosaur etrigansaurian, with their Analysis
2 placing Berberosaurus there and Analysis 4 placing Eoabelisaurus
there. The term Etrigansauria is best left unused, with
neoceratosaurs either including or excluding noasaurids based on the
analysis. At one point Delcourt states "in some analyses,
Berberosaurus is considered as a ... sister-taxa of cornisauria" with a
reference to Wang et al.'s topology, suggesting Cornisauria was an
early name for Etrigansauria.
A supposed neoceratosaur distal femur from the Sinpetru Beds of Romania described
by Csiki and Grigorescu (1998) is a probably hadrosaurid distal metatarsal based
on a matching complete specimen (MAFI Ob.3120a) (Kessler et al., 2005).
References- Novas, 1989. Los dinosaurios carnivoros de la Argentina.
PhD thesis. Universidad Nacional de La Plata. 510 pp.
Novas, 1991. Phylogenetic relationships of ceratosaurian theropod dinosaurs.
Ameghiniana. 28, 401.
Csiki and Grigorescu, 1998. Small theropods from the Late Cretaceous of the
Hateg Basin (Western Romania) - an unexpected diversity at the top of the food
chain. Oryctos. 1, 87-104.
Wilson, Sereno, Srivastava, Bhatt, Khosla and Sahni, 2003. A new abelisaurid
(Dinosauria, Theropoda) from the Lameta Formation (Cretaceous, Maastrichtian)
of India. Contributions from the Museum of Paleontology. 31(1), 1-42.
Kessler, Grigorescu and Csiki, 2005. Elopteryx revisited - a new bird-like
specimen from the Maastrichtian of the Hateg Basin. Acta Palaeontologica Romaniae.
5, 249-258.
Hendrickx, Hartman and Mateus, 2015. An overview of non-avian theropod discoveries
and classification. PalArch's Journal of Vertebrate Palaeontology. 12(1), 1-73.
Wang, Stiegler, Amiot, Wang, Du, Clark and Xu, 2017 (online 2016). Extreme ontogenetic
changes in a ceratosaurian theropod. Current Biology. 27(1), 144-148.
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.
Delcourt, 2018. Ceratosaur palaeobiology: New insights on evolution and
ecology of the southern rulers. Scientific Reports. 8:9730.
undescribed neoceratosaur (Company, Torices, Pereda-Suberbiola and Ruiz-Omenaca,
2009)
Late Campanian-Early Maastrichtian, Late Cretaceous
Sierra Perenchiza Formation, Valencia, Spain
Material- teeth (to 50 mm)
Comments- Described as moderately labiolingually compressed and distally
recurved; both mesial and distal serrations; mesial carina laterally displaced;
serrations chisel-shaped, wider labiolingually than mesiodistally; mesial serrations
slightly smaller than distal ones (~3 denticles per mm); enamel slightly stretch-marked.
Reference- Company, Torices, Pereda-Suberbiola and Ruiz-Omenaca, 2009.
Theropod teeth from the Late Cretaceous of Chera (Valencia, Eastern Spain).
Journal of Vertebrate Paleontology. 29(3), 81A.
unnamed possible neoceratosaur (Young, 1958)
Early Cretaceous
Tsanmakou, Shanxi, China
Material- (IVPP V969) caudal vertebra, distal scapula, distal ischia,
proximal tibia, proximal fibula, metatarsal II (~300 mm), pedal phalanx III-1?
(95 mm)
Comments- This was referred to Allosauridae indet. by Young (1958). Carrano
et al. (2012) noted the oblique proximal margin to the proximomedial fibular
fossa is unlike carnosaurs, and that the fossa's depth is unlike megalosauroids.
They referred it to Averostra indet., but noted the large
m. iliofibularis tubercle was like neoceratosaurs.
Reference- Young, 1958. The first record of dinosaurian remains from
Shansi. Vertebrata PalAsiatica. 2(4), 231-236.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
unnamed probable neoceratosaur (Maganuco et al., 2005)
Bathonian, Middle Jurassic
Isalo Formation IIIb, Madagascar
Material- (MSNM V5778) anterior tooth (25.2 mm)
(MSNM V5779) lateral tooth (24.4 mm)
(MSNM V5780) lateral tooth (10.2 mm)
(MSNM V5781) lateral tooth (9.1 mm)
(MSNM V5782) lateral tooth (14.9 mm)
(MSNM V5783) lateral tooth (12.3 mm)
(MSNM V5784) lateral tooth (14.1 mm)
(MSNM V5788) lateral tooth (11.2 mm)
(MSNM V5790) lateral tooth (8.5 mm)
(MSNM V5794) lateral tooth (10.2 mm)
(MSNM V5798) lateral tooth (10.4 mm)
(MSNM V5799) lateral tooth (11.8 mm)
(MSNM V5806) lateral tooth (>9.7 mm)
(MSNM V5807) anterior tooth (28 mm)
(MSNM V5809) lateral tooth (>17.3 mm)
(MSNM V5810) lateral tooth (21.6 mm)
(MSNM V5814) lateral tooth (16.1 mm)
(MSNM V5817) lateral tooth (13.7 mm)
(MSNM V5818) lateral tooth (>14.6 mm)
(MSNM V5820) anterior tooth (16.4 mm)
(MSNM V5821) lateral tooth (9.7 mm)
(MSNM V5957) lateral tooth (>14 mm)
(MSNM V5962) lateral tooth (>19.1 mm)
Comments- These teeth resemble ceratosaurids in being extremely labiolingually
compressed, and having the mesial carina vary in basal extent. The premaxillary
teeth lack lingual fluting, unlike ceratosaurids. They are similar to abelisaurids'
in having well developed basally angled blood grooves, short crowns, a fairly
high DSDI (mean of 1.24). In addition to possessing the ceratosaurid similarities
noted above, they differ from abelisaurids in having more serrations and lacking
a drop-shaped cross section.
Reference- Maganuco, Cau, and Pasini, 2005. First description of theropod
remains from the Middle Jurassic (Bathonian) of Madagascar. Atti della Società
Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano.
146(2), 165-202.
undescribed possible Neoceratosauria (Bonaparte and Powell, 1980)
Late Campanian-Maastrichtian, Late Cretaceous
El Brete, Lecho Formation, Salta, Argentina
Material- (PVL 4062) few teeth (15-33 mm)
Comments-
Bonaparte and Powell (1980) mentioned "A few isolated carnivorous teeth
were discovered whiIe excavating the Titanosaurid remains", there
described as the type material of Saltasaurus loricatus.
They stated "all of them are of the same type, laterally compressed,
with anterior and posterior borders serrated and asymmetrical, and with
the end of the tooth partially projected towards the rear." While they
described the material as "Carnosauria Genus and species indet.", they
also admitted "we have no evidence to exclude them from the carnivorous
Coelurosauria." Thus saying "they are of the type present in Genyodectes ... or in Majungasaurus" does not necessarily indicate a ceratosaurian identity within Averostra.
Reference- Bonaparte and Powell, 1980. A continental assemblage of tetrapods
from the upper Cretaceous beds of El Brete, northwestern Argentina (Sauropoda-Coelurosauria-Carnosauria-Aves).
Mémoires de la Société Géologique de France. Nouvelle
Série 59, 19-28.
Ceratosauridae Marsh, 1884
Definition- (Ceratosaurus nasicornis <- Abelisaurus comahuensis)
(modified from Rauhut, 2004)
Other definitions- (Ceratosaurus nasicornis <- Noasaurus
leali, Carnotaurus sastrei) (Hendrickx, Hartman and Mateus, 2015)
(Ceratosaurus nasicornis <- Noasaurus leali, Abelisaurus comahuensis) (Dal Sasso, Maganuco and Cau, 2018)
(Ceratosaurus nasicornis <- Carnotaurus sastrei) (Delcourt, 2018)
= Neoceratosauroidea Marsh, 1884 sensu Madsen and Welles, 2000
= Ceratosauridae sensu Hendrickx, Hartman and Mateus, 2015
Definition- (Ceratosaurus nasicornis <- Noasaurus leali, Carnotaurus
sastrei)
= Ceratosauridae sensu Dal Sasso, Maganuco and Cau, 2018
Definition- (Ceratosaurus nasicornis <- Noasaurus leali, Abelisaurus comahuensis)
= Ceratosauridae sensu Delcourt, 2018
Definition- (Ceratosaurus nasicornis <- Carnotaurus sastrei)
Diagnosis- (after Rauhut, 2000) extremely labiolingually compressed maxillary
teeth; longest maxillary tooth exceeds minimum dentary height.
(after Chure, 2000) lingually fluted premaxillary teeth
Comments- Marsh (1884) named this family to include only Ceratosaurus,
and it has been used infrequently since to include taxa like Proceratosaurus,
Sarcosaurus and Genyodectes.
Madsen and Welles (2000) listed Neoceratosauroidea "including only the
family Ceratosauridae", but this in invalid as there is no genus Neoceratosaurus.
References- Marsh, 1884. Principal characters of American Jurassic dinosaurs.
Part VIII. The order Theropoda. The American Journal of Science. 27, 329-340.
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.
Rauhut, 2004. Provenance and anatomy of Genyodectes serus, a large-toothed
ceratosaur (Dinosauria: Theropoda) from Patagonia. Journal of Vertebrate Paleontology.
24(4), 894-902.
Hendrickx, Hartman and Mateus, 2015. An overview of non-avian theropod discoveries
and classification. PalArch's Journal of Vertebrate Palaeontology. 12(1), 1-73.
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.
Delcourt, 2018. Ceratosaur palaeobiology: New insights on evolution and
ecology of the southern rulers. Scientific Reports. 8:9730.
Ceratosaurus? roechlingi
Janensch, 1925
Late Tithonian, Late Jurassic
Upper Dinosaur Member of the Tendaguru Formation, Tanzania
Lectotype- (MB R 2162; = MW 4) partial mid caudal centrum
Diagnosis- (after Rauhut, 2011) Provisionally indeterminate relative
to Ceratosaurus nasicornis.
Comments- Janensch (1925) based this species on a partial quadrate (MB
R 2160), three partial caudal vertebrae (MB R 1934, 1935, 2162) and a proximal
fibula (MB R 3627), thought to belong to a single individual, as well as two
caudals from earlier sediments (MB R 1938, 2166). Additionally, he labeled an
astragalus and calcaneum (MB R 1926) as belonging to this taxon in the museum
collections, though the specimen is not mentioned in the literature until Carrano
and Sampson (2008). Madsen and Welles (2000) attempted to make the distal quadrate
MB R 2160 the holotype of Ceratosaurus roechlingi,
but as pointed out by Rauhut (2011) this is not valid as they only used
the word 'type' (ICZN 74.7.1). Rauhut instead made the partial caudal
MB R 2162 the lectotype because it shows ceratosaurian characters
unlike the other material. He also found that while the other two
associated vertebrae are of the right size to belong to the lectotype
(though they are only referrable to Averostra indet.), the quadrate
(also Averostra indet.) and hindlimb elements (Megalosauroidea indet.)
are from a larger individual. The earlier caudals were made the type of
his new carcharodontosaurid Veterupristisaurus, described as "a very large probable allosauroid"
in his 2006 abstract. Rauhut found roechlingi to be indeterminate, but
a ceratosaur based on the central cavity and possibly ceratosaurid based on
the wide and deep ventral median groove (the latter also present in some basal
tetanurines).
References- Janensch, 1925. Die Coelurosaurier und Theropoden der Tendaguru-Schichten
Deutsch-Ostafrikas. Palaeontographica. 1(supp. 7), 1-99.
Madsen and Welles, 2000. Ceratosaurus (Dinosauria, Theropoda) a revised
osteology. Miscellaneous Publication 00-2 Utah Geological Survey. 80 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.
Carrano and Sampson, 2008 (online 2007). The Phylogeny of Ceratosauria (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 6, 183-236.
Rauhut, 2011. Theropod dinosaurs from the Late Jurassic of Tendaguru (Tanzania).
Palaeontology. 86, 195-239.
Genyodectes Woodward, 1901
G. serus Woodward, 1901
Aptian-Albian, Early Cretaceous
Cerro Castano Member(?) of Cerro Barcino Formation, Chubut, Argentina
Holotype- (MLP 26-39) premaxillae, anteroventral maxillae, anterior dentaries,
partial supradentaries, splenial fragment, teeth
Diagnosis- (after Rauhut, 2004) ceratosaurid synapomorphies (extremely
labiolingually compressed maxillary teeth; longest maxillary tooth exceeds minimum
dentary height) combined with plesiomorphic presence of four premaxillary teeth.
References- Woodward, 1901. On some extinct reptiles from Patagonia,
of the genera Miolania, Dinilysia, and Genyodectes. Proceedings
of the Zoological Society of London. 1901, 169-184.
Huene, 1929. Los saurisquios y ornitisquios del Cretacéo Argentino. Anales
del Museo de La Plata (series 3). 3, 1-196.
Rauhut, 2004. Provenance and anatomy of Genyodectes serus, a large-toothed
ceratosaur (Dinosauria: Theropoda) from Patagonia. Journal of Vertebrate Paleontology.
24(4), 894-902.
Ostafrikasaurus Buffetaut, 2011
O. crassiserratus
Buffetaut, 2011
Tithonian, Late Jurassic
Upper Dinosaur Member of the Tendaguru Formation, Tanzania
Holotype- (MB R 1084; type a of Janensch; paralectotype of Labrosaurus
stechowi) premaxillary tooth (46 mm)
Late Kimmeridgian, Late Jurassic
Middle Dinosaur Member of the Tendaguru Formation, Tanzania
Referred- (MB R 1091; type d of Janensch; paralectotype of Labrosaurus
stechowi) premaxillary tooth
Diagnosis- (after Rauhut, 2011) differs from stechowi in having
more lingual ridges (11 vs. 2-3); lingual ridges well developed apically; labial
ridges present.
Comments- Though referred to Labrosaurus stechowi by Janensch
(1925; as type a), Rauhut (2011) noted differences that suggest it may be a
different taxon. Another possibility is positional variation. Buffetaut (2008)
suggested this (and possibly MR B 1091) could be a basal spinosaurid, and all
Ostafrikasaurus teeth were proposed by Fowler (2007) as being potential
links between ceratosaurs and baryonychines. Rauhut found the shape, cross section
and serration morphology was the same as stechowi, that it lacks spinosaurid-style
granulated enamel and that unlike baryonychines the distal crown is least ridged.
Buffetaut (2011) later described it as a new species of spinosaurid- Ostafrikasaurus
crassiserratus. It is retained in Ceratosauridae here pending further comparison.
References- Janensch, 1925. Die Coelurosaurier und Theropoden der Tendaguru-Schichten
Deutsch-Ostafrikas. Palaeontographica. 1(supp. 7), 1-99.
Fowler, 2007. Recently rediscovered baryonychine teeth (Dinosauria Theropoda):
New morphologic data, range extension and similarity to ceratosaurs. Journal
of Vertebrate Paleontology. 27(3), 76A.
Fowler, 2007 online. http://www.denverfowler.com/publications/Fowler_2007_SVP.htm
Buffetaut, 2008. Spinosaurid teeth from the Late Jurassic of Tengaduru, Tanzania,
with remarks on the evolutionary and biogeographical history of the Spinosauridae.
In Mazin, Pouch, Hantzpergue and Lacombe (eds.). Mid-Mesozoic life and environments.
Documents des Laboratoires de Geologie Lyon, 164. 26-28.
Buffetaut, 2011. An early spinosaurid dinosaur from the Late Jurassic of Tendaguru
(Tanzania) and the evolution of the spinosaurid dentition. Oryctos. 10, 1-8.
Rauhut, 2011. Theropod dinosaurs from the Late Jurassic of Tendaguru (Tanzania).
Palaeontology. 86, 195-239.
O? stechowi (Janensch, 1920)
new combination
= Labrosaurus stechowi Janensch, 1920
= Allosaurus stechowi (Janensch, 1920) Glut, 1997
Late Kimmeridgian, Late Jurassic
Middle Dinosaur Member of the Tendaguru Formation, Tanzania
Lectotype- (MB R 1083; type b of Janensch) premaxillary tooth (45 mm)
Paralectotypes- (MB R 1086-1090, 1092) three premaxillary teeth, two
lateral teeth (36-38 mm)
Referred- (MB R 1093) tooth (Rauhut, 2011)
Diagnosis- (after Chure, 2000) differs from Ceratosaurus in having
mesial serrations on probable anterior premaxillary teeth; lower, less laterally
compressed lateral teeth; lingual ridges on lateral teeth.
Comments- Janensch (1920) originally described this taxon based on ten
syntype teeth and only illustrated MB R 1083. He later (1925) stated "The
teeth that I already designated as type-teeth of the species (1920, p. 233,
Fig. 7-8) comes from the Middle Saurian Bed in Mahimbwi Valley at Tendaguru",
referencing the illustration of MB R 1083. Rauhut (2011) listed that tooth as
the holotype, but this cannot be true as Janensch never chose a holotype in
1920 and one can not be subsequently designated. Janensch's 1925 statement is
provisionally considered a lectotype designation for MB R 1083, based on ICZN
Article 74.6.1.1- "The inference that the specimen is a "holotype"
or "the type" may be by reference to an illustration or description
of a specimen [Art. 74.4]."
Chure (2000) noted this resembles Ceratosaurus in the lingual fluting
of premaxillary teeth, but also that it differs in some ways. As the species
is diagnosable, it is not a "nomen vanum" (a term unused by the ICZN
but synonymous with nomen dubium), contra Madsen and Welles (2000). While Madsen
and Welles claimed stechowi is probably a junior synonym of roechlingi,
this is impossible as the latter was named five years later. It is possible
the species are synonymous though, as both seem to be ceratosaurid though they
cannot be compared.
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.
Glut, 1997. Dinosaurs - The Encyclopedia. McFarland Press. 1076 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.
Madsen and Welles, 2000. Ceratosaurus (Dinosauria, Theropoda) a revised
osteology. Miscellaneous Publication 00-2 Utah Geological Survey. 80 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.
Ceratosaurus Marsh, 1884
Diagnosis- (after Rauhut, 2000) narrow rounded horn core centrally placed
on the fused nasals; median oval groove on nasals behind horn core; premaxilla
with three teeth; premaxillary teeth with reduced extent of mesial serrations;
chevrons extremely long; pubis with large, rounded notch underneath the obturator
foramen; small epaxial osteoderms.
Ex-Ceratosaurus- Janensch (1920) tentatively referred three dorsals
from the Tendaguru Formation (MB R 1936, 2163 and a lost one previously called
TL 8) to Ceratosaurus, but Rauhut (2011) subsequently identified them
as tetanurine. Similarly, Janensch stated a femur (MB R 3621) and two tibiae
(MB R 3625, 3626) from that formation were very similar to Ceratosaurus,
but Rauhut found these to be more similar to abelisaurids. For Ceratosaurus
roechlingi (Janensch, 1925), see its entry as a possible ceratosaurid with
several misassigned or questionably assigned elements (Rauhut, 2011). White
(1964) assigned a maxilla (DNM coll.) to Ceratosaurus, but Madsen and
Welles (2000) reidentified it as Allosaurus. Paul (1988) referred Megalosaurus
ingens to Ceratosaurus, but Rauhut (2011) showed that these teeth
are more like derived carcharodontosaurids. Madsen and Welles refer Labrosaurus
stechowi to Ceratosaurus sp., but these differ and are all identified
as Ostafrikasaurus here (a genus proposed for some of the paratypes by
Buffetaut, 2011). Meyer and Thuring (2003) reported a supposedly Ceratosaurus-like
tooth from the Oxfordian Vellerat Formation of Switzerland, but Meyer later
doubted this (Meyer pers. comm. 2006 to Soto and Perea, 2008).
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.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
464 pp.
Madsen and Welles, 2000. Ceratosaurus (Dinosauria, Theropoda) a revised
osteology. Miscellaneous Publication 00-2 Utah Geological Survey. 80 pp.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria,
Saurischia). PhD thesis. University of Bristol. 440 pp.
Meyer and Thüring, 2003. Dinosaurs of Switzerland. Comptes Rendus Palevol.
2, 103-117.
Soto and Perea, 2008. A ceratosaurid (Dinosauria, Theropoda) from the Late Jurassic-Early
Cretaceous of Uruguay. Journal of Vertebrate Paleontology. 28(2), 439-444.
Buffetaut, 2011. An early spinosaurid dinosaur from the Late Jurassic of Tendaguru
(Tanzania) and the evolution of the spinosaurid dentition. Oryctos. 10, 1-8.
Rauhut, 2011. Theropod dinosaurs from the Late Jurassic of Tendaguru (Tanzania).
Palaeontology. 86, 195-239.
C. nasicornis Marsh, 1884
= Megalosaurus nasicornis (Marsh, 1884) Cope, 1892
= Labrosaurus "sulcatus" Marsh, 1896
= Labrosaurus sulcatus Marsh, 1896 vide Hay, 1908
= Ceratosaurus "dentisulcatus" Anonymous, 1995
= Ceratosaurus "willisobrienorum" Welles, Powell and Pickering
vide Pickering, 1995
= Ceratosaurus dentisulcatus Madsen and Welles, 2000
= Ceratosaurus magnicornis Madsen and Welles, 2000
Tithonian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Colorado, US (Callison's
Quarries, Dry Mesa Quarry, Marsh-Felch Quarry 1, Mygatt-Moore Quarry)
Holotype- (USNM 4735) (5.46 m, 524 kg, adult) skull (625 mm), mandibles
(650 mm), three teeth, hyoids, atlantal centrum, atlantal neural arches, axial
intercentrum, axis (88 mm), third cervical vertebra (60 mm), fourth cervical
vertebra (60 mm), fifth cervical vertebra (64 mm), sixth cervical vertebra (68
mm), seventh cervical vertebra (68 mm), partial eighth cervical vertebra, partial
ninth cervical neural spine, fragmentary cervical ribs, second dorsal vertebra
(62 mm), third dorsal vertebra (~73 mm), fourth dorsal vertebra (~83 mm), fifth
dorsal vertebra (~80 mm), ninth dorsal vertebra (~93 mm), tenth dorsal vertebra
(88 mm), eleventh dorsal vertebra (81 mm), twelfth dorsal vertebra (80 mm),
eleventh dorsal rib, first sacral vertebra (85 mm), second sacral vertebra (82
mm), fused third-seventh sacral vertebra (84, 73, 71, 72, 80 mm), first caudal
vertebra (79 mm), second caudal vertebra (80 mm), third caudal vertebra (80
mm), fourth caudal vertebra (79 mm), fifth caudal vertebra (78 mm), sixth caudal
vertebra (76 mm), seventh caudal vertebra (81 mm), eighth caudal vertebra (79
mm), ninth caudal vertebra (78 mm), tenth caudal vertebra (78 mm), eleventh
caudal vertebra (77 mm), twelfth caudal vertebra (76 mm), thirteenth caudal
vertebra (74 mm), fourteenth caudal vertebra, fifteenth caudal vertebra, sixteenth
caudal vertebra, seventeenth caudal vertebra, eighteenth caudal vertebra (62
mm), nineteenth caudal vertebra (58 mm), twentieth caudal vertebra, twenty-first
caudal vertebra, twenty-second caudal vertebra (59 mm), twenty-third caudal
vertebra (62 mm), twenty-fourth caudal vertebra (58 mm), twenty-fifth caudal
vertebra (58 mm), twenty-sixth caudal vertebra (56 mm), twenty-eighth caudal
vertebra (55 mm), twenty-ninth caudal vertebra (51 mm), thirtieth caudal vertebra
(48 mm), thirty-first caudal vertebra (49 mm), thirty-second caudal vertebra
(48 mm), thirty-third caudal vertebra (50 mm), thirty-fourth caudal vertebra
(47 mm), thirty-fifth caudal vertebra (47 mm), thirty-sixth caudal vertebra
(43 mm), thirty-seventh caudal vertebra (41 mm), thirty-eighth caudal vertebra
(40 mm), thirty-ninth caudal vertebra (39 mm), fourtieth caudal vertebra (39
mm), fourty-first caudal vertebra (36 mm), fourty-second caudal vertebra (35
mm), fourty-third caudal vertebra (30 mm), fourty-fourth caudal vertebra (29
mm), fourty-fifth caudal vertebra (32 mm), fourty-sixth caudal vertebra (28
mm), fourty-seventh caudal vertebra (26 mm), fourty-eighth caudal vertebra (24
mm), fourty-ninth caudal vertebra (20 mm), fifthieth caudal vertebra (18 mm),
fifty-first caudal vertebra (18 mm), second to ninth chevron (3rd 258, 6th 221
mm), ~fifteenth chevron (132 mm), twenty-ninth to fortieth chevrons (30th 53,
31st 49, 33rd 43, 35th 34, 39th 24, 40th 23 mm), proximal scapulocoracoid, radius
(149 mm), incomplete ulna (181 mm), metacarpals I (42 mm; one incomplete), metacarpals
II (69 mm; one distal), phalanx II-1 (28 mm; distal half lost), metacarpal III
(68 mm), phalanges III-1 (29, 32 mm), metacarpals IV (48, 48mm), phalanx IV-1
(18 mm), fused pelves (ilia, pubes and ischia incomplete; ilia ~620, pubes ~670,
ischia ~505 mm), femora (620 mm), tibia (555 mm), fibula, astragalocalcaneum
(130 mm wide), distal tarsal III, fused metatarsus (II 230, III 254, IV 220
mm), pedal ungual IV, several axial osteoderms
Referred- (BYUVP 4838) caudal vertebra (80 mm) (Britt, 1991)
(BYUVP 4853) anterior caudal vertebra (Britt, 1991)
(BYUVP 4908) caudal vertebra (95 mm) (Britt, 1991)
(BYUVP 4951) posterior dorsal vertebra (Britt, 1991)
(BYUVP 4952) posterior dorsal vertebra (96 mm) (Britt, 1991)
(BYUVP 5092) caudal vertebra (94 mm) (Britt, 1991)
(BYUVP 5103) middle caudal vertebra (62 mm) (Britt, 1991)
(BYUVP 5254) caudal vertebra (84 mm) (Britt, 1991)
(BYUVP 8907) dorsal vertebra (88 mm) (Britt, 1991)
(BYUVP 8910) middle caudal vertebra, (Britt, 1991)
(BYUVP 8937) caudal vertebra (92 mm) (Britt, 1991)
(BYUVP 8938) caudal vertebra (76 mm) (Britt, 1991)
(BYUVP 8974) posterior caudal vertebra (91 mm) (Britt, 1991)
(BYUVP 8982) caudal vertebra (70 mm) (Britt, 1991)
(BYUVP 9108) caudal vertebra (83 mm) (Britt, 1991)
(BYUVP 9136) caudal vertebra (Britt, 1991)
(BYUVP 9141) caudal vertebra (90 mm) (Britt, 1991)
(BYUVP 9142) middle or posterior dorsal vertebra (83 mm) (Britt, 1991)
(BYUVP 9143) middle or posterior dorsal vertebra (85 mm) (Britt, 1991)
(BYUVP 9144) middle or posterior dorsal vertebra (Britt, 1991)
(BYUVP 9152) caudal vertebra (Britt, 1991)
(BYUVP 9161) caudal vertebra (Britt, 1991)
(BYUVP 9162) caudal vertebra (Britt, 1991)
(BYUVP 9163) posterior caudal vertebra (36 mm) (Britt, 1991)
(BYUVP 5010) metatarsal III (Britt, 1991)
(BYUVP 5008) metatarsal III (Britt, 1991)
(BYUVP 13024) scapulocoracoid (Madsen and Welles, 2000)
(BYUVP 17550) five sacral vertebrae, ossified tendons, ilia, pubes, proximal ischia (Stiegler, 2019)
(BYUVP? coll.) teeth (Kirkland pers. comm. to Madsen and Welles, 2000)
(CM 21706) incomplete dentary (McIntosh, 1981)
(MWC 1; material of Ceratosaurus "willisobrienorum"; holotype
of Ceratosaurus magnicornis) (subadult) (5.6 m) skull (~600 mm), fifth
cervical vertebra (65 mm), sixth cervical vertebra (80 mm), seventh cervical
vertebra (75 mm), eighth cervical vertebra (60 mm), ninth cervical vertebra
(68 mm), tenth cervical vertebra, cervical rib, first dorsal vertebra (170 mm),
second dorsal vertebra (75 mm), third dorsal vertebra (65 mm), fourth dorsal
vertebra (65 mm), sixth dorsal vertebra (110 mm), seventh dorsal vertebra (100
mm), eighth dorsal vertebra (85 mm), ninth dorsal vertebra (95 mm), posterior
dorsal vertebra (98 mm), proximal caudal vertebra (70 mm), mid caudal vertebra,
four distal caudal vertebrae, mid chevron, humerus (292 mm), manual ungual,
incomplete femora (630 mm), tibiae (520 mm), astragalocalcaneum (127 mm wide),
metatarsal II, metatarsal III (234 mm), phalanx II-1, phalanx III-2, phalanx
IV-1, five dermal ossicles (Pickering, 1995; described by Madsen and Welles,
2000)
(MWC 9504) tooth (Kane, 2020)
(MWC coll.) sixteen teeth (Kane, 2020)
teeth (Madsen and Welles, 2000)
Tithonian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Oklahoma, US (Stovall's Pit
1)
(OMNH 1491) tooth (OMNH online)
(OMNH 73096) caudal vertebra (OMNH online)
? several teeth (~51 mm), cervical vertebra (89 mm), two incomplete dorsal vertebrae
(124 mm), several ribs (~300-500 mm), fused metatarsus fragment (Stovall, 1938)
Tithonian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Utah, US (Carnegie Quarry,
Cleveland-Lloyd Quarry, Sand Bench site)
(BYUVP 12893) (adult) (7.1 m) skull (~810 mm), seven fragmentary dorsal vertebrae,
sacrum, ossified tendons, incomplete pelvic elements including pubis and ischium
(Britt, Chure, Holtz, Miles and Stadtman, 2000)
(DNM 972) premaxilla (154 mm) (Madsen and Welles, 2000)
(UMNH 5278; = UMNH 6329; holotype of Ceratosaurus dentisulcatus) (6.7
m) premaxillae, maxilla, jugal, quadratojugal, quadrate, pterygoid, incomplete
dentaries, incomplete angular, incomplete splenials, teeth (to 93x30x15 mm),
atlantal intercentrum (28 mm), axis (89 mm), third cervical vertebra (65 mm),
fourth cervical vertebra, fifth cervical vertebra (90 mm), incomplete sixth
cervical vertebra (75 mm), seventh cervical vertebra (64 mm), eighth cervical
vertebra, ninth cervical vertebra (66 mm), tenth cervical vertebra (50 mm),
two incomplete cervical ribs, sixth dorsal vertebra (63 mm), eighth dorsal vertebra
(94 mm), incomplete tenth dorsal vertebra (99 mm), three incompete dorsal ribs,
eight proximal caudal vertebrae (102 mm), three mid caudal vertebrae (90 mm),
eleven distal caudal vertebrae (72 mm), two proximal chevrons (276 mm), three
mid chevrons, six distal chevrons, scapulocoracoid (405 mm), humerus (333 mm),
metacarpal II, phalanx II-1, metacarpal III, femur (759 mm), tibiae (594 mm),
fibulae (564 mm), astragalocalcaneum (165 mm wide), distal tarsal IV, metatarsal
IV, fifteen axial osteoderms (Madsen and Stokes, 1963; described by Madsen and
Welles, 2000)
material (Turner and Peterson, 1999)
Tithonian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Wyoming, US (Nail Quarry,
Reed’s Quarry 9)
(YPM 1936; holotype of Labrosaurus sulcatus) anterior dentary tooth (30x12.5x12
mm)
scapulocoracoid (Madsen and Welles, 2000)
Kimmeridigian, Late Jurassic
Salt Wash Member of the Morrison Formation, Wyoming, US
Material- (TPII coll.) (juvenile) (3.6 m) complete skull (413 mm), 30% complete skeleton
including vertebrae and pelvis (Britt, Cloward, Miles and Madsen, 1999)
Late Jurassic?
Morrison Formation?, US?
Material- ("YPM 4681") tibia (557.0 mm) (Carrano, 1998)
Comments- The holotype was discovered in 1883-1884. Initially described
by Marsh (1884) and Hay (1908), it was extensively described by Gilmore (1920)
and has had its forelimb recently redescribed by Carrano and Choiniere (2016).
While initially described as natural, the fused metatarsus is now seen as pathological.
Labrosaurus sulcatus is based on a tooth (YPM 1936) discovered in 1878
from Reed's Quarry 9, and first published as a figure caption by Marsh (1896).
As the caption doesn't qualify as as description, definition or indication (ICZN
Article 12.1), the name as published by Marsh is a nomen nudum (Rauhut, 2011).
Hay (1908) was thus the first author to officially name the species, as he provides
a description. Hay questioned the rationale for Marsh's referring it to Labrosaurus,
which is based on L. lucaris, not thought to preserve teeth at the time.
Chure (2000) discovered that three uncatalogued teeth belong to the L. lucaris
holotype, though these have never been described. Regardless, L. lucaris
is an allosaurid so is not congeneric with sulcatus (see below). Olshevsky
(1991) was the first author to synonymize it with Ceratosaurus nasicornis,
though Glut (1997) states Madsen informed him of this in 1986. Current authors
(e.g. Chure, 2000; Madsen and Welles, 2000) agree it cannot be distinguished
from other Ceratosaurus. Note while Gilmore (1920) and Chure identified
it as a premaxillary tooth, Madsen and Welles believe it is more probably from
the anterior dentary.
A specimen was discovered in 1976 from Callison's Quarries (Fruita area, Colorado)
and initially published as C. "willisobrienorum" by Pickering
(1995). It was described by Madsen and Welles (2000) as C. magnicornis.
The latter authors diagnosed it relative to C. nasicornis based on numerous
characters- more massive; having a longer and lower skull (H:L ratio 40); anterior
border of premaxilla straighter; maxilla longer (412 mm); maxillary anterior
edge almost vertical; maxillary ventral edge more convex; promaxillary fenestra
more deeply impressed; deeperpneumatic in maxillary ascending process; dorsal
edge of posterior maxillary process angles 15 degrees; anterior edge of maxilla
lower at front of fenestra; nasal horn core is longer and lower; teeth are longer
and stouter, especially posteriorly; lacrimal is more massive; lacrimal bears
high, rugose horn core; lacrimal horn has longer base; much larger lacrimal
recess; quadratojugal more massive ventrally; quadrate has much larger, lower
articular surface; quadrate body more concave posteriorly; dentary much more
concave dorsally and convex ventrally; dentary chin much more rounded; dentary
more massive; dentary 148 mm high at surangular contact at 546 mm from chin;
11-12 dentary teeth; sixth cervical 80 mm long; sixth cervical neural spine
much higher (145 mm); sixth cervical neural spine longer anteroposteriorly (52
mm); sixth cervical neural spine table slants more steeply posteriorly; infrapostzygapopyhseal
fossa much shorter; sixth cervical diapophysis much higher above parapophysis;
stout epipophysis on sixth cervical vertebra; femur 630 mm; femoral head 120
mm broad; distal femur 135 mm broad; femoral shaft 75 mm broad below trochanteric
shelf; femoral shaft straighter; tibia 520 mm long; cnemial crest less well
developed; proximal tibia 135 mm in diameter; distal tibia 132 mm in diameter;
astragalar ascending process completely fills tibial facet; calcaneum broader
anteriorly, occupying 43% of astragalocalcanear breadth; astragalocalcanear
suture runs dorsolaterally; calcaneum broader in lateral view. As Carrano and
Sampson (2008) note, many of these are merely measurements which are taxonomically
useless in differently sized individuals, while others are expected to vary
ontogenetically, and others are proportional differences expected to vary individually.
They synonymized the species, which is accepted here.
A Cleveland-Lloyd Quarry Ceratosaurus was first announced by Madsen and
Stokes (1963). The new name was first mentioned in a sales pamphlet (Anonymous,
1995), and Madsen and Welles (2000) officially erected C. dentisulcatus
for UMNH 5278. This specimen was found disarticulated during discontinuous excavations
starting in 1927, and each element has its own UUVP number as well. Carrano
and Choiniere (2016) reidentified the pedal phalanx listed by Madsen and Welles
as manual phalanx II-1. The species distinction from C. nasicornis was
based on numerous characters- ventral border of premaxilla arched and almost
horizontal; dorsal premaxillary process lower; body of premaxilla longer; premaxilla
with several large foramina; maxilla more massive; maxillary alveolar border
more convex; maxillary fossa more pronounced; posterior edge of maxillary ascending
process rises more steeply; front of antorbital fenestra wider; promaxillary
fenestra and pneumatic fossa in ascending process present; 12 maxillary teeth;
maxillary teeth more massive; maxillary teeth more strongly recurved; dentary
more massive; dentary more upturned from tooth 6 forward; dentary teeth more
massive; 11 dentary teeth; atlas-axis 100 mm long; odontoid more prominent;
axial centrum much shorter; axial centrum ventral edge less downcurved; axial
neural spine higher; axial neural spine very much shorter anteroposteriorly;
dorsal edge of axial neural spine straighter; no axial prezygapophysis; axial
epipophysis extending far behind neural spine; third cervical centrum shorter;
third cervical centrum ends almost vertical; third cervical spine shorter; third
cervical neural spine almost vertical; third cervical epipophysis much larger;
tibia longer (594 mm); tibia more massive; cnemial crest heavier; astragalar
buttress angles five degrees; distal tibia 165 mm broad; fibula longer (564
mm); proximal fibula vertical; proximal fibula broader; iliofibularis tubercle
angles posteriorly; iliofibularis tubercle's proximal edge projects anteriorly;
distal fibula broader (81 mm); distal fibula convex, evenly rounded; weak horizontal
groove across astragalar body; astragalar ascending process ossified. Carrano
and Sampson (2008) correctly dismiss these for the same reasons as in C.
magnicornis, and their synonymization is similarly accepted on this site.
Note Carrano and Choiniere (2016) accidentally listed UMNH 6329 as a separate
metacarpal II than that in the type material, but there is only one metacarpal
II present (Carrano, pers. comm. 2016).
Stovall (1938) describes several elements from Oklahoma as ?Ceratosaurus,
but Madsen and Welles (2000) rejected this and viewed the material as Theropoda
indet.. However, the illustrated tooth has the ceratosaurid character of high
labiolingual compression (BW/FABL ratio 31%), the cervical is anteriorly flat
unlike other large Morrison theropods (megalosaurids and allosaurids) and is
similar to Ceratosaurus in shape, and the metatarsals are fused. Thus
the material may be Ceratosaurus after all, but deserves further study.
BYUVP 5103 was initially assigned by Britt (1991) to Stokesosaurus, but
later to Ceratosaurus (Britt in Curtice and Wilhite, 1996). Stiegler (2019) writes "... Ceratosaurus (BYUVP 17550), and Eoabelisaurus
(MPEF PV 3990) all have ossified tendons adjacent to the lateral aspect
of the posterior dorsal, sacral, and/or anterior caudal neural spines",
but Rhodes and Currie's (2020) Table 1 lists "Allosaurus fragilis BYUVP 17550 Fused, partial pubes (apron and boot)". Scheetz (pers. comm. 7-2022) confirms "BYU 17550 is the Ceratosaurus pelvis (pelvis consisting of 5
sacral verts, both illia, fused pair of pubes and fused prox ends of
ischia) from the Brushy Basin Member of the Morrison, Dry Mesa Quarry,
western Colorado." Carrano (1998)
lists YPM 4681 as a Ceratosaurus nasicornis tibia in his table, but the
YPM online catalog shows no results for this number and no Ceratosaurus
material besides the sulcatus type tooth.
References- Dollo, 1884. Les métatarsiens du Ceratosaurus.
Revue des questions scientifiques. 16, 646-648.
Marsh, 1884. Principal characters of American Jurassic dinosaurs. Part VIII.
The order Theropoda. The American Journal of Science, series 3. 27, 329-340.
Marsh, 1884. On the united metatarsal bones of Ceratosaurus. The American
Journal of Science, series 3. 28, 161-162.
Cope, 1892. On the skull of the dinosaurian Laelaps incrassatus (Cope).
Proceedings of the American Philosophical Society. 30, 240-245.
Marsh, 1892. Restorations of Claosaurus and Ceratosaurus. The
American Journal of Science, series 3. 44, 343-350.
Marsh, 1893. Restorations of Anchisaurus, Ceratosaurus, and Claosaurus.
Geological Magazine, series 3. 10, 150-157.
Marsh, 1896. The dinosaurs of North America. U.S. Geological Survey, 16th Annual
Report. 1894-95, 133-244.
Hay, 1908. On certain genera and species of carnivorous dinosaurs, with special
reference to Ceratosaurus nasicornis Marsh. Proceedings of the United
States National Museum. 35, 351-366.
Gilmore, 1920. Osteology of the carnivorous Dinosauria in the United States
National Museum, with special reference to the genera Antrodemus (Allosaurus)
and Ceratosaurus. Bulletin of the United States National Museum. 110,
1-154.
Madsen and Stokes, 1963. New information on the Jurassic dinosaur Ceratosaurus.
Geological Society of America. Special Paper 73, 90.
McIntosh, 1981. Annotated catalogue of the dinosaur (Reptilia, Archosauria)
in the collections of the Carnegie Museum of Natural History. Bulletin of Carnegie
Museum of Natural History. 18, 1-67.
Britt, 1991. Theropods of Dry Mesa Quarry (Morrison Formation, Late Jurassic),
Colorado, with emphasis on the osteology of Torvosaurus tanneri. Brigham
Young University Geology Studies. 37, 1-72.
Olshevsky, 1991. A revision of the parainfraclass Archosauria Cope, 1869, excluding
the advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.
Anonymous, 1995. Price list of specimens available from Dinolab, Salt Lake City,
Utah.
Pickering, 1995. Jurassic Park: Unauthorized Jewish Fractals in Philopatry.
A Fractal Scaling in Dinosaurology Project, 2nd revised printing. Capitola,
California. 478 pp.
Curtice and Wilhite, 1996. A re-evaluation of the Dry Mesa Dinosaur Quarry sauropod
fauna with a description of juvenile sauropod elements. In Huffman, Lund and
Godwin (eds.). Geology and Resources of the Paradox Basin. Utah Geological Association
Guidebook. 25, 325-338.
Glut, 1997. Dinosaurs - The Encyclopedia. McFarland Press. 1076 pp.
Carrano, 1998. The evolution of dinosaur locomotion: Functional
morphology, biomechanics, and modern analogs. Volume One. PhD Thesis,
The University of Chicago. 424 pp.
Britt, Cloward, Miles and Madsen, 1999. A juvenile Ceratosaurus (Theropoda,
Dinosauria) from Bone Cabin Quarry West (Upper Jurassic, Morrison Formation),
Wyoming. Journal of Vertebrate Paleontology. 19(3), 33A.
Turner and Peterson, 1999. Biostratigraphy of dinosaurs in the Upper Jurassic
Morrison Formation of the Western Interior, U.S.A.. In Gillette (ed.). Vertebrate
Paleontology in Utah. Utah Geological Survey Miscellaneous Publication. 99-1,
77-114.
Welles and Pickering, 1999. An Extract From: Archosauromorpha: Cladistics and
Osteologies. 70 pp.
Britt, Chure, Holtz, Miles and Stadtman, 2000. A reanalysis of the phylogenetic
affinities of Ceratosaurus (Theropoda, Dinosauria) based on new specimens
from Utah, Colorado, and Wyoming. Journal of Vertebrate Paleontology. 20(3),
32A.
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.
Madsen and Welles, 2000. Ceratosaurus (Dinosauria, Theropoda) a revised
osteology. Miscellaneous Publication 00-2 Utah Geological Survey. 80 pp.
Sanders and Smith, 2005. The endocranium of the theropod dinosaur Ceratosaurus
studied with computed tomography. Acta Palaeontologica Polonica. 50(3), 601-616.
Carrano and Sampson, 2008 (online 2007). The Phylogeny of Ceratosauria (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 6, 183-236.
Rauhut, 2011. Theropod dinosaurs from the Late Jurassic of Tendaguru (Tanzania).
Palaeontology. 86, 195-239.
Carrano and Choiniere, 2015. New information on theropod forelimb evolution
from the forearm and manus of Ceratosaurus nasicornis (Dinosauria, Theropoda).
Journal of Vertebrate Paleontology. Program and Abstracts 2015, 103.
Carrano and Choiniere, 2016. New information on the forearm and manus of Ceratosaurus
nasicornis Marsh, 1884 (Dinosauria, Theropoda), with implications for theropod
forelimb evolution. Journal of Vertebrate Paleontology. 36(2), e1054497.
Stiegler, 2019. Anatomy, systematics, and paleobiology of noasaurid
ceratosaurs from the Late Jurassic of China. PhD thesis, The George
Washington University. 693 pp.
Kane, 2020. Identifying Jurassic theropod genera using GIS maps of tooth serrations. The Society
of Vertebrate Paleontology 80th
Annual Meeting, Conference Program. 197.
Rhodes and Currie, 2020. The homology, form, and function of the
microraptorine lateral pubic tubercle. Journal of Vertebrate
Paleontology. e1755866.
C. meriani (Greppin, 1870) new
comb.
= Megalosaurus meriani Greppin, 1870
= Labrosaurus meriani (Greppin, 1870) Janensch, 1920
= Antrodemus meriani (Greppin, 1870) Steel, 1970
= Allosaurus meriani (Greppin, 1870) Olshevsky, 1978
Early Kimmeridgian, Late Jurassic
Reuchenette Formation, Switzerland
Holotype- (MH 350) premaxillary tooth (60x18x? mm)
Diagnosis- Provisionally indeterminate relative to Ceratosaurus nasicornis.
Comments- Meyer and Thuring (2003) note this is probably not from the
Virgulla Beds as reported by Janensch and others.
Originally Megalosaurus meriani was described as including more teeth,
two caudals (MH 276, 280), a femur (MH 372) and an ?osteoderm, though the postcrania
were later removed as the holotype of Ornithopsis greppini once Janensch
realized it was sauropod in 1920/1921 (Huene, 1922).
Olshevsky (1978) refers this to Ceratosaurus based on resemblance to
"Labrosaurus" sulcatus. The tooth is nearly identical to the
first premaxillary tooth of Ceratosaurus dentisulcatus' holotype, and
differs from most other theropods in having lingual ridges. It differs from
Genyodectes and Ostafrikosaurus in lacking mesial serrations.
As it is of identical size and found in temporally equivalent beds, I believe
it should be called Ceratosaurus meriani, though the species is indeterminate
within the genus Ceratosaurus.
References- Greppin, 1870. Description geologique du Jura bernois et
de quelques districts adjacents. Beiträge zur geologischen Karte der Schweiz.
8, 1-357.
Janensch, 1920. Uber Elaphrosaurus bambergi und die Megalosaurier aus
den Tendaguru-Schichten Deutsch-Ostafricas. Sitzungsberichte Gesellschaft Naturforschender
Freunde Berlin. 8, 225-235.
Huene, 1922. Ueber einen Sauropoden im obern Malm des Berner Jura. Eclogae Geologicae
Helvetiae. 17, 80-94.
Steel, 1970. Part 14. Saurischia. Handbuch der Paläoherpetologie/Encyclopedia
of Paleoherpetology. Gustav Fischer Verlag, Stuttgart. 1-87.
Olshevsky, 1978. The archosaurian taxa (excluding the Crocodylia). Mesozoic
Meanderings. 1, 1-50.
Meyer and Thüring, 2003. Dinosaurs of Switzerland. Comptes Rendus Palevol.
2, 103-117.
C. sp. (Mateus and Antunes, 2000)
Late Kimmeridgian, Late Jurassic
Praia da Amoreira Member of Alcobaca Formation, Portugal
(IPFUB Gui D 191) tooth (Rauhut, 2000)
(IPFUB Gui D coll.) several teeth (Rauhut, 2000)
(ML 342) tooth (Mateus et al., 2006)
(ML 352) femur (647 mm), tibia (570 mm) (Mateus and Antunes, 2000)
....(SHN(JJS)-65) femur (650 mm), tibia (570 mm), partial fibula (Malafaia et
al., 2014)
(ML 737) tooth (Mateus et al., 2006)
(ML 809) tooth (Mateus et al., 2006)
Comments- Mateus and Antunes (2000) initially reported a femur and tibia
as cf. Ceratosaurus sp., while Rauhut (2000) referred several teeth to
Ceratosaurus. Antunes and Mateus (2003) illustrated the hindlimb elements
as Ceratosaurus sp.. Mateus et al. (2006) reported additional teeth and
ascribed the Portuguese material to C. dentisulcatus based on the "relative
position of the [tibial] epiphysial expansions" and "posterior intercondylar
bridge of the femur", but Carrano and Sampson (2008) state "the supposed
similarities are with distorted features of the right tibia of C. dentisulcatus;
the left tibia of the same individual is identical to those of other Ceratosaurus
'species'." Malafaia et al. (2014) later described additional hindlimb
elements from the same individual as ML 352, this time as "Ceratosaurus
aff. Ceratosaurus nasicornis, the only valid species recognised for the
genus." While it is probable the European material is taxonomically distinct
from C. nasicornis, this has yet to be demonstrated morphologically,
so they are retained as C. sp. here.
References- Mateus and Antunes, 2000. Late Jurassic dinosaurs of Portugal.
Abstracts of the First Symposium of European Dinosaurs. [pp?]
Rauhut, 2000. The dinosaur fauna from the Guimarota mine. In Martin and Krebs
(eds.). Guimarota - A Jurassic Ecosystem. Verlag Dr. Friedrich Pfeil. 75-82.
Antunes and Mateus, 2003. Dinosaurs of Portugal. Comptes Rendus Palevol. 2(1),
77-95.
Mateus, Walen and Antunes, 2006. The large theropod fauna of the Lourinha Formation
(Portugal) and its similarity to the Morrison Formation, with a description
of a new species of Allosaurus. In Foster and Lucas (eds.). Paleontology
and Geology of the Upper Jurassic Morrison Formation. New Mexico Museum of Natural
History and Science Bulletin. 36, 1-7.
Carrano and Sampson, 2008 (online 2007). The Phylogeny of Ceratosauria (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 6, 183-236.
Malafaia, Ortega, Escaso and Silva, 2015. New evidence of Ceratosaurus
(Dinosauria: Theropoda) from the Late Jurassic of the Lusitanian Basin, Portugal.
Historical Biology. 27(7), 938-946.
C? sp. (Soto and Perea, 2008)
Kimmeridgian-Tithonian, Late Jurassic
Lower member of Tacuarembo Formation, Uruguay
Material- (FC-DPV 1950) premaxillary tooth (15.2 x 6.5 x 5.3 mm)
(FC-DPV 2150) posterior premaxillary tooth (10 x 5.9 x 4.1 mm)
(FC-DPV coll.) lateral teeth
Comments- These teeth have serrationless mesial carinae, and are contemporaneus
with Ceratosaurus, so are tentatively referred to that genus here.
Reference- Soto and Perea, 2008. A ceratosaurid (Dinosauria, Theropoda)
from the Late Jurassic-Early Cretaceous of Uruguay. Journal of Vertebrate Paleontology.
28(2), 439-444.
Abelisauroidea Bonaparte and Novas, 1985
sensu Bonaparte, 1991
Definition- (Abelisaurus comahuensis <- Ceratosaurus nasicornis)
(modified from Holtz, 1994)
Other definitions- (Carnotaurus sastrei <- Ceratosaurus
nasicornis) (Tykoski and Rowe, 2004; modified from Padian, Hutchinson and Holtz, 1999)
(Carnotaurus sastrei + Noasaurus leali) (Wilson, Sereno, Srivastava, Bhatt, Khosla and Sahni, 2003)
(Abelisaurus comahuensis + Noasaurus leali) (Dal Sasso, Maganuco and Cau, 2018)
= "Abelisauroidea" Novas, 1989
= Abelisauroidea sensu Padian, Hutchinson and Holtz, 1999
Definition- (Carnotaurus sastrei <- Ceratosaurus nasicornis)
= Abelisauridae sensu Delcourt, 2018
Definition- (Carnotaurus sastrei <- Ceratosaurus nasicornis)
Comments- Novas (1989) proposed Abelisauroidea to group abelisaurids
with noasaurids.
References- Bonaparte and Novas, 1985. Abelisaurus comahuensis,
n. g., n. sp., Carnosauria from the Late Cretaceous of Patagonia. Ameghiniana.
21, 259-265.
Novas, 1989. Los dinosaurios carnivoros de la Argentina.
PhD thesis. Universidad Nacional de La Plata. 510 pp.
Bonaparte, 1991. The Gondwanian theropod families Abelisauridae and Noasauridae.
Historical Biology. 5, 1-25.
Holtz, 1994. The phylogenetic position of the Tyrannosauridae: Implications
for theropod systematics. Journal of Paleontology. 68(5), 1100-1117.
Padian, Hutchinson and Holtz, 1999. Phylogenetic definitions and nomenclature
of the major taxonomic categories of the carnivorous Dinosauria (Theropoda).
Journal of Vertebrate Paleontology. 19(1), 69-80.
Wilson, Sereno, Srivastava, Bhatt, Khosla and Sahni, 2003. A new abelisaurid
(Dinosauria, Theropoda) from the Lameta Formation (Cretaceous, Maastrichtian)
of India. Contributions from the Museum of Paleontology. The University of Michigan.
31, 1-42.
Tykoski and Rowe, 2004. Ceratosauria. In Weishampel, Dodson and Osmolska (eds.).
The Dinosauria Second Edition. University of California Press. 47-70.
Ezcurra and Agnolin, 2012. An abelisauroid dinosaur from the Middle Jurassic
of Laurasia and its implications on theropod palaeobiogeography and evolution.
Proceedings of the Geologists' Association. 123(3), 500-507.
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.
Delcourt, 2018. Ceratosaur palaeobiology: New insights on evolution and
ecology of the southern rulers. Scientific Reports. 8:9730.
de Souza-Júnior, Candeiro, da Silva Vidal, Brusatte and Mortimer, 2023.
Abelisauroidea (Theropoda, Dinosauria) from Africa: A review of the
fossil record. Papéis Avulsos de Zoologia. 63, e202363019.
unnamed Abelisauroidea (Lasseron, 2020)
Early Bathonian, Middle Jurassic
GEA 6, Guelb el Ahmar,
Anoual Formation, Morocco
Material- (MNHN GEA6-5) lateral tooth (2.58x1.86x1.39 mm)
Early Bathonian, Middle Jurassic
GEA 7, Guelb el Ahmar,
Anoual Formation, Morocco
(MNHN GEA7-14) lateral tooth (3.75x2.51x1.59 mm)
(MNHN GEA7-17) lateral tooth (2.60x2.06x1.09 mm)
Comments- Discovered in 2015 and/or 2018, Lasseron (2020) concluded "The convexity of the two carinae is a remarkable feature. It
recalls the Abelisauridae (Smith, 2007; Hendrickx & Mateus, 2014),
but also the Noasauridae (Carrano et al., 2002), an identification
which would be consistent with the small size of these teeth. We therefore attribute these teeth to an Abelisauroidea gen. and sp. indet." (translated).
Reference- Lasseron, 2020. Paleobiodiversite, evolution et paleobiogeographie des
vertebres mesozoiques africans et gondwaniens : apport des gisements du
Maroc oriental. Doctoral thesis, Museum National D'Histoire Naturelle.
493 pp.
unnamed abelisauroid (Lasseron, 2020)
Berriasian, Early Cretaceous
KM-A1, Ksar Metlili, Ksar Metlili Formation, Morocco
Material- (FSAC-KM-A1-12) lateral tooth (3.41x2.26x1.47 mm)
Comments- Collected in 2010, 2015 or 2018, Lasseron (2020) concluded "The convexity of the two carinae is a remarkable feature. It
recalls the Abelisauridae (Smith, 2007; Hendrickx & Mateus, 2014),
but also the Noasauridae (Carrano et al., 2002), an identification
which would be consistent with the small size of these teeth. We therefore attribute these teeth to an Abelisauroidea gen. and sp. indet." (translated).
Reference- Lasseron, 2020. Paleobiodiversite, evolution et paleobiogeographie des
vertebres mesozoiques africans et gondwaniens : apport des gisements du
Maroc oriental. Doctoral thesis, Museum National D'Histoire Naturelle.
493 pp.
undescribed possible abelisauroid (Smith, Lamanna, Dodson, Attia and Lacovara,
2001)
Cenomanian, Late Cretaceous
Bahariya Oasis, Baharija Formation, Egypt
Material- teeth (FABL 7.5 mm)
Comments- These "recently recovered" teeth are stated to have significantly more serrations per mm than similarly sized Deinonychus or Dromaeosaurus
and have "denticle morphologies and the average crown base width (CBW)
and CBL9crown base length] relationships ... more reminiscant of
abelisaurids than dromaeosaurids."
Reference- Smith, Lamanna, Dodson, Attia and Lacovara, 2001. Evidence
of a new theropod from the Late Cretaceous of Egypt. Journal of Vertebrate Paleontology.
21(3), 102A.
unnamed Abelisauroidea (Rauhut, 2005)
Late Kimmeridgian, Late Jurassic
Middle Dinosaur Member of the Tendaguru Formation, Tanzania
Material- (HMN MB.R 1750) tibia
Tithonian, Late Jurassic
Upper Dinosaur Member of the Tendaguru Formation, Tanzania
(HMN MB.R 1751) tibia (255 mm)
...?(HMN MB.R 1756) distal ischium
Comments- Originally described as 'coelurosaurier B and C', Rauhut (2005)
has identified the tibiae as abelisauroids. MB.R 1750 differs from 1751 in having
a sharp ridge extend proximally from the lateral malleolus, slightly broader
facet for the astragalar ascending process, and narrower lateral fibular facet.
They may be from different taxa, especially as MB.R 1750 comes from earlier
sediments. Carrano and Sampson (2008) stated "HMN MB.R.1756, a distal left
ischium, may also pertain to" Elaphrosaurus.
This element was figured by Rauhut (2005) who referred it to
Abelisauroidea and suggested it may belong to the same individual as
tibia HMN MB.R.1751 since it comes from the same locality and is of
similar size and preservation. It is roughly similar to Elaphrosaurus but has a
much deeper shaft relative to ischial foot depth.
References- Janensch, 1925. Die Coelurosaurier und Theropoden der Tendaguru-Schichten
Deutsch-Ostafrikas. Palaeontographica. Supplement VII, 1-99.
Rauhut, 2005. Post-cranial remains of 'coelurosaurs' (Dinosauria, Theropoda)
from the Late Jurassic of Tanzania. Geological Magazine. 142(1), 97-107.
Carrano and Sampson, 2008 (online 2007). The Phylogeny of Ceratosauria (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 6, 183-236.
Eoabelisaurus Pol and Rauhut,
2012
E. mefi Pol and Rauhut, 2012
Late Toarcian-Bajocian, Early-Middle Jurassic
Cañadón Asfalto Formation, Chubut, Argentina
Holotype- (MPEF PV 3990) (adult) maxillary fragment, posterior skull, five
cervical vertebrae, nine dorsal vertebrae, four dorsal ribs, sacrum (520 mm),
twenty-seven caudal vertebrae, twenty-sex chevrons, scapulocoracoids (~720 mm),
humeri (335 mm), radii (165 mm), ulnae (215 mm), distal carpals II, metacarpals
I, phalanx I-1, manual unguals I, metacarpals II (74 mm), phalanges II-1, metacarpals
III, phalanges III-1, phalanx III-2(?), metacarpals IV, phalanx, fused pelves,
femora (640 mm), tibiotarsi (550 mm), fibulae, metatarsals I, phalanx I-1, metatarsals
II, phalanx II-1, phalanx II-2, metatarsals III (317 mm), phalanges III-1, phalanx
III-2, phalanges III-3, pedal unguals III, metatarsals IV, phalanx IV-1, phalanx
IV-2, seven pedal phalanges, pedal ungual, metatarsals V, osteoderms
Diagnosis- (after Pol and Rauhut, 2012) quadrate with thickened medial
distal articular end and subparallel articular condyles; mid dorsal vertebrae
with double, V-shaped lamina extending from the parapophysis to the prezygodiapophyseal
lamina; ulna with hypertrophied olecranon process, accounting for more than
30% of its length; pubic foramen elongate, more than twice as long as high;
ambiens process of pubis developed as a large, anterolaterally directed, convex
expansion.
Comments- This specimen was discovered in 2009. Stiegler (2019) stated "Eoabelisaurus (MPEF PV 3990) [has] a single row of osteoderms dorsal to postaxial vertebrae."
Pol and Rauhut (2012)
found it to be a basal abelisaurid less derived than Kryptops and Rugops. However, only one extra step
was needed to place it as a non-abelisauroid abelisaurian. Novas et al. (2013)
noted several supposed abelisaurid characters are present in non-abelisaurids,
and the analysis of Tortosa et al. (2014) recovered it as sister to Abelisauria.
Reference- 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.
Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution of the carnivorous
dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research.
45, 174-215.
Tortosa, Buffetaut, Vialle, Dutour, Turini and Cheylan, 2014. A new abelisaurid
dinosaur from the Late Cretaceous of southern France: Palaeobiogeographical
implications. Annales de Paléontologie. 100(1), 63-86.
Stiegler, 2019. Anatomy, systematics, and paleobiology of noasaurid
ceratosaurs from the Late Jurassic of China. PhD thesis, The George
Washington University. 693 pp.
Ligabueino Bonaparte, 1996
= "Ligabueino" Bonaparte, 1995
L. andesi Bonaparte, 1996
Barremian, Early Cretaceous
Puesto Antigual Member of La Amarga Formation, Neuquen, Argentina
Holotype- (MACN-N 42) (.74 m) posterior cervical neural arch, dorsal
centrum, two posterior dorsal neural arches, caudal vertebra, two manual phalanges
III-?, ilium, incomplete pubes, femur (62 mm)
Comments- Bonaparte (1996) initially referred this to Abelisauroidea,
possibly as a noasaurid, an assignment with which Carrano and Sampson (2008)
concurred. Tortosa t al. (2014) recovered it as a noasaurid, but the more recent
version of this analysis in Filippi et al. (2016) found it to be a non-abelisaurian
abelisauroid closer to abelisaurs than Eoabelisaurus.
References- Bonaparte, 1995. Dinosaurios de America del Sur. Buenos Aires.
174 pp.
Bonaparte, 1996. Cretaceous tetrapods of Argentina. Muenchner Geowissenschaftliche
Abhandlungen. 30A, 73-130.
Carrano and Sampson, 2008 (online 2007). The Phylogeny of Ceratosauria (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 6, 183-236.
Agnolin and Chiarelli, 2010. The position of the claws in Noasauridae (Dinosauria:
Abelisauroidea) and its implications for abelisauroid manus evolution. Paläontologische
Zeitschrift. 84, 293-300.
Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution of the carnivorous
dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research.
45, 174-215.
Tortosa, Buffetaut, Vialle, Dutour, Turini and Cheylan, 2014. A new abelisaurid
dinosaur from the Late Cretaceous of southern France: Palaeobiogeographical
implications. Annales de Paléontologie. 100(1), 63-86.
Filippi, Mendez, Juarez Valieri and Garrido, 2016. A new brachyrostran with
hypertrophied axial structures reveals an unexpected radiation of latest Cretaceous
abelisaurids. Cretaceous Research. 61, 209-219.
Abelisauria Novas, 1992
Definition- (Abelisaurus comahuensis + Noasaurus leali)
(modified from Novas, 1997)
= Abelisauroidea sensu Wilson, Sereno, Srivastava, Bhatt, Khosla and Sahni, 2003
Definition- (Carnotaurus sastrei + Noasaurus leali)
= Abelisauroidea sensu
Dal Sasso, Maganuco and Cau, 2018
Definition- (Abelisaurus comahuensis + Noasaurus leali)
= Ceratosauroidea sensu Delcourt, 2018
Definition- (Carnotaurus sastrei + Noasaurus leali)
Diagnosis- (after Carrano et al., 2002) enlarged external mandibular
fenestra; anterior end of external mandibular fenestra ventral to last dentary
tooth; large socket in surangular for articulation with dentary; posteroventral
dentary process extends posteriorly subequally to posterodorsal process; cervical
neural spines anteroposteriorly short; humeral head globular; hypertrophied
and flange-like femoral medial epicondyle; fibula fused to astragalar ascending
process; double vascular grooves in pedal unguals; pedal ungual II asymmetrical.
References- Novas, 1992. La evolucion de los dinosaurios carnivoros. In Sanz and Buscalioni (eds.). Los Dinosaurios y Su Entorno Biotico:
Actas del Segundo Curso de Paleontologia in Cuenca. Instituto "Juan Valdez",
Cuenca, Argentina. 126-163.
Novas, 1997. Abelisauridae. In Currie and Padian (eds.). Encyclopedia of Dinosaurs. Academic Press. 1-2.
Wilson, Sereno, Srivastava, Bhatt, Khosla and Sahni, 2003. A new abelisaurid
(Dinosauria, Theropoda) from the Lameta Formation (Cretaceous, Maastrichtian)
of India. Contributions from the Museum of Paleontology. The University of Michigan.
31, 1-42.
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.
Delcourt, 2018. Ceratosaur palaeobiology: New insights on evolution and
ecology of the southern rulers. Scientific Reports. 8:9730.
Abelisauria incertae sedis
Betasuchus Huene, 1932
B. bredai (Seeley, 1883) Huene, 1932
= Megalosaurus bredai Seeley, 1883
Maastrichtian, Late Cretaceous
Maastricht Beds, Netherlands
Holotype- (NHMUK 42997) incomplete femur (353.0 mm)
Comments- The term Ornithomimidorum is not a generic name, but rather
the latinized form of ornithomimid. Ornithomimidorum gen. B refers to Huene
transfering Megalosaurus bredai to the Ornithomimidae.
References- Seeley, 1883. On the dinosaurs from the Maastricht beds.
Quarterly Journal of the Geological Society of London. 39, 246-253.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte.
Monographien zur Geologie und Palaeontologie. 4(1), viii + 361 pp.
unnamed possible abelisaurian (Lydekker, 1879)
Campanian-Maastrichtian, Late Cretaceous
Ariyalur Group, Trichinopoli, India
Material- (Indian Museum coll.) tooth (~41x23x10 mm)
Comments- Lydekker (1879) described this as a "tooth of a species of Megalosaurus", stating that compared to the first maxillary tooth of M. bucklandii
(OUMNH J.13506 as poorly illustrated by Phillips 1871- Diagram LVI),
"the former is somewhat less curved than the latter, while in cross
section the antero-posterior diameter of the former is of considerably
less relative length than in the latter." He concluded "it
indicates that it belonged
to a species distinct from the European species; on such very slender
evidence, however, I do not think it advisable to assign a distinct
name to the former."
Gemmellaro (1921) later stated "The Arialur-Group of Trichinopolis
(British Indies) tooth, illustrated by Lydekker and generically
referred to as Megalosaurus, must in fact belong to the Megalosaurus crenatissimus.
The slightly bigger dimensions of the Indian sample, compared to the
Madagascar one, are confirmed by some of the teeth I have studied"
[Egyptian MGUP coll. Abelisauridae indet. here] (translated).
However, most Majungasaurus teeth have less distal concavity as typical of abelisaurid teeth, with the seventh maxillary tooth being most similar.
References- Lydekker, 1879. Fossil Reptilia and Batrachia. Memoirs of
the Geological Survey of India. Palaeontologia Indica, Series IV. Indian Pretertiary
Vertebrata. 1(3), 1-36.
Gemmellaro, 1921. Rettili maëstrichtiani d'Egitto. Giornale di Scienze
Naturali ed Economiche. 32, 339-351.
unnamed probable Abelisauria (Huene and Matley, 1933)
Late Maastrichtian, Late Cretaceous
Lameta Formation, India
Material- (GSI 296) proximal femur (Novas et al., 2004)
(GSI K19/574) mid caudal vertebra (70 mm) (Huene and Matley, 1933)
(GSI K20/336A) distal metatarsal III (Huene and Matley, 1933)
(GSI K20/336B) distal metatarsal III (Huene and Matley, 1933)
(GSI K20/337 A) pedal phalanx (Huene and Matley, 1933)
(GSI K20/362) mid chevron (Huene and Matley, 1933)
(GSI K20/396) unknown element (previously identified as calcaneum) (Huene and
Matley, 1933)
(GSI K20/399) pedal ungual (~140 mm) (Huene and Matley, 1933)
(GSI K20/612 in part) distal caudal vertebra (35 mm) (Huene and Matley, 1933)
(GSI K20/615; syntype of Dryptosauroides grandis) proximal dorsal rib
(Huene and Matley, 1933)
(GSI K20/620) tooth (Huene and Matley, 1933)
(GSI K20/625) pedal ungual (Huene and Matley, 1933)
(GSI K20/670) two teeth
(GSI K27/476) basioccipital
(GSI K27/524) pedal phalanx (Huene and Matley, 1933)
(GSI K27/527) articular (Huene and Matley, 1933)
(GSI K27/530) caudal vertebra (Huene and Matley, 1933)
(GSI K27/533) four sacral centra (Huene and Matley, 1933)
(GSI K27/534) pedal phalanx IV-? (Huene and Matley, 1933)
(GSI K27/535) jugal fragment (lost) (Huene and Matley, 1933)
(GSI K27/536) caudal vertebra (115 mm) (Huene and Matley, 1933)
(GSI K27/537) pedal ungual IV
(GSI K27/540) partial fibula (Huene and Matley, 1933)
(GSI K27/542) distal caudal vertebra (25 mm) (Huene and Matley, 1933)
(GSI K27/543) pedal ungual (Huene and Matley, 1933)
(GSI K27/545) mid chevron (Huene and Matley, 1933)
(GSI K27/546) proximal ischium (Huene and Matley, 1933)
(GSI K27/547; syntype of Dryptosauroides grandis) dorsal rib
(GSI K27/551) pedal ungual (Huene and Matley, 1933)
(GSI K27/552; lost) tibia (430 mm) (Huene and Matley, 1933)
(GSI K27/554) four sacral centra (Huene and Matley, 1933)
(GSI K27/555; syntype of Dryptosauroides grandis) incomplete anterior
cervical vertebra (120 mm) (Huene and Matley, 1933)
(GSI K27/556; lost) tibia (Huene and Matley, 1933)
(GSI K27/557) distal metatarsal III (Huene and Matley, 1933)
(GSI K27/559) unknown element (originally identified as ilial fragment) (Huene
and Matley, 1933)
(GSI K27/560; lost) femur (Huene and Matley, 1933)
(GSI K27/561) distal caudal vertebra (65 mm) (Huene and Matley, 1933)
(GSI K27/563; lost) femur (Huene and Matley, 1933)
(GSI K27/564; lost) femur (Huene and Matley, 1933)
(GSI K27/566) mid chevron (120 mm) (Huene and Matley, 1933)
(GSI K27/567) ectopterygoid (Huene and Matley, 1933)
(GSI K27/569; lost) femur (Huene and Matley, 1933)
(GSI K27/572; lost) cervical vertebra (75 mm) (Huene and Matley, 1933)
(GSI K27/573) dentary (lost) (Huene and Matley, 1933)
(GSI K27/576) basioccipital (Huene and Matley, 1933)
(GSI K27/579) tooth (34 x 20 x 9 mm) (Huene and Matley, 1933)
(GSI K27/581) partial jugal (lost) (Huene and Matley, 1933)
(GSI K27/582) tooth (35 x 17 x 6.5 mm) (Huene and Matley, 1933)
(GSI K27/583) five teeth including premaxillary tooth (37 x 13.7 x 20 mm) (Huene
and Matley, 1933)
(GSI K27/584) tooth (Huene and Matley, 1933)
(GSI K27/585) tooth (Huene and Matley, 1933)
(GSI K27/590) partial anterior dorsal vertebra (~90 mm) (Huene and Matley, 1933)
(GSI K27/591) axial centrum (Huene and Matley, 1933)
(GSI K27/592) vertebra (Huene and Matley, 1933)
(GSI K27/593) caudal vertebra (Huene and Matley, 1933)
(GSI K27/598) first sacral centrum (~120 mm) (Huene and Matley, 1933)
(GSI K27/603) caudal vertebra (Huene and Matley, 1933)
(GSI K27/605) distal caudal vertebra (38 mm) (Huene and Matley, 1933)
(GSI K27/606) distal caudal vertebra (60 mm) (Huene and Matley, 1933)
(GSI K27/607) distal caudal vertebra (70 mm) (Huene and Matley, 1933)
(GSI K27/608) distal caudal vertebra (40 mm) (Huene and Matley, 1933)
(GSI K27/609) distal caudal vertebra (45 mm) (Huene and Matley, 1933)
(GSI K27/610) distal caudal vertebra (45 mm) (Huene and Matley, 1933)
(GSI K27/611) distal caudal vertebra (55 mm) (Huene and Matley, 1933)
(GSI K27/612) caudal vertebra (Huene and Matley, 1933)
(GSI K27/613) vertebra (Huene and Matley, 1933)
(GSI K27/615) vertebra (Huene and Matley, 1933)
(GSI K27/616) mid caudal vertebhra (80 mm) (Huene and Matley, 1933)
(GSI K27/617) caudal vertebra (Huene and Matley, 1933)
(GSI K27/620) incomplete fibula (Huene and Matley, 1933)
(GSI K27/621; lost) distal femur (Huene and Matley, 1933)
(GSI K27/623; syntype of Dryptosauroides grandis) proximal dorsal rib
(Huene and Matley, 1933)
(GSI K27/624; syntype of Dryptosauroides grandis) proximal dorsal rib
(Huene and Matley, 1933)
(GSI K27/625; syntype of Dryptosauroides grandis) proximal dorsal rib
(Huene and Matley, 1933)
(GSI K27/626) limb bone (lost) (originally identified as tibia)
(GSI K27/627; lost; caudal vertebra in Huene and Matley, 1933) proximal femur
(Huene and Matley, 1933)
(GSI K27/628) basioccipital
(GSI K27/630) pedal ungual (Huene and Matley, 1933)
(GSI K27/631) pedal ungual (Huene and Matley, 1933)
(GSI K27/633; manual ungual in Huene and Matley, 1933) pedal ungual II (60 mm)
(Huene and Matley, 1933)
(GSI K27/634) pedal ungual III
(GSI K27/635) pedal ungual II
(GSI K27/636) pedal ungual (Huene and Matley, 1933)
(GSI K27/639) pedal phalanx (Huene and Matley, 1933)
(GSi K27/640) pedal phalanx (Huene and Matley, 1933)
(GSI K27/641) pedal phalanx (Huene and Matley, 1933)
(GSI K27/642) pedal phalanx IV-I (31 mm) (manual in Huene and Matley, 1933)
(GSI K27/643) pedal phalanx (Huene and Matley, 1933)
(GSI K27/645) unknown element (originally identified as manual phalanx) (Huene
and Matley, 1933)
(GSI K27/649) pedal phalanx (Huene and Matley, 1933)
(GSI K27/650) pedal phalanx (Huene and Matley, 1933)
(GSI K27/651) pedal phalanx I-1 (70 mm) (Huene and Matley, 1933)
(GSI K27/652) pedal phalanx I-1 (70 mm) (Huene and Matley, 1933)
(GSI K27/655) pedal phalanx (Huene and Matley, 1933)
(GSI K27/656) distal metatarsal IV (Huene and Matley, 1933)
(GSI K27/657) distal phalanx (Huene and Matley, 1933)
(GSI K27/660) metatarsal IV (240 mm) (Huene and Matley, 1933)
(GSI K27/661) metatarsal IV (180 mm) (Huene and Matley, 1933)
(GSI K27/662; lost) tibia (200 mm) (Huene and Matley, 1933)
(GSI K27/663) metatarsal IV (220 mm) (Huene and Matley, 1933)
(GSI K27/664) metatarsal IV (Huene and Matley, 1933)
(GSI K/27/668) carpal or tarsal (Huene and Matley, 1933)
(GSI K27/669) distal limb bone (originally identified as tibia) (Huene and Matley,
1933)
(GSI K27/670) tibia (lost)
(GSI K27/672) incomplete proximal chevron (Huene and Matley, 1933)
(GSI K27/673) distal chevron (Huene and Matley, 1933)
(GSI K27/674) distal chevron (Huene and Matley, 1933)
(GSI K27/675) mid chevron (Huene and Matley, 1933)
(GSI K27/675A) mid chevron (Huene and Matley, 1933)
(GSI K27/676) mid chevron (Huene and Matley, 1933)
(GSI K27/677) distal chevron (Huene and Matley, 1933)
(GSI K27/678) incomplete mid chevron (Huene and Matley, 1933)
(GSI K27/679) mid chevron (Huene and Matley, 1933)
(GSI K27/680) distal chevron (Huene and Matley, 1933)
(GSI K27/683) caudal vertebra (Huene and Matley, 1933)
(GSI K27/686) proximal ischium (Huene and Matley, 1933)
(GSI K27/687) basioccipital (Huene and Matley, 1933)
(GSI K27/688) unknown element (originally identified as ectopterygoid) (Huene
and Matley, 1933)
(GSI K27/689) ectopterygoid (Huene and Matley, 1933)
(GSI K27/691) distal quadrate (Huene and Matley, 1933)
(GSI K27/692) partial medial gastralium (Huene and Matley, 1933)
(GSI K27/693) surangular (Huene and Matley, 1933)
(GSI K27/694) pedal phalanx (Huene and Matley, 1933)
(GSI K27/698) ectopterygoid (Huene and Matley, 1933)
(GSI K27/699) caudal vertebra (Huene and Matley, 1933)
(GSI K27/700) tooth (Huene and Matley, 1933)
(GSI K27/701) tooth (Huene and Matley, 1933)
(GSI K27/702) tooth (Huene and Matley, 1933)
(GSI K27/703) tooth (Huene and Matley, 1933)
(GSI K27/707) partial lateral gastralium (Huene and Matley, 1933)
(GSI K27/708) unknown element (originally identified as partial lacrimal) (Huene
and Matley, 1933)
(GSI K27/711) angular fragment (Huene and Matley, 1933)
(GSI K27/712) distal caudal vertebra (55 mm) (Huene and Matley, 1933)
Comments- Basioccipital GSI K27/687 shows the exoccipitals floored the
foramen magnum, and has an elongate occipital condyle neck, unlike GSI K27/628.
Cervical vertebra GSI K27/572 differs from abelisauroids in its axially elongate
and transversely narrow neural spine and and poorly developed epipophyses. Sacral
vertebrae GSI K27/533 and 554 resemble Masiakasaurus, Rajasaurus and
material referred to Lametasaurus in their transverse width and clear
contact between successive centra. At least one chevron has open haemal canals,
like Aucasaurus, but unlike Carnotaurus and Ilokelesia.
Femora GSI K/560, 563, 564, 569, 621 and 627 are slender, unlike the robust
femora assigned to Abelisauridae. They may belong to Noasauridae. Much of the
material above is probably indeterminate at levels higher than Abelisauroidea,
but is retained in this section for simplicity and because all diagnostic Lameta
Formation theropod material appears to belong to this clade.
References- Huene and Matley, 1933. The Cretaceous Saurischia and Ornithischia
of the central provinces of India. Palaeontologica Indica. 21, 1-74.
Novas, Agnolin and Bandyopadhyay, 2004. Cretaceous theropods from India: A review
of specimens described by Huene and Matley (1933). Revista del Museo Argentino
del Ciencias Naturales. 6(1), 67-103.
unnamed Abelisauria (Azevedo, Simbras, Furtado, Candeiro and Bergqvist,
2012)
Campanian-Maastrichtian, Late Cretaceous
Alfredo Marcondes, Presidente Prudente Formation, Brazil
Material-
(UFRJ-DG 598-R) tooth fragment (Alves, Bergqvist and Brito, 2016)
(UFRJ-DG 599-R) tooth fragment (Alves, Bergqvist and Brito, 2016)
Campanian-Maastrichtian, Late Cretaceous
Alvares Machado, Presidente Prudente Formation, Brazil
(UFRJ-DG 487-R) tooth (Alves, Bergqvist and Brito, 2016)
Campanian-Maastrichtian, Late Cretaceous
Flórida Paulista area (FP1 site), Presidente Prudente Formation, Brazil
(UFRJ-DG 254-R) ilial fragment (Azevedo, Simbras, Furtado, Candeiro
and Bergqvist, 2012)
Comments- These were all referred to Abelisauroidea.
References- Azevedo, Simbras, Furtado, Candeiro and Bergqvist, 2012.
First Brazilian carcharodontosaurid and other new theropod dinosaur fossils
from the Campanian-Maastrichtian Presidente Prudente Formation, São Paulo
State, southeastern Brazil. Cretaceous Research. 40, 131-142.
Alves, Bergqvist and Brito, 2016. New occurrences of microvertebrate fossil
accumulations in Bauru Group, Late Cretaceous of western São Paulo state,
Brazil. Journal of South American Earth Sciences. 69, 80-90.
Noasauridae Bonaparte and Powell, 1980
Definition- (Noasaurus leali <- Carnotaurus sastrei)
(Wilson et al., 2003)
Other definitions- (Noasaurus leali <- Coelophysis bauri,
Carnotaurus sastrei, Passer domesticus) (Sereno, in press)
= Noasaurinae Bonaparte and Powell, 1980 sensu Paul, 1988
Definition- (Noasaurus leali <- Elaphrosaurus bambergi, Abelisaurus comahuensis, Ceratosaurus nasicornis, Allosaurus fragilis) (modified from Rauhut and Carrano, 2016)
Other definitions- (Noasaurus leali <- Elaphrosaurus bambergi) (modified from Delcourt, 2018)
= Velocisauridae Bonaparte, 1991
= Velocisaurinae Bonaparte, 1991 sensu Agnolin, Novas and Apesteguia, 2003
= Noasauridae sensu Sereno, in press
Definition- (Noasaurus leali <- Coelophysis bauri, Carnotaurus
sastrei, Passer domesticus)
Diagnosis- (after Carrano et al., 2002; Sereno et al., 2004) palatal
process of maxilla simple; anteroventral border of antorbital fenestra demarcated
by a raised ridge; interdental plates obscured medially; presacral vertebrae
without posterior pleurocoels; cervical neural spines located in the anterior
half of centra; distal condyle of metatarsal IV <50% width of metatarsal
II distal condyle.
Comments- Neither Coelophysis nor Passer seem particularily
useful as external specifiers in Sereno's (in press) redefinition, as noasaurids
have been universally considered abelisauroids since Paul (1988) and Bonaparte
et al. (1990), and were never placed anywhere else besides the polyphyletic
pre-cladistic Coelurosauria.
References- 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, 546-610.
Delcourt, 2018. Ceratosaur palaeobiology: New insights on evolution and
ecology of the southern rulers. Scientific Reports. 8:9730.
unnamed Noasauridae (Russell, 1996)
Cenomanian, Late Cretaceous
Kem Kem beds, Morocco
Material- (CMN 41873; bone taxon H) distal humerus (48 mm wide) (Russell,
1996)
?(CMN 50811; bone taxon B) (~2.4 m; juvenile) posterior cervical centrum (34 mm) (Russell,
1996)
(FSAC-KK-5016) (~1.5 m) incomplete ~fourth cervical vertebra (21 mm) (Smyth, Ibrahim, Kao and Martill, 2020)
(ROM 64666) (~2.1 m; <1 year old juvenile) femur (204 mm) (Evans, Barrett, Brink
and Carrano, 2015)
Comments- The humerus is very similar to Masiakasaurus.
The cervical CMN 50811 was considered a possible indeterminate
noasaurid by McFeeters (2013). Smyth et al. agreed it was
ceratosaurian but stated "the specimen lacks any noasaurid
synapomorphies so can only be referred to Abelisauroidea indet.."
Evans et al. (2015) described femur ROM 64666 as Noasauridae gen. et
sp. indet., proposing it may be a juvenile Deltadromeus
based on its young age, small size and supposedly medially inclined
femoral head. However, the head is declined in its medioal
portion as in other noasaurids while that of Deltadromeus
is incomplete dorsally so indeterminate for this trait. The lack
of an anteriorly projected medial condyle may be due to erosion, but
the well preserved posterolateral edge shows no trace of Deltadromeus'
enlarged M. adductor femoris 2 insertion scar. This could
plausibly be ontogenetic, but in the absence of positive evidence the
femur is retained as Noasauridae indet. here. Smyth et al.
described cervical FSAC-KK-5016 and recovered it as a noasaurid.
They believed it was most similar to Masiakasaurus
but differed in having- articular surfaces of centrum approximately as
tall as wide; anterior articular surface of centrum convex;
prezygapophyseal facets not medially angled; ventral surface of centrum
rounded and with keel.
References- Russell, 1996. Isolated dinosaur bones from the Middle Cretaceous
of the Tafilalt, Morocco. Bulletin du Museum national d'Histoire naturelle.
18, 349-402.
McFeeters, 2013. Bone "taxon" B: Reevaluation of a supposed small
theropod dinosaur from the Mid-Cretaceous of Morocco. Kirtlandia. 58, 38-41.
Evans, Barrett, Brink and Carrano, 2015 (online 2014). Osteology and bone microstructure of
new, small theropod dinosaur material from the early Late Cretaceous of Morocco.
Gondwana Research. 27(3), 1034-1041.
Smyth, Ibrahim, Kao and Martill, 2020 (online 2019). Abelisauroid
cervical vertebrae from the Cretaceous Kem Kem beds of southern Morocco
and a review of Kem Kem abelisauroids. Cretaceous Research. 108, 104330.
unnamed Noasauridae (Huene and Matley, 1933)
Late Maastrichtian, Late Cretaceous
Lameta Formation, India
Material- (GSI K20/337B) pedal phalanx IV-2 (24 mm) (Huene and Matley,
1933)
(GSI K20/337C) distal metatarsal IV (metatarsal I of Huene and Matley, 1933)
(GSI K20/619) incomplete premaxilla (Huene and Matley, 1933)
(GSI K20/626B) pedal phalanx IV-1 (48 mm) (manual in Huene and Matley, 1933)
(GSI K27/524) pedal phalanx II-1 (56 mm), pedal ungual IV (Huene and Matley,
1933)
(GSI K27/525) pedal phalanx III-1 (76 mm)
(GSI K27/526) tibia (330 mm) (Huene and Matley, 1933)
(GSI K27/587) mid caudal vertebra (80 mm) (Huene and Matley, 1933)
(GSI K27/589) mid caudal vertebra (90 mm) (Huene and Matley, 1933)
(GSI K27/599) mid caudal vertebra (85 mm) (Huene and Matley, 1933)
(GSI K27/629) pedal ungual III (Huene and Matley, 1933)
(GSI K27/632) pedal ungual II or IV (Huene and Matley, 1933)
(GSI K27/637) pedal phalanx IV-4 (28 mm) (Huene and Matley, 1933)
(GSI K27/638) pedal phalanx IV-3 (38 mm) (Huene and Matley, 1933)
(GSI K27/644) pedal phalanx III-1 (46 mm) (Huene and Matley, 1933)
(GSI K27/646) pedal phalanx III-2 or 3 (36 mm) (Huene and Matley, 1933)
(GSI K27/647) pedal phalanx IV-4 (Huene and Matley, 1933)
(GSI K27/648) pedal phalanx IV-I (26 mm) (manual in Huene and Matley, 1933)
(GSI K27/659) metatarsal IV (Huene and Matley, 1933)
(GSI K27/665) distal metatarsal III (Huene and Matley, 1933)
(GSI K27/666) distal metatarsal IV (metacarpal in Huene and Matley, 1933)
(GSI K27/667) distal metatarsal II (manual in Huene and Matley, 1933)
(GSI K27/671) distal metatarsal II (Huene and Matley, 1933)
(GSI K27/681) metatarsal III (Huene and Matley, 1933)
....(GSI K27/697) metatarsal III (Huene and Matley, 1933)
(GSI K27/684; identified as an astragalus by Huene and Matley, 1933) quadrate
(Huene and Matley, 1933)
material including dentary (Wilson, 2012)
Comments- The premaxilla GSI K20/619 is excluded from Abelisauridae due
to the lack of external texturing. The quadrate GSI K27/684 is excluded due
to the lack of fusion with the quadratojugal. These are provisionally assumed
to be noasaurid based on the apparent presence of only abelisaurians in the
Lameta Forrmation. The pedal elements closely resemble Velocisaurus,
so maybe from a noasaurid more derived than Deltadromeus. Carrano et
al. (2011) noted they were more robust than Masiakasaurus. Carrano et
al. reidentified GSI K27/648 as IV-1 (not IV-3), 646 as III-2 or 3 (not 1),
644 as III-1 (not III-3) and 629 as pedal ungual III (not IV). Wilson (2012)
noted new noasaurid remains including a dentary with procumbant anterior teeth
as in Masiakasaurus. The above material may be referrable to Laevisuchus,
Jubbulpuria, Ornithomimoides and/or Coeluroides.
References- Huene and Matley, 1933. The Cretaceous Saurischia and Ornithischia
of the central provinces of India. Palaeontologica Indica. 21, 1-74.
Novas, Agnolin and Bandyopadhyay, 2004. Cretaceous theropods from India: A review
of specimens described by Huene and Matley (1933). Revista del Museo Argentino
del Ciencias Naturales. 6(1), 67-103.
Carrano, Loewen and Sertich, 2011. New materials of Masiakasaurus knopfleri
Sampson, Carrano, and Forster, 2001, and implications for the morphology of
the Noasauridae (Theropoda: Ceratosauria). Smithsonian Contributions to Paleobiology.
95, 53 pp.
Wilson, 2012. Small theropod dinosaurs from the Latest Cretaceous of India.
Journal of Vertebrate Paleontology. Program and Abstracts 2012, 194.
undescribed Noasauridae (Canale, Apesteguia, Gallina, Haluza, Gianechini
and Pazo, 2014)
Late Berriasian-Valanginian, Early Cretaceous
Bajada Colorada Formation, Neuquen, Argentina
Material- (MMCh-PV-67) (~2 m) axis
(MMCh-PV-68-2) tooth
(MMCh-PV-68-3) tooth
(MMCh-PV-68-4) tooth
(MMCh-PV-68-5) tooth
Reference- 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.
undescribed possible noasaurid (Novas, Cladera and Puerta, 1996)
Cenomanian-Early Coniacian, Late Cretaceous
Rio Neuquen Subgroup, Neuquen, Argentina
Material- proximal humerus
Comments- Novas et al. (1996) mentioned this as an ornithomimid based
on the ball-shaped articular head and shallow deltopectoral crest. However,
these character are also found in abelisauroids, which are common in Late Cretaceous
South America, unlike ornithomimosaurs.
Reference- Novas, Cladera and Puerta, 1996. New theropods from the Late
Cretaceous of Patagonia. Journal of Vertebrate Paleontology. 16(3), 56A.
Noasauridae indet. (Motta, Aranciaga Rolando, Rozadilla, Agnolin, Chimento, Brisson
Egli and Novas, 2016)
Middle Cenomanian-Early Turonian, Late Cretaceous
Huincul Formation of the Rio Limay Subgroup, Rio Negro, Argentina
Material- (MPCA-Pv 800) distal quadrate
(MPCA-PV 801) metatarsal IV
Comments- Motta et al. say these are similar to noasaurids and
abelisaurids, referring them to Abelisauroidea gen. et sp. indet., but
the lack of quadrate-quadratojugal fusion and the very slender
metatarsal IV are like noasaurids.
Reference- Motta, Aranciaga Rolando, Rozadilla, Agnolin, Chimento, Brisson
Egli and Novas, 2016. New theropod fauna from the Upper Cretaceous (Huincul
Formtation) of northwestern Patagonia, Argentina. In Khosla and Lucas (eds.).
Cretaceous period: Biotic diversity and biogeography. New Mexico Museum of Natural
History and Science Bulletin. 71, 231-253.
unnamed noasaurid (Novas, Agnolin,
Rozadilla, Aranciaga-Rolando, Brissón-Eli, Motta, Cerroni, Ezcurra,
Martinelli, D'Angelo, Álvarez-Herrera, Gentil, Bogan, Chimento,
García-Marsà, Lo Coco, Miquel, Brito, Vera, Perez Loinaze, Fernandez
and Salgado, 2019)
Late Campanian-Early Maastrichtian, Late Cretaceous
Chorrillo Formation, Santa Cruz, Argentina
Material- (MPM 21547) pedal phalanx IV-2? (15 mm)
Comments- Discovered between
January and March 2019, Novas et al. assign this to Noasauridae.
Reference- Novas, Agnolin,
Rozadilla, Aranciaga-Rolando, Brissón-Eli, Motta, Cerroni, Ezcurra,
Martinelli, D'Angelo, Álvarez-Herrera, Gentil, Bogan, Chimento,
García-Marsà, Lo Coco, Miquel, Brito, Vera, Perez Loinaze, Fernandez
and Salgado, 2019. Paleontological discoveries in the Chorrillo
Formation (upper Campanian-lower Maastrichtian, Upper Cretaceous),
Santa Cruz Province, Patagonia, Argentina. Revista del Museo Argentino
de Ciencias Naturales. 21(2), 217-293.
unnamed noasaurid (Lindoso, Medeiros, Carvalho and Marinho, 2012)
Cenomanian, Late Cretaceous
Alcantara Formation, Brazil
Material- (UFMA 1.20.554) nine teeth (6-16 mm)
Reference- Lindoso, Medeiros, Carvalho and Marinho, 2012. Masiakasaurus-like
theropod teeth from the Alcântara Formation, São Luís Basin
(Cenomanian), northeastern Brazil. Cretaceous Research. 36, 119-124.
unnamed noasaurid (Brum, Machado, Campos and Kellner, 2015)
Late Santonian-Campanian, Late Cretaceous
Adamantina Formation of the Bauru Group, Brazil
Material- (DGM 929-R) (~2-3 m, ~20 kg) incomplete mid cervical vertebra (43.7 mm)
Comments- This vertebra was
donated to the DGM in 1950 and described in detail by Brum et al.
(2018). They added it to Tortosa et al.'s ceratosaur analysis and
recovered it as a noasaurid.
References- Brum, Machado, Campos and Kellner, 2015. The first record
of Noasauridae (Theropoda) from the Adamantina Formation (Campanian-Maastrichtian),
Bauru Group, Brazil. Journal of Vertebrate Paleontology. Program and Abstracts
2015, 98.
Brum, Machado, Campos and Kellner, 2018. Description of uncommon
pneumatic structures of a noasaurid (Theropoda, Dinosauria) cervical
vertebra from the Bauru Group (Upper Cretaceous), Brazil. Cretaceous
Research. 85, 193-206.
unnamed probable noasaurid (Fitzgerald, Carrano, Holland, Wagstaff,
Pickering, Rich and Vickers-Rich, 2012)
Early Aptian, Early Cretaceous
San Remo Member of Wonthoggi Formation of Strzelecki Group, Victoria, Australia
Material- (NMV P221202) tibial fragment, astragalocalcaneum (60 mm wide)
Comments- Discovered on October 15 2006, Fitzgerald et al. (2012) described this and referred it to
Ceratosauria, most similar to Elaphrosaurus, Deltadromeus
and noasaurids. Novas et al. (2013) questioned this identification,
only placing it as Averostra indet.. Brougham et al. (2020)
effectively argued against Novas et al.'s critiques, and included it in
Cau's megamatrix to recover it as a noasaurid
References- Fitzgerald, Carrano, Holland, Wagstaff, Pickering, Rich and
Vickers-Rich, 2012. First ceratosaurian dinosaur from Australia. Naturwissenschaften.
99, 397-405.
Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution of the carnivorous
dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research.
45, 174-215.
Brougham, Smith and Bell, 2020. Noasaurids are a component of the
Australian 'mid'-Cretaceous theropod fauna. Scientific Reports. 10:1428.
unnamed noasaurid (Poropat, Pentland, Duncan, Bevitt, Vickers-Rich and Rich, 2020)
Early Albian, Early Cretaceous
Eumeralla Formation of the Otway Group, Victoria, Australia
Material- (NMV P252004) (~2 m) ~fifth cervical vertebra (38.69 mm)
Comments- Poropat et al. (2020) added this to Rauhuit and Carrano's ceratosaur analysis and recovered it as an elaphrosaurine sister to Elaphrosaurus plus CCG 20011.
Reference- Poropat, Pentland,
Duncan, Bevitt, Vickers-Rich and Rich, 2020. First elaphrosaurine
theropod dinosaur (Ceratosauria: Noasauridae) from Australia - A
cervical vertebra from the Early Cretaceous of Victoria. Gondwana
Research. 84, 284-295.
unnamed noasaurid (Brougham, Smith and Bell, 2020)
Early Cenomanian, Late Cretaceous
Wallangulla Sandstone Member of Griman Creek Formation, New South Wales, Australia
Material- (LRF 3050.AR) mid-posterior cervical centrum (75 mm)
Comments- Brougham et al. (2020)
included this in
Cau's megamatrix to recover it as a noasaurid.
References-
Brougham, Smith and Bell, 2020. Noasaurids are a component of the
Australian 'mid'-Cretaceous theropod fauna. Scientific Reports. 10:1428.
Afromimus Sereno, 2017
A. tenerensis Sereno, 2017
Aptian-Albian, Early Cretaceous
El Rhaz Formation, Niger
Holotype- (MNBH GAD112) (adult)
dorsal rib fragment, posterior ?nineteenth caudal vertebra, ~twentieth
caudal vertebra (52.9 mm), partial ?twenty-second caudal vertebra
(~49.4 mm), ~twenty-fourth caudal vertebra (50.9 mm), posterior
~twenty-sixth caudal vertebra, ~twenty-eighth caudal vertebra (~40 mm),
~thirty-first caudal vertebra (39.6 mm), two partial proximal chevrons,
incomplete tibiotarsus (tibia 404.5 mm), incomplete fibula (~389.8 mm),
distal pedal phalanx II-2, incomplete pedal ungual II (32.2 mm),
proximal phalanx III-1
Diagnosis- (after Cerroni et
al., 2019) posterodorsal process on chevrons larger than anterodorsal
process; anterior rim of proximomedial fibular fossa extensively covers
fossa;
Other diagnoses- Sereno (2017) proposed several characters as diagnostic of an ornithomimosaurian Afromimus
that Cerroni et al. (2019; asterisked characters first noted by Cerroni
et al. 2018 in their meeting abstract) noted are common in ceratosaurs-
rugose texture on non-articular surfaces of pre- and postzygapophyses
of mid caudal vertebrae (also in Ceratosaurus, Majungasaurus and Ekrixnatosaurus); elevated zygapophyseal facets on pre-and
postzygapophyses of mid caudal vertebrae (also in Ceratosaurus and Viavenator); fused pedicels on proximal chevrons that are
less than one half as long anteroposteriorly as broad transversely* (also in Ceratosaurus, Masiaksaurus and Majungasaurus); large elliptical attachment scar on posterior tibial shaft distal to lateral condyle* (also in Masiakasaurus, Xenotarsosaurus, Ekrixinatosaurus, Skorpiovenator and Carnotaurus); rugose attachment surface on fibula posterior to tibial crest (also in Ceratosaurus, Masiakasaurus, Arcovenator and Skorpiovenator); trough on anteromedial fibular shaft ventral to iliofibularis tubercle (also in Ceratosaurus, Masiakasaurus and Skorpiovenator).
Comments- This was discovered in 1997 and described by Sereno (2017) as an ornithomimosaur more basal than Harpymimus or Shenzhousaurus.
Cerroni et al. (2018) first noted in an abstract some characters were
shared with ceratosaurs (asterisked above and below) and proposed it
was an abelisauroid. The resulting 2019 publication of Cerroni et
al. found that all of Afromimus'
proposed diagnostic characters among ornithomimosaurs were common in
ceratosaurs, and that other characters supported this too (short mid
caudal prezygapophyses; laterally bowed tibial shaft; proximally
extended fibular crest*; medial buttress for astragalar ascending
process on tibia; large m. iliofibularis tubercle on fibula*;
transversely broad distal fibula; astragalocalcaneum fused to tibia;
narrow and low astragalar ascending process*). Contrary to both
Sereno and Cerroni et al. (2019), I don't think the fibular crest is
placed notably more distally than in Gallimimus. Also, a slight overlap of the astragalar ascending process by the fibula is common in tetanurines (e.g. Ornithomimus velox),
even if it was first described in ceratosaurs sensu lato.
Finally, while Cerroni et al. (2019) claim Sereno said some
ornithomimosaurs have a dorsal vascular groove on their pedal unguals,
Sereno actually says it has "not been reported before among
ornithomimosaurs", so this would be another ceratosaur-like character
of Afromimus. Note Sereno assigned caudal numbers "based on other ornithomimosaurs (e.g., Harpymimus; Kobayashi and Barsbold, 2005b)", whereas I have revised them based on comparison to Masiakasaurus so that e.g. caudal 16 is now caudal 20.
Cerroni et al. (2018) initially added Afromimus
to Carrano's tetanurine matrix where it emerged as a noasaurid, though
no maniraptoriforms such as ornithomimosaurs were included. They
later (2019) recovered it more generally as an abelisauroid in that
analysis, and also tested it in Choiniere's coelurosaur matrix where it
emerged as a noasaurid. The latter matrix has numerous scoring
issues however, and forcing it to be ornithomimosaurian only took four
extra steps. Cerroni et al. (2019) finally added it to Pol and
Rauhut's ceratosaur matrix and found it to be in a polytomy with other
noasaurids, but of course no ornithomimosaurs were included. The
maniraptoromorph matrix of Hartman et al. (2019) recovered Afromimus as a noasaurid sister to Deltadromeuswith
extensive taxon sampling despite the absence of characters
designed to group ceratosaurs, though that portion of the tree was not
figured. Baiano et al. (2020) added Afromimus to Wang et al.'s
basal theropod analysis and Cau's megamatrix and recovered it as a
noasaurine in the first and an abelisauroid in the second.
References-
Sereno, 2017. Early Cretaceous ornithomimosaurs (Dinosauria: Coelurosauria) from Africa. Ameghiniana. 54, 576-616.
Cerroni, Agnolin, Brissón Egli and Novas, 2018. The phylogenetic affinities of Afromimus tenerensis Sereno, 2017. XXXII Jornadas Argentinas de Paleontologia de Vertebrados. R13.
Cerroni, Agnolin, Brissón Egli and Novas, 2019. The phylogenetic position of Afromimus tenerensis Sereno, 2017 and its paleobiogeographical implications. Journal of African Earth Sciences. 159, 103572.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247.
Baiano, Coria and Cau, 2020. A new abelisauroid (Dinosauria: Theropoda)
from the Huincul Formation (lower Upper Cretaceous, Neuquén Basin) of
Patagonia, Argentina. Cretaceous Research. 110, 104408.
Coeluroides Huene and Matley,
1933
= "Coeluroides" Huene, 1932
C. largus Huene and Matley, 1933
= "Coeluroides largus" Huene, 1932
Maastrichtian, Late Cretaceous
Lameta Formation, India
Syntypes- (GSI K27/562) proximal caudal vertebra (92 mm)
(GSI K27/574) proximal caudal vertebra
Referred- (AMNH 1957) caudal vertebra (Chatterjee, 1978)
?(GSI K27/695) partial vertebra (Huene and Matley, 1933)
Diagnosis- (after Novas et al., 2004) wide, almost horizontally oriented
and well separated mid-caudal zygapophyses; expanded and triangular mid caudal
transverse processes with deeply excavated dorsal surface; transversely robust
and axially elongate mid caudal neural spines.
Comments- Novas et al. (2004) believe the specimens closely resemble
AMNH 1957 and Jubbulpuria, so may be synonymous with the latter, Compsosuchus
and/or Laevisuchus. However, Wilson (2012) believes this is synonymous
with (both species of?) Ornithomimoides as they share autapomorphies
and are distinct from Jubbulpuria. He proposed sinking Ornithomimoides
into Coeluroides, as the latter has page priority, but this is not acknowledged
by the ICZN. As first revisor (Article 24.2.2), his determination would have
force under the ICZN once properly published though. The taxa are kept separate
here pending publication of Wilson's detailed reasoning.
References- Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre
Entwicklung und Geschichte. Monographien zur Geologie und Palaeontologie. 4(1),
viii + 361 pp.
Huene and Matley, 1933. The Cretaceous Saurischia and Ornithischia of the central
provinces of India. Palaeontologica Indica. 21, 1-74.
Novas, Agnolin and Bandyopadhyay, 2004. Cretaceous theropods from India: A review
of specimens described by Huene and Matley (1933). Revista del Museo Argentino
del Ciencias Naturales. 6(1), 67-103.
Wilson, 2012. Small theropod dinosaurs from the Latest Cretaceous of India.
Journal of Vertebrate Paleontology. Program and Abstracts 2012, 194.
Compsosuchus Huene and Matley,1933
= "Compsosuchus" Huene, 1932
C. solus Huene and Matley, 1933
= "Compsosuchus solus" Huene, 1932
Maastrichtian, Late Cretaceous
Lameta Formation, India
Holotype- (GSI K27/578) atlas, axis (40 mm)
Referred- (ISI I91/1) atlas, axis (Chatterjee and Rudra, 1996)
Comments- Novas et al. (2004) concluded that Compsosuchus is an
indeterminate abelisaurid, since there are apparently no major differences from
an atlas-axis complex referred to Indosaurus by Chatterjee and Rudra
(1996). However, the latter is not necessarily Indosaurus, and may be
Compsosuchus, making this reason moot. Carrano et al. (2011) described
the axis of Masiakasaurus and noted Compsosuchus' was very similar
except for a slightly more upturned atlantal intercentrum and a proportionally
less projected odontoid, thus placing the taxon in Noasauridae instead.
References- Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre
Entwicklung und Geschichte. Monographien zur Geologie und Palaeontologie. 4(1),
viii + 361 pp.
Huene and Matley, 1933. The Cretaceous Saurischia and Ornithischia of the central
provinces of India. Palaeontologica Indica. 21, 1-74.
Chatterjee and Rudra, 1996. KT events in India: Impact, rifting, volcanism and
dinosaur extinction. In Novas and Molnar (eds.). Proceedings of the Gondwanan
Dinosaur Symposium. Memiors of the Queensland Museum. 39(3), 489-532.
Novas and Bandyopaphyay, 1999. New approaches on the Cretaceous theropods from
India. VII International Symposium on Mesozoic Terrestrial Ecosystems, abstracts.
46-47.
Novas, Agnolin and Bandyopadhyay, 2004. Cretaceous theropods from India: A review
of specimens described by Huene and Matley (1933). Revista del Museo Argentino
del Ciencias Naturales. 6(1), 67-103.
Carrano, Loewen and Sertich, 2011. New materials of Masiakasaurus knopfleri
Sampson, Carrano, and Forster, 2001, and implications for the morphology of
the Noasauridae (Theropoda: Ceratosauria). Smithsonian Contributions to Paleobiology.
95, 53 pp.
Jubbulpuria Huene and Matley,
1933
= "Jubbulpuria" Huene, 1932
J. tenuis Huene and Matley, 1933
= "Jubbulpuria tenuis" Huene, 1932
Maastrichtian, Late Cretaceous
Lameta Formation, India
Syntypes- (GSI K20/612 in part) mid caudal vertebra (18 mm)
(GSI K27/614; lost) mid caudal vertebra (42 mm)
Referred- (GSI K27/599) distal caudal vertebra (Huene and Matley, 1933)
Diagnosis- (after Wilson, 2012) in prep.
Comments- Novas et al. (2004) determined the vertebrae were distal caudals,
not dorsals as suggested by Huene and Matley (1933). Carrano et al. (2011) refined
their placement to mid caudals. Jubbulpuria may be synonymous with Coeluroides
since both taxa possess large, triangular, dorsally excavated transverse processes.
Both may be synonymous with the noasaurid Laevisuchus based on resemblence
to Masiakasaurus and Ligabueino. Wilson (2012) proposed Jubbulpuria
may be synonymous with Leavisuchus as they are of similar size and differ
in features attributable to intracolumnar variation, though they lack overlapping
remains. He also stated Jubbulpuria is diagnostic.
References- Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre
Entwicklung und Geschichte. Monographien zur Geologie und Palaeontologie. 4(1),
viii + 361 pp.
Huene and Matley, 1933. The Cretaceous Saurischia and Ornithischia of the central
provinces of India. Palaeontologica Indica. 21, 1-74.
Novas and Bandyopaphyay, 1999. New approaches on the Cretaceous theropods from
India. VII International Symposium on Mesozoic Terrestrial Ecosystems, abstracts.
46-47.
Novas, Agnolin and Bandyopadhyay, 2004. Cretaceous theropods from India: A review
of specimens described by Huene and Matley (1933). Revista del Museo Argentino
del Ciencias Naturales. 6(1), 67-103.
Carrano, Loewen and Sertich, 2011. New materials of Masiakasaurus knopfleri
Sampson, Carrano, and Forster, 2001, and implications for the morphology of
the Noasauridae (Theropoda: Ceratosauria). Smithsonian Contributions to Paleobiology.
95, 53 pp.
Wilson, 2012. Small theropod dinosaurs from the Latest Cretaceous of India.
Journal of Vertebrate Paleontology. Program and Abstracts 2012, 194.
Ornithomimoides? barasimlensis
Huene and Matley, 1933
= "Ornithomimoides barasimlensis" Huene, 1932
Maastrichtian, Late Cretaceous
Lameta Formation, India
Syntypes- (GSI K27/531) proximal caudal vertebra
(GSI K27/541) proximal caudal vertebra
(GSI K27/604) proximal caudal vertebra (55 mm)
(GSI K27/682) proximal caudal vertebra (50 mm)
Diagnosis- (After Wilson, 2012) Possible junior synonym of Coeluroides
largus.
Comments- Novas et al. (2004) determined the vertebrae were proximal
caudals, not dorsals as suggested by Huene and Matley (1933). They suggested
the material is indistinguishable from Dryptosauroides and Ornithomimoides
mobilis. Carrano et al. (2011) retained GSI K/531 and 541 as dorsals, but
604 as a mid caudal. Wilson (2012) synonymized this with Coeluroides
(and presumably O. mobilis) based on shared autapomorphies and differences
from Jubbulpuria, seemingly as noasaurids. He proposed sinking Ornithomimoides
into Coeluroides, as the latter has page priority, but this is not acknowledged
by the ICZN. As first revisor (Article 24.2.2), his determination would have
force under the ICZN once properly published though. The taxa are kept separate
here pending publication of Wilson's detailed reasoning.
References- Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre
Entwicklung und Geschichte. Monographien zur Geologie und Palaeontologie. 4(1),
viii + 361 pp.
Huene and Matley, 1933. The Cretaceous Saurischia and Ornithischia of the central
provinces of India. Palaeontologica Indica. 21, 1-74.
Novas and Bandyopaphyay, 1999. New approaches on the Cretaceous theropods from
India. VII International Symposium on Mesozoic Terrestrial Ecosystems, abstracts.
46-47.
Novas, Agnolin and Bandyopadhyay, 2004. Cretaceous theropods from India: A review
of specimens described by Huene and Matley (1933). Revista del Museo Argentino
del Ciencias Naturales. 6(1), 67-103.
Carrano, Loewen and Sertich, 2011. New materials of Masiakasaurus knopfleri
Sampson, Carrano, and Forster, 2001, and implications for the morphology of
the Noasauridae (Theropoda: Ceratosauria). Smithsonian Contributions to Paleobiology.
95, 53 pp.
Wilson, 2012. Small theropod dinosaurs from the Latest Cretaceous of India.
Journal of Vertebrate Paleontology. Program and Abstracts 2012, 194.
Ornithomimoides Huene and
Matley, 1933
= "Ornithomimoides" Huene, 1932
O. mobilis Huene and Matley, 1933
= "Ornithomimoides mobilis" Huene, 1932
Maastrichtian, Late Cretaceous
Lameta Formation, India
Syntypes- (GSI K20/610) proximal caudal vertebra
(GSI K20/614B) proximal caudal vertebra
(GSI K27/586) proximal caudal vertebra (85 mm)
(GSI K27/597) proximal caudal vertebra
(GSI K27/600) proximal caudal vertebra
Diagnosis- (After Wilson, 2012) Possible junior synonym of Coeluroides
largus.
Comments- Novas et al. (2004) determined the vertebrae were proximal
caudals, not dorsals as suggested by Huene and Matley (1933). They suggested
the material is indistinguishable from Dryptosauroides and Ornithomimoides?
barasimlensis. Wilson (2012) synonymized this with Coeluroides (and
presumably O? barasimlensis) based on shared autapomorphies and differences
from Jubbulpuria, seemingly as noasaurids. He proposed sinking Ornithomimoides
into Coeluroides, as the latter has page priority, but this is not acknowledged
by the ICZN. As first revisor (Article 24.2.2), his determination would have
force under the ICZN once properly published though. The taxa are kept separate
here pending publication of Wilson's detailed reasoning.
References- Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre
Entwicklung und Geschichte. Monographien zur Geologie und Palaeontologie. 4(1),
viii + 361 pp.
Huene and Matley, 1933. The Cretaceous Saurischia and Ornithischia of the central
provinces of India. Palaeontologica Indica. 21, 1-74.
Novas and Bandyopaphyay, 1999. New approaches on the Cretaceous theropods from
India. VII International Symposium on Mesozoic Terrestrial Ecosystems, abstracts.
46-47.
Novas, Agnolin and Bandyopadhyay, 2004. Cretaceous theropods from India: A review
of specimens described by Huene and Matley (1933). Revista del Museo Argentino
del Ciencias Naturales. 6(1), 67-103.
Wilson, 2012. Small theropod dinosaurs from the Latest Cretaceous of India.
Journal of Vertebrate Paleontology. Program and Abstracts 2012, 194.
"Sidormimus" Molina-Perez and Larramendi, 2019
= "Dogosaurus" Anonymous, 2000 online
Aptian-Albian, Early Cretaceous
Gadoufaoua, Elrhaz Formation, Niger
Material- (MNN GAD coll.) (~1 m) partial skeleton including maxilla, cervical ribs, dorsal vertebrae,
dorsal ribs, uncinate processes, sacral vertebrae, scapula, coracoid, sternal
plates, sternal ribs, humerus, radius, ulna, carpus, manus including manual
digit II, phalanges, manual unguals and manual claw sheath, pelvis including
pubis and hindlimb including femur, tibia, fibula, calcaneum, metatarsus, pedal digit II, pedal
digit III, pedal digit IV and pedal unguals
Comments-
This specimen was discovered on September 13 2000 and announced by Lyon (2000 online) as "a brand
new, dog-sized theropod, which the team
has affectionately (but unofficially) named
"Sidormimus."" and a "new,
dog-sized carnivore - perhaps one of the smallest dinosaurs ever", with
a photo of the specimen in situ. The same photo was labeled Dogosaurus on a dispatch from Project
Exploration on the National Geographic website, which also stated it was "no more than three feet long. Its slender neck
and ribcage were preserved poking out of the rock. Its long, clawed
hind leg was uncovered as we dug around the exposed bones."
Sereno et al. (2004) first announced the
specimen in print, as "a small (1 m) articulated skeleton showing many
abelisauroid and noasaurid synapomorphies, including pneumatized
presacral and sacral neural arches, proportionately long presacral
centra, and others" and "preserves a maxilla and pelvic girdle, the
former of which bears the distinctive abelisaurid pit-and-groove
texturing of the skull bones." They include it in their cladogram
as "Gadoufaoua noasaurid", but it is not an OTU in the matrix. It
is also listed
in Sereno and Brusatte (2008) as "undescribed noasaurid" in a faunal
list, and is noted to have a pubic boot more with limited expansion
than the "Kryptops" postcrania MNN GAD1-2. Sidor
(pers. comm. 2005) confirms the "Sidormimus" specimen is the Elrhaz
noasaurid. Sereno (2010) devoted an SVP abstract to it, noting the specimen has- long and
robust posterior cervical ribs, dorsal centra more than twice as long
as tall, five uncinate processes (unique among non-maniraptoran
theropods), posteriorly directed glenoid, enlarged coracoid,
ossified sternal plates and ribs, forelimb 18% of hindlimb length,
robust deltopectoral crest and olecranon process, manual digit II
longer than III, straight manual unguals, tibia longer than femur,
pedal digits II and IV much shorter than III, and short flat pedal
unguals. Sereno (2017) provided further information- straight
tibia, weak iliofibularis tubercle on fibula, fibula distally fused
with calcaneum, no dorsal vascular groove on pedal unguals. The
co-occuring Afromimus
shared fibular-calcaneum fusion but differs in the other characters, in
addition to having a more robust proximal fibula and distal fibula
which is expanded transversely. Molina-Perez and Larramendi
(2019) first used the name in print, specified to be a nomen nudum.
References- Lyon, online 2000. http://web.archive.org/web/20121024124915/http://www.projectexploration.org/niger2000/9_15_2000.htm
Anonymous, online 2000.
http://web.archive.org/web/20001208070300/http://www.nationalgeographic.com/dinoquest/profile_01_dispatch2b.html
Sereno, Wilson and Conrad, 2004. New dinosaurs link southern landmasses in the
Mid-Cretaceous. Proceedings of the Royal Society, Series B. 271, 1325-1330.
Sereno and Brusatte, 2008. Basal abelisaurid and carcharodontosaurid theropods
from the Lower Cretaceous Elrhaz Formation of Niger. Acta Palaeontologica Polonica.
53(1), 15-46.
Keillor, Sereno and Masek, 2010. Range of movement in a noasaurid forelimb:
In situ data and joint reconstruction. Journal of Vertebrate Paleontology. Program
and Abstracts 2010, 114A.
Sereno, 2010. Noasaurid (Theropoda: Abelisauroidea) skeleton from Africa shows
derived skeletal proportions and function. Journal of Vertebrate Paleontology.
Program and Abstracts 2010, 162A.
Sereno, 2017. Early Cretaceous ornithomimosaurs (Dinosauria: Coelurosauria) from Africa. Ameghiniana. 54, 576-616.
Molina-Perez and Larramendi, 2019. Dinosaur Facts and Figures: The
Theropods and Other Dinosauriformes. Princeton University Press. 288 pp.
Laevisuchus Huene and Matley, 1933
= "Laevisuchus" Huene, 1932
L. indicus Huene and Matley, 1933
= "Laevisuchus indicus" Huene, 1932
Maastrichtian, Late Cretaceous
Lameta Formation, India
Syntypes- (GSI K27/588; lost) mid dorsal vertebra (35 mm)
(GSI K20/613; lost) seventh or eighth cervical vertebra (40 mm)
(GSI K20/614; lost) fifth cervical vertebra
(GSI K27/696) sixth cervical vertebra (42 mm)
Diagnosis- (after Novas et al., 2004) differs from Noasaurus in-
shallower antediapophyseal, diapophyseal and postdiapophyseal fossae in cervical
vertebrae; wider and less ventrally directed cervical diapophyses; cervical
neural spines less excavated anteriorly and posteriorly; shorter cervical prezygopophyses;
postzygapophyses posteriorly rounded in dorsal view. Differs from Masiakasaurus
in- less excavated space between postzygapophyses; thinner prezygapophyses;
shallower infrapostzygapophyseal and infraprezygapophyseal fossae.
Comments- Carrano et al. (2011) identified the vertebral positions based
on comparison to Masiakasaurus.
Wilson (2012) proposed Jubbulpuria may be synonymous with Leavisuchus
as they are of similar size and differ in features attributable to intracolumnar
variation, though they lack overlapping remains.
References- Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre
Entwicklung und Geschichte. Monographien zur Geologie und Palaeontologie. 4(1),
viii + 361 pp.
Huene and Matley, 1933. The Cretaceous Saurischia and Ornithischia of the central
provinces of India. Palaeontologica Indica. 21, 1-74.
Novas and Bandyopaphyay, 1999. New approaches on the Cretaceous theropods from
India. VII International Symposium on Mesozoic Terrestrial Ecosystems, abstracts.
46-47.
Novas, Agnolin and Bandyopadhyay, 2004. Cretaceous theropods from India: A review
of specimens described by Huene and Matley (1933). Revista del Museo Argentino
del Ciencias Naturales. 6(1), 67-103.
Carrano, Loewen and Sertich, 2011. New materials of Masiakasaurus knopfleri
Sampson, Carrano, and Forster, 2001, and implications for the morphology of
the Noasauridae (Theropoda: Ceratosauria). Smithsonian Contributions to Paleobiology.
95, 53 pp.
Wilson, 2012. Small theropod dinosaurs from the Latest Cretaceous of India.
Journal of Vertebrate Paleontology. Program and Abstracts 2012, 194.
Austrocheirus Ezcurra, Agnolin
and Novas, 2010
A. isasii Ezcurra, Agnolin and Novas, 2010
Middle Campanian, Late Cretaceous
Cerro Fortaleza Formation, Santa Cruz, Argentina
Holotype- (MPM-PV 10003) (~7-8 m; young adult) incomplete mid caudal centra,
three partial mid caudal neural arches, incomplete metacarpal III (?), incomplete
phalanx III-1(?), distal tibia, metatarsal II fragment, distal metatarsal III,
three distal pedal phalanges
Diagnosis- (after Ezcurra et al., 2010) metacarpal III with dorsoventrally
tapering shaft towards proximal end; posteriorly displaced collateral tendon
fossae located at same level of the proximal end of distal condyles on metacarpal
III; pedal phalanges with conspicuous longitudinal crest delimitating dorsal
margin of distal collateral tendon fossae.
Comments- Varela (2011) revised the age and nomenclature of the formation
Orkoraptor was found in, which was later done again by Novas et al. (2019). The holotype was discovered in 2002 and described
in 2010 as an abelisauroid outside Abelisauridae by
Ezcurra et al.. Rauhut (2012) later doubted its ceratosaurian identity as no
megaraptorans or coelurosaurs were included in Ezcurra et al.'s analysis, and
several supposedly ceratosaurian characters have a wider distribution within
those clades. Further, he doubted the identification of metacarpal III and suggested
the manual phalanx may be pedal. Rauhut thus placed Austrocheirus as
Theropoda indet., but it is here tentatively retained as a valid abelisauroid
as Cau's (online, 2010) larger unpublished analysis including megaraptorans
and coelurosaurs still recovered it in that clade. Novas et al. (2013)
defended the identity of metacarpal III and the manual phalanx. Most recently,
Tortosa et al. (2014) recovered Austrocheirus as a noasaurid. As metatarsal
II is unreduced in width, it is here placed basal to Velocisaurus, Masiakasaurus
and Noasaurus.
References- Cau, online 2010. http://theropoda.blogspot.com/2010/05/austrocheirus-ezcurra-et-al-2010-una.html
Ezcurra, Agnolin and Novas, 2010. An abelisauroid dinosaur with a non-atrophied
manus from the Late Cretaceous Pari Aike Formation of southern Patagonia. Zootaxa.
2450, 1-25.
Varela, 2011. Sedimentología y modelos deposicionales de la Formación
Mata Amarilla, Cretácico de la cuenca austral, Argentina. PhD thesis,
Universidad Nacional de La Plata. 287 pp.
Rauhut, 2012. A reappraisal of a putative record of abelisauroid theropod dinosaur
from the Middle Jurassic of England. Proceedings of the Geologists' Association.
123(5), 779-786.
Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution of the carnivorous
dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research.
45, 174-215.
Novas, Agnolin,
Rozadilla, Aranciaga-Rolando, Brissón-Eli, Motta, Cerroni, Ezcurra,
Martinelli, D'Angelo, Álvarez-Herrera, Gentil, Bogan, Chimento,
García-Marsà, Lo Coco, Miquel, Brito, Vera, Perez Loinaze, Fernandez
and Salgado, 2019. Paleontological discoveries in the Chorrillo
Formation (upper Campanian-lower Maastrichtian, Upper Cretaceous),
Santa Cruz Province, Patagonia, Argentina. Revista del Museo Argentino
de Ciencias Naturales. 21(2), 217-293.
Velocisaurus Bonaparte, 1991
V. unicus Bonaparte, 1991
Santonian, Late Cretaceous
Bajo de la Carpa Formation of the Rio Colorado Subgroup, Neuquen, Argentina
Holotype- (MUCPv 41) (gracile) tibia (140 mm), astragalus, partial metatarsal
II, phalanx II-1 (23 mm), phalanx II-2 (15 mm), metatarsal III (90 mm), phalanx
III-1 (23 mm), phalanx III-2 (20 mm), phalanx III-3 (16 mm), partial metatarsal
IV, phalanx IV-1 (15 mm), phalanx IV-2 (12 mm), phalanx IV-3 (7 mm), phalanx
IV-4 (5 mm), pedal ungual IV (15 mm)
Santonian, Late Cretaceous
Bajo de la Carpa Formation of the Rio Colorado Subgroup, Rio Negro, Argentina
Referred- (MPCN-PV-370) (robust) incomplete femur (112.1 mm), incomplete
tibia (142.7 mm), fibular fragment, partial metatarsal II, phalanx II-1 (21
mm), incomplete metatarsal III (85.7 mm), phalanx III-1 (16.8 mm), partial metatarsal
IV, phalanx IV-1 (12.4 mm), phalanx IV-2 (12 mm), pedal ungual Iv (11.2 mm)
(Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013; described by Brissón
Egli, Agnolin and Novas, 2016)
Diagnosis- (after Brisson Egli et al., 2016) femur with lateral and medial
surfaces converging anteriorly and resulting in thick dorsoventral crest; tibia
with distal end anteriorly flat, with large surface to accommodate tall and
wide astragalar ascending process; pedal phalanx IV-1 medially curved (also
in GSI K20/626B).
Other diagnoses- Brisson Egli et al. (2016) listed the strongly subtriangular
cross-section of the proximal femur as diagnostic, but noted this is also present
in Masiaksaurus (unknown in other noasaurids). The long and slender tibia
is true of most small theropods. The rod-like metatarsal II and IV shafts are
plesiomorphic compared to Masiakasaurus and Noasaurus. Masiakasaurus
also has a pedal phalanx IV-1 which is transversely compressed, probably narrow
dorsally (described as elliptical), and described as short.
Comments- Brisson Egli et al. (2014) note the side metatarsals are plesiomorphically
rounded, unlike the strap-like elements in Masiakasaurus and Noasaurus,
thus it is placed outside a clade formed by the latter here.
References- Bonaparte, 1991. Los vertebrados fósiles de la Formación
Rio Colorado, de la Ciudad de Neuquén y Cercanías, Cretácico
Superior, Argentina. Revista del Museo Argentino de Ciencias Naturales "Bernardino
Rivadavia" e Instituto Nacional de Investigación de las Ciencias
Naturales: Paleontología. 4(3), 17-123.
Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution of the carnivorous
dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research.
45, 174-215.
Brisson Egli, Agnolin and Novas, 2014. New specimen of Velocisaurus unicus
(Theropoda, Abelisauroidea) from the Paso Cordova locality (Santonian), Rio
Negro. Jornadas Argentinas de Paleontologia de Vertebrados. Ameghiniana. 51(6)
suplemento, 5.
Brissón Egli, Agnolin and Novas, 2016. A new specimen of Velocisaurus
unicus (Theropoda, Abelisauroidea) from the Paso Córdoba locality
(Santonian), Río Negro, Argentina. Journal of Vertebrate Paleontology.
36(4), e1119156.
Vespersaurus Langer, Martins, Manzig, Ferreira, Marsola, Fortes, Lima, Sant'ana, Vidal, Lorençato Ezcurra, 2019
V. paranaensis Langer, Martins, Manzig, Ferreira, Marsola, Fortes, Lima, Sant'ana, Vidal, Lorençato Ezcurra, 2019
Aptian-Albian, Early Cretaceous
Cemitério dos Pterossauros Quarry, Goio Erê Formation, Brazil
Holotype- (MPCO.V 0065d;
proximal ends of metatarsals II-IV lost) (~1-1.5 m) three mid-posterior
dorsal centra, three sacral centra, three mid caudal vertebrae,
incomplete ilium, incomplete ischium, metatarsal I (18 mm), phalanx
I-1, pedal ungual I, distal metatarsal II, phalanx II-1, phalanx II-2,
pedal ungual II, distal metatarsal III, phalanx III-1, phalanx III-2,
phalanx III-3, pedal ungual III, distal metatarsal IV, phalanx IV-1,
phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV
Paratypes- (MPCO.V 0006d) humerus
....(MPCO.V 0006f) scapulocoracoid fragment
....(MPCO.V 0006j) radius
(MPCO.V 0006k) pedal phalanx IV-2, phalanx IV-3
(MPCO.V 0010) mid-posterior dorsal centrum
(MPCO.V 0011) incomplete scapulocoracoid
(MPCO.V 0013) incomplete scapulocoracoid
(MPCO.V 0014) incomplete ischium
(MPCO.V 0016) proximal metatarsal II, distal tarsal III fused with proximal metatarsal III
(MPCO.V 0017) ~eighth-ninth cervical vertebra
(MPCO.V 0018) (~11 kg) incomplete tibia
(MPCO.V 0020) distal caudal vertebra
(MPCO.V 0020c) anterior tooth (5x4x? mm)
(MPCO.V 0022) pedal ungual IV
(MPCO.V 0024) distal caudal vertebra
(MPCO.V 0025) distal caudal vertebra
(MPCO.V 0026) distal caudal vertebra
(MPCO.V 0027) distal caudal vertebra
(MPCO.V 0029) distal caudal vertebra
(MPCO.V 0034a) ~fourth-fifth cervical neural arch
?...(MPCO.V 0034c) ~fourth-fifth cervical centrum
(MPCO.V 0034b) partial posterior cervical neural arch
?...(MPCO.V 0034d) posterior cervical centrum
(MPCO.V 0035) ~third-fourth cervical vertebra
(MPCO.V 0036a) pedal ungual I (9 mm)
(MPCO.V 0040) mid-posterior dorsal centrum
(MPCO.V 0042) incomplete pubis
(MPCO.V 0044) pedal phalanx III-2
(MPCO.V 0048) partial ~fifth-sixth cervical vertebra
(MPCO.V 0049) pedal phalanx III-1
(MPCO.V 0052) distal caudal vertebra
(MPCO.V 0054) pedal phalanx III-1
(MPCO.V 0055) distal metatarsal II
(MPCO.V 0056a) pedal phalanx IV-1
....(MPCO.V 0056b) pedal phalanx IV-2
(MPCO.V 0057a) pedal phalanx IV-2
(MPCO.V 0057b) metatarsal I (13 mm)
(MPCO.V 0059) pedal phalanx III-2
(MPCO.V 0060) pedal phalanx IV-1
(MPCO.V 0061) caudal vertebra
(MPCO.V 0062a) mid-posterior dorsal centrum
(MPCO.V 0062b) mid-posterior dorsal centrum
(MPCO.V 0063a) incomplete metatarsal II
(MPCO.V 0063b) incomplete frontal
(MPCO.V 0064a) fused first to third sacral centra, ilial fragment
(MPCO.V 0064b) manual phalanx I-1, manual ungual I, phalanx II-2, manual ungual II, metacarpal III, phalanx III-1, phalanx III-2
(MPCO.V 0066) partial mid dorsal neural arch
(MPCO.V 0067) axial neural arch
Diagnosis- (after Langer et al., 2019) frontal lacking well-rimed supratemporal fossa; anterior margin of
axial diapophysis not separated from lateral surface of neural arch;
axial epipophysis projecting distinctly beyond postzygapophyseal facet;
no cranial or caudal peduncular fossae in anterior post-axial cervical
vertebrae; mid cervical vertebrae with prezygoepipophyseal lamina not
anteriorly extensive (also in Masiaksaurus); mid cervical epipophysis
shorter than length of postzygapophyseal facet (also in Masiakasaurus);
scapular blade not narrowing at its distal two-thirds; coracoid with
drop-shaped groove on tuber; nearly straight posterior coracoid margin;
humeral head more than twice lateromedially broader than
anteroposteriorly deep in proximal view; humerus with convex distal
condyles; ischiadic peduncle of ilium not significantly projected
ventrally relative to pubic peduncle; pubis with ambiens tuber on
dorsal surface; medial surface of pubis with semilunate depression
posterior to obturator foramen; ischium with no obturator notch;
metatarsal II with extremely transversely compressed shaft; distal
articulation of metatarsal II with shallower ventral sulcus and
lateromedialy broader medial flange; lateromedially compressed
metatarsal IV (also in Velocisaurus); pedal ungual IV with proximal end dorsoventrally taller than distal end of phalanx IV-1.
Comments- Langer et al. state
the assignment of the holotype to one individual "to the same
individual is tentative and based on the absence of duplicated
elements, their close association, and matching phylogenetic
signal." Similarly, the referred elements "are tentatively
assigned to Ves. paranaensis based on their matching size and phylogenetic signal." Vespersaurus was discovered in October 2012 (Kellner et al., 2019) in a bonebed intermixed with two pterosaurs, the tapejarid Caiaujara dobruski and basal tapejaromorph Keresdrakon vilsoni. Langer et al. (2019) added Vespersaurus to Rauhut and Carrano's ceratosaur matrix and recovered it sister to Velocisaurus in Noasauridae.
References- Kellner,
Weinschutz, Holgado, Bantim and Sayao, 2019. 2019. A new toothless
pterosaur (Pterodactyloidea) from southern Brazil with insights into
the paleoecology of a Cretaceous desert. Anais da Academia Brasileira
de Ciências. 91(Suppl. 2), e20190768.
Langer, Martins, Manzig, Ferreira, Marsola, Fortes, Lima, Sant'ana,
Vidal, Lorençato Ezcurra, 2019. A new desert-dwelling dinosaur
(Theropoda, Noasaurinae) from the Cretaceous of south Brazil.
Scientific Reports. 9:9379.
Masiakasaurus Sampson, Carrano
and Forster, 2001
M. knopfleri Sampson, Carrano and Forster, 2001
Middle Maastrichtian, Late Cretaceous
Anembalemba Member of Maevarano Formation, Madagascar
Holotype- (UA 8680) incomplete dentary
Paratypes- (FMNH PR 2108) pubis
(FMNH PR 2109) pubis
(FMNH PR 2110) mid caudal vertebra
(FMNH PR 2111) ?dorsal centrum
(FMNH PR 2112) (gracile) tibia (191.3 mm)
(FMNH PR 2113) dorsal centrum
(FMNH PR 2114) dorsal centrum
(FMNH PR 2115) (gracile) femur
(FMNH PR 2116) (robust) tibia (189.2 mm), partial fibula, astragalocalcaneum
(FMNH PR 2117) (robust) femur (189.4 mm)
(FMNH PR 2118) (gracile) tibia
(FMNH PR 2119) (robust) tibia
(FMNH PR 2120) (gracile) femur (190.8 mm)
(FMNH PR 2121) (gracile) tibia (171.5 mm)
(FMNH PR 2122) (robust) tibia, partial fibula, astragalocalcaneum
(FMNH PR 2123) (robust) femur (202.5 mm)
(FMNH PR 2124; mistakenly labeled 2183 by Carrano et al., 2002 in fig. 4) splenial
(FMNH PR 2125) mid caudal
(FMNH PR 2126) mid caudal vertebra
(FMNH PR 2127) distal caudal vertebra
(FMNH PR 2128) distal caudal vertebra
(FMNH PR 2129) pedal phalanx II-1
(FMNH PR 2130) pedal phalanx IV-3
(FMNH PR 2131) pedal phalanx IV-3
(FMNH PR 2132) manual phalanx I-1(?)
(FMNH PR 2133) proximal caudal vertebra
(FMNH PR 2134) pedal ungual
(FMNH PR 2135) pedal ungual II/IV
(FMNH PR 2136) manual ungual, pedal phalanx II-2
(FMNH PR 2137) dorsal centrum
(FMNH PR 2138) dorsal centrum
(FMNH PR 2139) seventh cervical vertebra
(FMNH PR 2140) tenth cervical vertebra
(FMNH PR 2141) eighth cervical vertebra
(FMNH PR 2142) third-fifth sacral vertebrae
(FMNH PR 2143) incomplete humerus
(FMNH PR 2144) fourth dorsal neural arch
....(FMNH PR 2145) dorsal centrum
(FMNH PR 2146; mistakenly labeled 2147 by Carrano et al. (2002) in fig. 19)
metatarsal III
(FMNH PR 2147) metatarsal II
(FMNH PR 2148) (gracile) femur
(FMNH PR 2149) (robust) femur
(FMNH PR 2150) (gracile) femur
(FMNH PR 2151) metatarsal II
(FMNH PR 2152) (gracile) distal tibia
(FMNH PR 2153) (gracile) femur
(FMNH PR 2154) metatarsal II
(FMNH PR 2155) metatarsal III
(FMNH PR 2156) distal caudal vertebra
(FMNH PR 2157) distal caudal vertebra
(FMNH PR 2158) pedal phalanx IV-1
(FMNH PR 2159) pedal phalanx III-1
(FMNH PR 2160) pedal phalanx II-1
(FMNH PR 2161) pedal phalanx II-1
(FMNH PR 2162) distal caudal vertebra
(FMNH PR 2163) distal caudal vertebra
(FMNH PR 2164) lateral tooth
(FMNH PR 2165) anterior tooth
(FMNH PR 2167) pedal phalanx III-1, phalanx III-2
(FMNH PR 2168) distal caudal vertebra
(FMNH PR 2169) manual ungual
(FMNH PR 2170) lateral tooth
(FMNH PR 2171) dorsal centrum
(FMNH PR 2172) pedal phalanx IV-1
(FMNH PR 2173) pedal phalanx III-2
(FMNH PR 2174) pedal phalanx IV-3
(FMNH PR 2175) distal metatarsal II
(FMNH PR 2176) pedal phalanx III-2
(FMNH PR 2177) dentary
(FMNH PR 2178) dentary
(FMNH PR 2179) dentary
(FMNH PR 2180) anterior tooth
(FMNH PR 2181) two lateral teeth
(FMNH PR 2182) lateral tooth
(UA 8681) (gracile) femur (202.5 mm)
(UA 8682) pedal phalanx (mistakenly listed as a dentary by Carrano et al., 2002-
Caranno et al., 2011)
(UA 8683) distal metatarsal II
(UA 8684) (robust) femur (201.6 mm)
(UA 8685) (robust) tibia (205.4 mm)
(UA 8686) pedal phalanx IV-4
(UA 8687) (gracile) two vertebrae, tibia
(UA 8688) mid caudal vertebra
(UA 8689) distal caudal vertebra
(UA 8690) distal caudal vertebra
(UA 8691) distal caudal vertebra
(UA 8692) mid caudal vertebra
(UA 8693) humeral shaft
(UA 8694) humeral shaft
(UA 8695) distal caudal vertebra
(UA 8696) distal caudal vertebra
Referred- (FMNH PR 2100) lateral tooth (Carrano et al., 2002)
(FMNH PR 2101) lateral tooth (Carrano et al., 2002)
(FMNH PR 2183) maxilla (Carrano et al., 2002)
(FMNH PR 2199) lateral tooth (Carrano et al., 2002)
(FMNH PR 2200) anterior tooth (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2201) latreral tooth (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2202) distal caudal vertebra (Carrano et al., 2002)
(FMNH PR 2203) distal caudal vertebra (Carrano et al., 2002)
(FMNH PR 2205) pedal phalanx (Carrano et al., 2002)
(FMNH PR 2206) metatarsal II (Carrano et al., 2002)
(FMNH PR 2207) dorsal centrum (Carrano et al., 2002)
(FMNH PR 2208) (robust) femur (Carrano et al., 2002)
(FMNH PR 2214) (gracile) tibia (151.2 mm), metatarsal IV (Carrano et al., 2002)
(FMNH PR 2215) (gracile) femur (180 mm) (Carrano et al., 2002)
(FMNH PR 2216) pedal phalanx IV-2 (Carrano et al., 2002)
(FMNH PR 2217) pedal phalanx II-2 (Carrano et al., 2002)
(FMNH PR 2218) pedal phalanx III-1 (Carrano et al., 2002)
(FMNH PR 2219) pedal phalanx III-2/3 (Carrano et al., 2002)
(FMNH PR 2220) anterior tooth (Carrano et al., 2002)
(FMNH PR 2221) lateral tooth (Carrano et al., 2002)
(FMNH PR 2222) dentary (Carrano et al., 2002)
(FMNH PR 2223) two pedal phalanges (Carrano et al., 2002)
(FMNH PR 2224) manual phalanx I-1(?) (Carrano et al., 2002)
(FMNH PR 2225) two manual phalanges (Carrano et al., 2002)
(FMNH PR 2227) manual phalanx (Carrano et al., 2002)
(FMNH PR 2229) dorsal centrum (Carrano et al., 2002)
(FMNH PR 2230) proximal caudal centrum (Carrano et al., 2002)
(FMNH PR 2234) proximal metatarsal IV (Carrano et al., 2002)
(FMNH PR 2235) calcaneum (Carrano et al., 2002)
(FMNH PR 2236) pedal ungual (Carrano et al., 2002)
(FMNH PR 2453) partial premaxilla (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2454) prearticular (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2455) incomplete angular (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2456) postorbital (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2458) dorsal neural arch, partial fourth dorsal rib, mid caudal vertebra,
distal caudal vertebra, scapula (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2459) metatarsal IV (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2460) fused third to sixth sacral centra (Carrano, Loewen and Sertich,
2011)
(FMNH PR 2461) proximal pubis (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2462) axis (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2463) incomplete pubis (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2464) cervical vertebra (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2465) fifth cervical vertebra (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2466) axis (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2467) distal caudal vertebra (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2468) partial ischium (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2469) proximal caudal vertebra (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2470) fused pubes (one incomplete, one distal) (Carrano, Loewen and
Sertich, 2011)
(FMNH PR 2471) dentary (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2472) ilium (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2473) partial lacrimal (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2475) frontal (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2476) lateral tooth (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2477) atlantal intercentrum (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2478) rib (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2479) pedal phalanx IV-2/3 (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2480) pedal phalanx III-2 (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2481) hyoid, atlantal intercentrum (5 mm), ~fifth cervical neural arch,
sixth cervical vertebra (32.7 mm), seventh cervical vertebra (29.4 mm), ninth
cervical vertebra (26.5 mm), proximal fourth cervical rib, proximal tenth cervical
rib, two proximal cervical ribs, fifth dorsal vertebra (26.9 mm), sixth dorsal
vertebra (28 mm), seventh dorsal vertebra (28.7 mm), ~eleventh dorsal vertebra
(29 mm), ~twelfth dorsal vertebra (29.6 mm), ~thirteenth dorsal vertebra (29.3
mm), ~fourteenth dorsal vertebra (28.9 mm), three partial dorsal ribs, three
rib fragments, fused anterior gastralia, sacrum (148.8 mm), proximal caudal
vertebra (29.3 mm), proximal caudal vertebra (29.1 mm), proximal caudal vertebra
(29 mm), mid caudal vertebra (28.9 mm), proximal caudal neural arch, mid caudal
vertebra (29.8 mm), distal caudal vertebra (30 mm), distal caudal vertebra (31.2
mm), distal caudal vertebra (31.1 mm), distal caudal vertebra, distal caudal
vertebra (31 mm), distal caudal vertebra (29.2 mm), three caudals, chevron,
scapulae (128.4, 128.2 mm), coracoids (one fragmentary; 38 mm), humerus (94.3
mm), ilia (187.2 mm), pubes (206 mm), femora (one distal; ~193.6 mm), tibiae
(195.8, 196.9 mm), fibulae (one incomplete; 189.6 mm), astragali, distal tarsal
III, metatarsals II (one distal; 93.2 mm), metatarsal III (112 mm), phalanx
III-1 (28.2 mm), phalanx III-2/3 (18.2 mm) (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2482) mid caudal vertebra (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2485) fourth cervical vertebra (24.7 mm), ~sixth cervical vertebra
(25.4 mm), ~seventh cervical vertebra (23.4 mm), fourth cervical rib, sixth
cervical rib, proximal seventh cervical rib, second dorsal vertebra, fourth
dorsal neural arch, two dorsal neural arches, ~fourth dorsal rib, dorsal rib,
five rib fragments, first sacral centrum (22.2 mm), proximal caudal vertebra
(28.3 mm), mid caudal vertebra (23.2 mm), distal caudal vertebra (25.4 mm),
distal caudal vertebra (27.3 mm), distal caudal vertebra (25.5 mm), two caudals,
vertebra, chevron, humerus (80.8 mm), incomplete ilium (146.3 mm), incomplete
ischia, femora (160.7, 159.8 mm), tibiae (173.7, 171.7 mm), proximal fibulae,
metatarsal III (96.4 mm), distal metatarsal IV, pedal phalanx (14.8 mm), pedal
ungual II/IV (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2486) pedal ungual II/IV (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2490) caudal centrum (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2491) tenth cervical vertebra (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2492) three distal caudal vertebrae (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2493) pedal phalanx (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2495) ?laterosphenoid (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2496) partial quadrate (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2603) rib fragments (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2604) pedal phalanx II-2 (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2605) pedal phalanx IV-1 (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2606) scapula (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2610) dorsal centrum (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2611) pedal phalanx (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2613) pedal phalanx III-? (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2614) pedal phalanx IV-1 (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2616) pedal phalanx II-1 (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2617) pedal phalanx III-? (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2618) neural arch (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2628) sixth cervical verterbra (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2630) antlantal intercentrum (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2632) rib fragment (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2634) proximal rib (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2636) fourth dorsal vertebra (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2638) distal caudal vertebra (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2642) distal caudal vertebra (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2667) pedal ungual II/IV (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2672) tenth cervical or first dorsal rib (Carrano, Loewen and Sertich,
2011)
(FMNH PR 2673) chevron (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2677) centrum (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2679) distal metatarsal II (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2684) third dorsal vertebra (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2685) rib fragment (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2686) tibia (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2687) distal tarsal III, proximal metatarsal III (Carrano, Loewen and
Sertich, 2011)
(FMNH PR 2696) lateral tooth (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2699) caudal (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2700) caudal (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2709) pedal phalanx II-1 (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2817) tibia (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2818) anterior tooth (Carrano, Loewen and Sertich, 2011)
(FMNH PR 2837) first dorsal vertebra (Carrano, Loewen and Sertich, 2011)
(MNHN.MAJ 249) tooth (Smith, 2007)
(MSNM V5378) fourth or fifth dentary tooth (13 mm) (Fanti and Therrien, 2007)
(UA 8700) pedal phalanx III-1 (Carrano et al., 2002)
(UA 8701) dorsal centrum (Carrano et al., 2002)
(UA 8702) distal caudal (Carrano et al., 2002)
(UA 8703) distal caudal vertebra (Carrano et al., 2002)
(UA 8710) (gracile) proximal tibia (Carrano et al., 2002)
(UA 8711) (gracile) tibia (167.6 mm) (Carrano et al., 2002)
(UA 8712) (robust) femur (168.3 mm) (Carrano et al., 2002)
(UA 8713) pedal phalanx (Carrano et al., 2002)
(UA 8714) pedal phalanx (Carrano et al., 2002)
(UA 9090) postorbital (Carrano, Loewen and Sertich, 2011)
(UA 9091) lateral tooth (Carrano, Loewen and Sertich, 2011)
(UA 9092) posterior dorsal vertebra (Carrano, Loewen and Sertich, 2011)
(UA 9093) centrum (Carrano, Loewen and Sertich, 2011)
(UA 9094) pedal ungual III (Carrano, Loewen and Sertich, 2011)
(UA 9095) frontal (Carrano, Loewen and Sertich, 2011)
(UA 9096) pedal phalanx IV-2 (Carrano, Loewen and Sertich, 2011)
(UA 9097) posterior dorsal vertebra (Carrano, Loewen and Sertich, 2011)
(UA 9098) partial fifth sacral vertebra, sixth sacral vertebra, incomplete fibula
(Carrano, Loewen and Sertich, 2011)
(UA 9099) proximal tibia (Carrano, Loewen and Sertich, 2011)
(UA 9100) incomplete pubis (Carrano, Loewen and Sertich, 2011)
(UA 9101) metatarsal IV (Carrano, Loewen and Sertich, 2011)
(UA 9102) distal tarsal III fused to metatarsal III (Carrano, Loewen and Sertich,
2011)
(UA 9103) posterior dorsal vertebra (Carrano, Loewen and Sertich, 2011)
(UA 9104) proximal ninth cervical rib (Carrano, Loewen and Sertich, 2011)
(UA 9105) rib fragments, metatarsal (Carrano, Loewen and Sertich, 2011)
(UA 9106) fourth cervical vertebra (Carrano, Loewen and Sertich, 2011)
(UA 9107) second dorsal neural arch (Carrano, Loewen and Sertich, 2011)
(UA 9108) dorsal transverse process (Carrano, Loewen and Sertich, 2011)
(UA 9109) neural arch (Carrano, Loewen and Sertich, 2011)
(UA 9110) caudal centrum (Carrano, Loewen and Sertich, 2011)
(UA 9111) seventh cervical vertebra (Carrano, Loewen and Sertich, 2011)
(UA 9112) proximal caudal vertebra (Carrano, Loewen and Sertich, 2011)
(UA 9113) caudal centrum (Carrano, Loewen and Sertich, 2011)
(UA 9115) first sacral vertebra (Carrano, Loewen and Sertich, 2011)
(UA 9116) scapula (Carrano, Loewen and Sertich, 2011)
(UA 9117) pubic shaft (Carrano, Loewen and Sertich, 2011)
(UA 9118) distal metatarsal ?I (Carrano, Loewen and Sertich, 2011)
(UA 9119) distal caudal vertebra (Carrano, Loewen and Sertich, 2011)
(UA 9120) mid caudal vertebra (Carrano, Loewen and Sertich, 2011)
(UA 9121) third cervical vertebra (Carrano, Loewen and Sertich, 2011)
(UA 9122) metatarsal II (Carrano, Loewen and Sertich, 2011)
(UA 9123) lateral tooth (Carrano, Loewen and Sertich, 2011)
(UA 9126) lateral tooth (Carrano, Loewen and Sertich, 2011)
(UA 9127) caudal (Carrano, Loewen and Sertich, 2011)
(UA 9128) lateral tooth (Carrano, Loewen and Sertich, 2011)
(UA 9130) axial neural arch (Carrano, Loewen and Sertich, 2011)
(UA 9132) partial fibula, pedal phalanx (Carrano, Loewen and Sertich, 2011)
(UA 9133) proximal ischium (Carrano, Loewen and Sertich, 2011)
(UA 9134) proximal fibula (Carrano, Loewen and Sertich, 2011)
(UA 9135) distal femur (Carrano, Loewen and Sertich, 2011)
(UA 9136) pubic shaft (Carrano, Loewen and Sertich, 2011)
(UA 9137) pedal phalanx IV-2 (Carrano, Loewen and Sertich, 2011)
(UA 9138) distal metatarsal II (Carrano, Loewen and Sertich, 2011)
(UA 9139) distal caudal vertebra (Carrano, Loewen and Sertich, 2011)
(UA 9140) caudal (Carrano, Loewen and Sertich, 2011)
(UA 9142) distal tibia (Carrano, Loewen and Sertich, 2011)
(UA 9143) partial fibula (Carrano, Loewen and Sertich, 2011)
(UA 9144) pedal ungual (Carrano, Loewen and Sertich, 2011)
(UA 9145) dentary (Carrano, Loewen and Sertich, 2011)
(UA 9146) proximal metacarpal ?IV (Carrano, Loewen and Sertich, 2011)
(UA 9147) incomplete angular (Carrano, Loewen and Sertich, 2011)
(UA 9148) caudal (Carrano, Loewen and Sertich, 2011)
(UA 9149) prearticular (Carrano, Loewen and Sertich, 2011)
(UA 9150) ninth cervical vertebra, tenth cervical vertebra (Carrano, Loewen
and Sertich, 2011)
(UA 9151) caudal centrum (Carrano, Loewen and Sertich, 2011)
(UA 9152) mid chevron (Carrano, Loewen and Sertich, 2011)
(UA 9153) proximal pubis (Carrano, Loewen and Sertich, 2011)
(UA 9154) pedal phalanx II-1 (Carrano, Loewen and Sertich, 2011)
(UA 9155) proximal dorsal rib (Carrano, Loewen and Sertich, 2011)
(UA 9156) pedal phalanx IV-1 (Carrano, Loewen and Sertich, 2011)
(UA 9157) pedal ungual III (Carrano, Loewen and Sertich, 2011)
(UA 9158) mid caudal vertebra (Carrano, Loewen and Sertich, 2011)
(UA 9159) lateral tooth, coracoid (Carrano, Loewen and Sertich, 2011)
(UA 9160) proximal ~fourth dorsal rib, incomplete scapulocoracoid (Carrano,
Loewen and Sertich, 2011)
(UA 9162) proximal pubis (Carrano, Loewen and Sertich, 2011)
(UA 9163) distal metatarsal IV (Carrano, Loewen and Sertich, 2011)
(UA 9164) dorsal neural arch (Carrano, Loewen and Sertich, 2011)
(UA 9165) proximal humerus (Carrano, Loewen and Sertich, 2011)
(UA 9166) prearticular, articular (Carrano, Loewen and Sertich, 2011)
(UA 9167) fibula (Carrano, Loewen and Sertich, 2011)
(UA 9168) ischium (Carrano, Loewen and Sertich, 2011)
(UA 9169) proximal sixth cervical rib (Carrano, Loewen and Sertich, 2011)
(UA 9170) partial ilium, femur (Carrano, Loewen and Sertich, 2011)
(UA 9171) eighth cervical vertebra (Carrano, Loewen and Sertich, 2011)
(UA 9172) ischium, incomplete fibula (Carrano, Loewen and Sertich, 2011)
(UA 9173) proximal caudal centrum (Carrano, Loewen and Sertich, 2011)
(UA 9174) caudal neural arch (Carrano, Loewen and Sertich, 2011)
(UA 9175) posterior dorsal vertebra (Carrano, Loewen and Sertich, 2011)
(UA 9176) posterior dorsal vertebra (Carrano, Loewen and Sertich, 2011)
(UA 9177) dentary (Carrano, Loewen and Sertich, 2011)
(UA 9178) basioccipital (Carrano, Loewen and Sertich, 2011)
(UA 9179) mid caudal vertebra (Carrano, Loewen and Sertich, 2011)
(UA 9180) chevron (Carrano, Loewen and Sertich, 2011)
(UA 9181) eighth cervical vertebra (Carrano, Loewen and Sertich, 2011)
(UA 9182) chevron (Carrano, Loewen and Sertich, 2011)
(UA 9183) lateral tooth (Carrano, Loewen and Sertich, 2011)
(UA 9185) pedal phalanx IV-1 (Carrano, Loewen and Sertich, 2011)
(UA 9186) pedal phalanx III-? (Carrano, Loewen and Sertich, 2011)
(UA 9187) caudal (Carrano, Loewen and Sertich, 2011)
(UA 9188) pedal phalanx II-1 (Carrano, Loewen and Sertich, 2011)
(UA 9189) fourth cervical vertebra (Carrano, Loewen and Sertich, 2011)
(UA 9190) mid caudal vertebra (Carrano, Loewen and Sertich, 2011)
(UA 9192) pedal phalanx III-? (Carrano, Loewen and Sertich, 2011)
(UA 9193) femur (Carrano, Loewen and Sertich, 2011)
(UA 9194) manual phalanx (Carrano, Loewen and Sertich, 2011)
(UA 9195) mid caudal vertebra (Carrano, Loewen and Sertich, 2011)
(UA 9196) chevron (Carrano, Loewen and Sertich, 2011)
(UA 9197) mid caudal vertebra (Carrano, Loewen and Sertich, 2011)
(UA 9198) pedal ungual (Carrano, Loewen and Sertich, 2011)
(UA 9199) pedal phalanx III-2/3 (Carrano, Loewen and Sertich, 2011)
(UA 9613) distal tibia (Carrano, Loewen and Sertich, 2011)
(UA 9734) lateral tooth (Carrano, Loewen and Sertich, 2011)
(UA 9773) distal metatarsal (Carrano, Loewen and Sertich, 2011)
(UA 9774) distal metatarsal II (Carrano, Loewen and Sertich, 2011)
(UA 9857) distal caudal vertebra (Carrano, Loewen and Sertich, 2011)
(UA 9858) mid caudal vertebra (Carrano, Loewen and Sertich, 2011)
(UA 9859) cervical centrum (Carrano, Loewen and Sertich, 2011)
(UA 9861) proximal rib (Carrano, Loewen and Sertich, 2011)
(UA 9862) proximal ~fourth dorsal rib (Carrano, Loewen and Sertich, 2011)
(UA 9863) proximal rib (Carrano, Loewen and Sertich, 2011)
Early Maastrichtian(?), Late Cretaceous
Masorobe Member of Maevarano Formation, Madagascar
(FMNH PR 2457) (subadult) posterior braincase (Carrano, Loewen and Sertich,
2011)
(UA 9184) caudal centrum (Carrano, Loewen and Sertich, 2011)
Diagnosis- (after Carrano et al., 2002) anterior four dentary teeth procumbent,
the first inclined at 108 above the horizontal and lying in an alveolus that
is slung below the ventral margin of the dentary; first alveolus large and ventrally
expanded, lying lateral to an anteroposteriorly long dentary symphysis; lower
dentition markedly heterodont: the first four teeth are weakly spoon-shaped,
elongate, and terminate in a posteriorly hooked, pointed apex; anterior dentary
teeth bear two weakly serrated posterior carinae and have faint posterior ridges;
more posterior teeth are transversely compressed, recurved, and have a serrated
posterior carina.
Comments- Carrano et al. (2011) reidentified a number of elements from
Sampson et al. (2001) and/or Carrano et al. (2002). The possible angular (FMNH
PR 2166) is an unknown element from an unknown taxon. FMNH PR 2198, 2226 and
2228 are reidentified as juvenile Majungasaurus teeth, contra Carrano
et al. (2002). FMNH PR 2204 is a crocodyliform proximal caudal vertebra. FMNH
PR 2136 and 2217 are manual phalanges instead of pedal phalanges.
References- Sampson, Nyit, Forster, Suny, Krause and Suny, 1998. The
Late Cretaceous dinosaurs of Madagascar. Journal of Vertebrate Paleontology.
18(3), 74A.
Sampson, Carrano, and Forster. 2000. A theropod dinosaur with bizarre dentition
from the Late Cretaceous of Madagascar. Journal of Vertebrate Paleontology.
20(3), 66A.
Sampson, Carrano and Forster, 2001. A bizarre predatory dinosaur from the Late
Cretaceous of Madagascar. Nature. 409, 504-506.
Carrano, Sampson and Forster, 2002. The osteology of Masiakasaurus knopfleri,
a small abelisauroid (Dinosauria: Theropoda) from the Late Cretaceous of Madagascar.
Journal of Vertebrate Paleontology. 22(3), 510-534.
Carrano, Sampson and Loewen, 2004. New discoveries of Masiakasaurus knopfleri
and the morphology of the Noasauridae (Dinosauria: Theropoda). Journal of Vertebrate
Paleontology. 24(3), 27A-28A.
Fanti and Therrien, 2007. Theropod tooth assemblages from the Late Cretaceous
Maevarano Formation and the possible presence of dromaeosaurids in Madagascar.
Acta Palaeontologica Polonica. 52(1), 155-166.
Smith, 2007. Dental morphology and variation in Majungasaurus crenatissimus
(Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. In Sampson
and Krause (eds.). Majungasaurus crenatissimus (Theropoda: Abelisauridae)
from the Late Cretaceous of Madagascar. SVP Memoir 8. 103-126.
Carrano, Loewen and Sertich, 2011. New materials of Masiakasaurus knopfleri
Sampson, Carrano, and Forster, 2001, and implications for the morphology of
the Noasauridae (Theropoda: Ceratosauria). Smithsonian Contributions to Paleobiology.
95, 53 pp.
Lee and O'Connor, 2011. Variation in bone histology and growth of the noasaurid
theropod Masiakasaurus knopfleri. Journal of Vertebrate Paleontology.
Program and Abstracts 2011, 142.
Noasaurus Bonaparte and Powell,
1980
N. leali Bonaparte and Powell, 1980
Late Campanian-Maastrichtian, Late Cretaceous
Lecho Formation, Salta, Argentina
Holotype- (PVL 4061) maxilla (78 mm), quadrate (45 mm), fourth cervical
neural arch, fourth cervical rib (48 mm), tenth cervical or first dorsal rib,
dorsal centrum, manual phalanx III-? (17 mm), manual ungual ?II (37 mm), metatarsal
II (113 mm)
....(MACN 622) (adult) anterior cervical vertebra (34.4 mm) (Frankfurt and Chiappe,
1999)
Referred- manual ungual ?II (Agnolin, Apesteguia and Chiarelli, 2004)
Diagnosis- (after Bonaparte and Powell, 1980) very deep and subtriangular
ventral concavity on manual ungual ?II[pedal ungual II in original].
(After Agnolin et al., 2004) both sides of manual ungual ?II sub-parallel in
dorsal view.
(after Angnolin and Chiarelli, 2009) median ventral ridge distal to proximoventral
concavity on manual ungual ?II.
(after Caranno et al., 2011) compared to Masiakasaurus- narrower antorbital
fossa; anterior margin of antorbital fenestra reaches to fifth alveolus; quadrate
shaft more strongly curved; medial collateral ligament pit on metatarsal IV
internally subdivided; more distinct distal intercondylar sulcus on metatarsal
IV.
Comments- The squamosal originally identified is a cervical rib (Bonaparte,
1991), further identified as the tenth (or the first dorsal) rib by Carrano
et al. (2011). Carrano et al. also identified the cervical neural arch and rib
as the fourth based on comparison to Masiakasaurus. A cervical vertebra
(MACN 622) discovered with the holotype was originally identified as an oviraptorosaur
(Frankfurt and Chiappe, 1999), but reidentified by Agnolin and Martinelli (2007)
as a noasaurid, and probably part of the Noasaurus holotype individual.
Noasaurus was originally described by Bonaparte and Powell (1980) as
preserving a hyperextendable pedal phalanx II-2 and enlarged and trenchant pedal
ungual II as in deinonychosaurs. However, Agnolin et al. (2004) and Agnolin
and Chiarelli (2009) found a series of manual characters and abelisauroid synapomorphies
in the phalanx and ungual. Similarly, Carrano et al. (2004; 2011) compared the
phalanx favorably to manual phalanges FMNH PR 2136 and 2217 of Masiakasaurus,
which has very different unguals on its pes. Agnolin et al. proposed the ungual
was from digit II based on the symmetry of the proximal articular facets, but
Agnolin and Chiarelli suggested it was from I or II based on its "strong
curvature and general morphology", though their figure caption lists it
as from II or III. Carrano and Sampson (2008) suggested Bonaparte and Powell
had the phalanx upside down, but Agnolin and Chiarelli noted several characters
indicating its manual identification and an abelisauroid synapomorphy that work
using the original orientation, so that is preferred here. Carrano et al. (2011)
suggested the proportions of the phalanx might indicate it was penultimate,
but Agnolin and Chiarelli noted the short and broad proportions were like digit
III and that contra the original description, it does not articulate well with
the ungual.
References- Bonaparte and Powell, 1980. A continental assemblage of tetrapods
from the upper Cretaceous beds of El Brete, northwestern Argentina (Sauropoda-Coelurosauria-Carnosauria-Aves).
Mémoires de la Société Géologique de France. Nouvelle
Série. 19, 19-28.
Bonaparte, 1991. The Gondwanian theropod families Abelisauridae and Noasauridae.
Historical Biology. 5, 1-25.
Frankfurt and Chiappe, 1999. A possible oviraptorosaur from the Late Cretaceous
of northwestern Argentina. Journal of Vertebrate Paleontology. 19(1), 101-105.
Agnolin, Apesteguia and Chiarelli, 2004. The end of a myth: The mysterious ungual
claw of Noasaurus leali. Journal of Vertebrate Paleontology. 24(3), 301A-302A.
Carrano, Sampson and Loewen, 2004. New discoveries of Masiakasaurus knopfleri
and the morphology of the Noasauridae (Dinosauria: Theropoda). Journal of Vertebrate
Paleontology. Program and Abstracts 2004. 28.
Agnolin and Martinelli, 2007. Did oviraptorosaurs (Dinosauria; Theropoda) inhabit
Argentina? Cretaceous Research. 28(5), 785-790.
Carrano and Sampson, 2008 (online 2007). The Phylogeny of Ceratosauria (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 6, 183-236.
Agnolin and Chiarelli, 2010. The position of the claws in Noasauridae (Dinosauria:
Abelisauroidea) and its implications for abelisauroid manus evolution. Paläontologische
Zeitschrift. 84, 293-300.
Carrano, Loewen and Sertich, 2011. New materials of Masiakasaurus knopfleri
Sampson, Carrano, and Forster, 2001, and implications for the morphology of
the Noasauridae (Theropoda: Ceratosauria). Smithsonian Contributions to Paleobiology.
95, 53 pp.
Abelisauridae Bonaparte and Novas, 1985
Definition- (Abelisaurus comahuensis <- Noasaurus leali, Ceratosaurus nasicornis, Elaphrosaurus bambergi, Allosaurus fragilis) (proposed)
Other definitions- (Abelisaurus comahuensis + Carnotaurus sastrei
+ Indosaurus matleyi + Indosuchus raptorius + Majungasaurus
crenatissimus + Xenotarsosaurus bonapartei) (Novas, 1997)
(Carnotaurus sastrei <- Elaphrosaurus
bambergi) (modified from Rowe, Tykoski and Hutchinson, 1997)
(Abelisaurus comahuensis + Carnotaurus sastrei) (Tykoski and Rowe,
2004; modified from Sereno, 1998)
(Carnotaurus sastrei <- Noasaurus leali) (Wilson, Sereno, Srivastava, Bhatt, Khosla and Sahni, 2003)
(Carnotaurus sastrei <- Coelophysis bauri, Noasaurus leali,
Passer domesticus) (Sereno, online 2005)
(Carnotaurus sastrei <- Ceratosaurus nasicornis) (Delcourt, 2018)
(Carnotaurus sastrei <- Noasaurus leali, Ceratosaurus nasicornis) (Hendrickx,
Mateus, Araújo and Choiniere, 2019)
= Abelisauridae sensu Wilson, Sereno, Srivastava, Bhatt, Khosla and Sahni, 2003
Definition- (Carnotaurus sastrei <- Noasaurus leali)
= Abelisauridae sensu Sereno, online 2005
Definition- (Carnotaurus sastrei <- Coelophysis bauri, Noasaurus
leali, Passer domesticus)
= Abelisauridae sensu Hendrickx,
Mateus, Araújo and Choiniere, 2019
Definition- (Carnotaurus sastrei <- Noasaurus leali, Ceratosaurus nasicornis)
Diagnosis- (after Carrano et al., 2002) long axis of postorbital slanted
anteroventrally/posterodorsally; frontals fused to parietals; quadratojugal
fused to quadrate; interdental plates striated medially.
Comments- Unfortunately, Abelisauridae lacks a potentially
valid published definition as currently conceived, as all stem-based
definitions use Carnotaurus as the internal specifier when Abelisaurus comahuensis must be one (Phylocode Article 11.10- "when a clade name is converted from a preexisting name that is typified
under a rank-based code or is a new or converted name derived from the
stem of a typified name, the definition of the clade name must use the
type species of that preexisting typified name or of the genus name from
which it is derived [or the type specimen of that species] as an
internal specifier"). If Abelisaurus
is a carcharodontosaurid (a possibility suggested by Lamanna et al., 2002)
it would leave Abelisauridae without Abelisaurus
which would not be accepted anyway, and the family would be renamed
Carnotauridae as that is the next oldest family level name associated
with the clade. Of the two published definitions which do include
Abelisaurus, Novas' (1997) includes the fragmentary Xenotarsosaurus whose precise relationships with the brachyrostran-majungasaurine clade and Rugops
cannot be resolved even after the most recent redescription (Ibiricu et
al., 2021), leaving the family with uncertain basal content. On
the other hand, Sereno's (1998) definition would limit it to
Carnotauruni using this site's phylogeny, and introduce an identical
problem to Xenotarsosaurus using Ibiricu et al.'s topology where Abelisaurus is outside the brachyrostran-majungasaurine clade instead of sister to Aucasaurus. Thus a stem-based definition of Abelisauridae is proposed here including Abelisaurus which still allows basal taxa like Rugops and fragmentary remains to be referred to the family.
References- Bonaparte and Novas, 1985. Abelisaurus comahuensis,
n. g., n. sp., Carnosauria from the Late Cretaceous of Patagonia. Ameghiniana.
21, 259-265.
Novas, 1997. Abelisauridae. In Currie and Padian (eds.). Encyclopedia of Dinosaurs. Academic Press. 1-2.
Rowe, Tykoski and Hutchinson, 1997. Ceratosauria. In Currie and Padian
(eds.). Encyclopedia of Dinosaurs. Academic Press. 106-110.
Sereno, 1998. A rationale for phylogenetic definitions, with application to
the higher-level taxonomy of Dinosauria. Neues Jahrbuch fur Geologie und Palaontologie.
210(1), 41-83.
Lamanna, Martinez and Smith, 2002. A definitive abelisaurid theropod dinosaur
from the early Late Cretaceous of Patagonia. Journal of Vertebrate Paleontology.
22(1), 58-69.
Wilson, Sereno, Srivastava, Bhatt, Khosla and Sahni, 2003. A new abelisaurid
(Dinosauria, Theropoda) from the Lameta Formation (Cretaceous, Maastrichtian)
of India. Contributions from the Museum of Paleontology. The University of Michigan.
31, 1-42.
Tykoski and Rowe, 2004. Ceratosauria. In Weishampel, Dodson and Osmolska (eds.).
The Dinosauria Second Edition. University of California Press. 47-70.
Sereno, online 2005. Stem Archosauria - TaxonSearch. http://www.taxonsearch.org/dev/file_home.php
[version 1.0, 2005 November 7]
Thompson, 2011. Comparative analysis of abelisaurid skulls
indicates dietary specialization. Journal of Vertebrate Paleontology. Program
and Abstracts 2011, 204.
Burch, 2013. The myological consequences of extreme limb reduction: New insights
from the forelimb musculature of abelisaurid theropods. Journal of Vertebrate
Paleontology. Program and Abstracts 2013, 97.
Delcourt, 2018. Ceratosaur palaeobiology: New insights on evolution and
ecology of the southern rulers. Scientific Reports. 8:9730.
Hendrickx,
Mateus, Araújo and Choiniere, 2019. The distribution of dental features
in non-avian theropod dinosaurs: Taxonomic potential, degree of
homoplasy, and major evolutionary trends. Palaeontologia Electronica.
22.3.74, 1-110.
Ibiricu, Baiano, Martinez, Alvarez, Lamanna and Casal, 2021. A detailed osteological description of Xenotarsosaurus bonapartei (Theropoda: Abelisauridae): implications for abelisauroid phylogeny. Cretaceous Research. 124, 104829.
"Carnosaurus" Huene, 1929
Late Cretaceous
east of Colhue Huapi Lake and north of Chico River, Chubut Group(?), Chubut,
Argentina
Material- (MACN coll.) tooth (23 x 12.5 x 6 mm)
Comments- Carnosaurus was listed by Huene (1929) in a faunal list
for three specimens- a metapodial (MLP CS 1240) from the Allen Formation, a
tooth (MACN coll.) perhaps from the Chubut group, and provisionally ("Cf.
Carnosaurus") a tooth (MLP coll.) from the Plottier Formation. As
Olshevsky (DML, 1999) noted, the name is probably a typographical error for
Carnosauria made when translating the paper from German to Spanish. This is
indicated by the fact he never attaches a name to these specimens in the description
or plates, and indeed states on of the specimens is taxonomically indistinguishable
from another named genus. Since "Carnosaurus" was apparently not meant
as a valid name when it was published (ICZN Article 11.5), it is a nomen nudum.
The metapodial is here identified as a sauropod or ankylosaur metatarsal, while
the Plottier tooth was only questionably referred by Huene, thus leaving the
Chubut tooth as the specimen best discussed under "Carnosaurus".
The Chubut tooth is merely described under the heading "tooth of a carnivorous
saurischian from the Chico River". The mesial edge is slightly concave
and the distal edge slightly convex. It has fine longitudinal ridges, and 30
serrations per 5 mm which are perpendicular to the edge. Huene stated there
was no difference between it and teeth he referred to Loncosaurus except
for the less curved mesial edge, which he felt could be explained by a different
position in the jaw. It compares well with abelisaurids in all variables except
the greater serration density, so may be a member of that clade.
References- Huene, 1929. Los saurisquios y ornitisquios del Cretacéo
Argentino. Anales del Museo de La Plata (series 3). 3, 1-196.
Olshevsky, DML 1999. https://web.archive.org/web/20200720012936/http://dml.cmnh.org/1999Nov/msg00507.html
Dryptosauroides Huene and
Matley, 1933
= "Dryptosauroides" Huene 1932
D. grandis Huene and Matley, 1933
= "Dryptosauroides grandis" Huene, 1932
Maastrichtian, Late Cretaceous
Lameta Formation, India
Syntypes- (GSI K20/334) proximal caudal vertebra
(GSI K20/609) proximal caudal vertebra
(GSI K27/549) proximal caudal vertebra
(GSI K27/601) proximal caudal vertebra
(GSI K27/602) proximal caudal vertebra
(GSI K27/626) proximal caudal vertebra
Diagnosis- Provisionally indeterminate at the level of Abelisauridae.
Comments- Novas et al. (2004) determined the vertebrae were proximal
caudals, not dorsals as suggested by Huene and Matley (1933). They are indistinguishable
from Ornithomimoides mobilis and O? barasimlensis.
References- Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre
Entwicklung und Geschichte. Monographien zur Geologie und Palaeontologie. 4(1),
viii + 361 pp.
Huene and Matley. 1933. The Cretaceous Saurischia and Ornithischia of the central
provinces of India. Palaeontologica Indica. 21, 1-74.
Novas and Bandyopaphyay, 1999. New approaches on the Cretaceous theropods from
India. VII International Symposium on Mesozoic Terrestrial Ecosystems, abstracts.
46-47.
Novas, Agnolin and Bandyopadhyay, 2004. Cretaceous theropods from India: A review
of specimens described by Huene and Matley (1933). Revista del Museo Argentino
del Ciencias Naturales. 6(1), 67-103.
Tarascosaurus Le Loeuff
and Buffetaut, 1991
T. salluvicus Le Loeuff and Buffetaut, 1991
Early Campanian, Late Cretaceous
Fuvelian beds, France
Holotype- (FSL 330201) proximal femur (~350 mm)
Paratype- ....(FSL 330202) fragment of ~sixth dorsal vertebra, partial
~seventh dorsal vertebra (~45 mm)
Referred- ?(FSL 330203) caudal centrum (Le Loeuff and Buffetaut, 1991)
Diagnosis- (after Le Loeuff and Buffetaut, 1991) very cavernous internal
structure of vertebrae; hyposphene-hypantrum articulations present; dorsal transverse
processes very wide anteroposteriorly; dorsal infradiapophyseal laminae diverge
beneath diapophyses; femoral neck particularly straight anteroposteriorly; femoral
head directed anteromedially; limited dorsal projection of anterior trochanter.
Comments- Though initially assigned to Abelisauridae by Le Loeuff and
Buffetaut (1991), it has more recently been assigned merely to Abelisauria (as
Abelisauroidea- Carrano and Sampson, 2008). However, Tortosa et al. (2014) recovered
it as closer to abelisaurids than noasaurids, and the improved version of that
study done by Filippi et al. (2016) further resolved it as outside the Rahiolisaurus+Abelisaurus
subclade of brachyrostrans.
References- Le Loeuff and Buffetaut, 1991. Tarascosaurus salluvicus
nov. gen., nov. sp., theropod dinosaur from the Upper Cretaceous of southern
France. Géobios - Paléontologie, Stratigraphie, Paléoécologie.
24, 585-594.
Carrano and Sampson, 2008 (online 2007). The Phylogeny of Ceratosauria (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 6, 183-236.
Tortosa, Buffetaut, Vialle, Dutour, Turini and Cheylan, 2014. A new abelisaurid
dinosaur from the Late Cretaceous of southern France: Palaeobiogeographical
implications. Annales de Paléontologie. 100(1), 63-86.
Filippi, Mendez, Juarez Valieri and Garrido, 2016. A new brachyrostran with
hypertrophied axial structures reveals an unexpected radiation of latest Cretaceous
abelisaurids. Cretaceous Research. 61, 209-219.
"Titanovenator" Boyd, 2020 online
"T. kenyaensis" Boyd, 2020 online
Maastrichtian, Late Cretaceous
Lapurr sandstone (= Turkana Grits), Turkana, Kenya
Material- (KNM-WT coll.) (~11-12 m) multiple specimens including cranial material (including
a partial skull - premaxilla, postorbital and braincase), axial material and
appendicular material (including astragalocalcaneum)
Comments-
Discovered in 2004, this is based on unassociated material referred to
one taxon "based on
morphological consistency and on the recovery of specimens from a
narrow stratigraphic and geographic area." Abelisaurid characters
listed by Sertich et al. (2013) include- "a tall, rugose premaxilla, an
anteroventrally inclined posterior border of the postorbital," ... "a
prominent dorsal projection of the parietals and supraoccipital" and an
"ascending process [that] is low and subrectangular, separated from the
anterior surface of the astragalus by a distinct fossa." They
further note the "skull is strongly coossified, with a thickened but
weakly sculptured skull roof", "no prominent cranial ornamentation is
evident" and "the astragalocalcaneum is completely coossified and
displays a prominent transverse sulcus on the anteroventral
surface." The museum collection is based on an azhdarchid
vertebra found by the same team from the same locality (O'Connor et
al., 2011).
Boyd (2020) included the
name Titanovenator kenyaensis in the pdf visual portion of her "Ask a
Geologist" presentation for RUGM. It accompanies a photo of the MCN Carnotaurus cast
that is also labeled Abelisauridae gen. et sp. nov., and based on
the name refers to Sertich et al.'s (2013) giant Kenyan abelisaurid
taxon. It is a nomen nudum as the pdf was not "issued for the
purpose
of providing a public and permanent scientific record" (ICZN Article
8.1.1) and doesn't "state the date of publication in the work itself"
(Article 8.5.2), while the name itself was not "registered in the
Official Register of Zoological Nomenclature (ZooBank)" (Article 8.5.3)
or "accompanied by a description or definition that states in words
characters that are purported to differentiate the taxon" (Article
13.1.1). The name (as "Titanovenator kenyanis") actually predates
Boyd's presentation, originating on a Jurassic Park message board from
2013 in a post under the pseudonym Rex Fan 684, but this is not
considered a source for catalogued nomina nuda here.
References- Sertich, Manthi, Sampson, Loewen and Getty, 2006. Rift Valley
dinosaurs: A new Late Cretaceous vertebrate fauna from Kenya. Journal of Vertebrate
Paleontology. 26(3), 124A.
O'Connor, Sertich and Manthi, 2011. A pterodactyloid pterosaur from the
Upper Cretaceous Lapurr sandstone, west Turkana, Kenya. Anais da
Academia Brasileira de Ciências. 83(1), 309-315.
McCoy, 2013 online. http://www.jurassicworld3.net/topic/28609
Sertich, Seiffert and Manthi, 2013. A giant abelisaurid theropod from the Latest
Cretaceous of northern Turkana, Kenya. Journal of Vertebrate Paleontology. Program
and Abstracts 2013, 211.
Boyd, 2020 online. Ask a Geologist: Geology & Paleontology of Kenya. Rutgers Geology Museum. 25 pp.
unnamed Abelisauridae (Astibia et al., 1990)
Late Campanian, Late Cretaceous
Laño, Sedano Formation, Spain
Material- (MCNA-1852; Morphotype 1) tooth (45x15x9 mm) (Torices, Currie, Canudo and Pereda-Suberbiola, 2015)
(MCNA-1853; Morphotype 2) tooth (24x13x7.3 mm) (Torices, Currie, Canudo and Pereda-Suberbiola, 2015)
(MCNA-4520; Morphotype 1) tooth (46x15x9 mm) (Torices, Currie, Canudo and Pereda-Suberbiola, 2015)
(MCNA-14521; Morphotype 1) tooth (62x16x8 mm) (Torices, Currie, Canudo and Pereda-Suberbiola, 2015)
(MCNA-14522; Morphotype 2) tooth (20x12x6.2 mm) (Torices, Currie, Canudo and Pereda-Suberbiola, 2015)
(MCNA-22051; Morphotype 1) tooth (43x19x8.9 mm) (Torices, Currie, Canudo and Pereda-Suberbiola, 2015)
(MCNA coll.) femora (one proximal) (Astibia et al., 1990)
Comments- Astibia et al. (1990) referred the femora to Abelisauridae?, while Le Loeuff and Buffetaut
(1991) called them cf. Tarascosaurus. The latter authors state (translated) the femora "show the same narrowness of the neck [as Tarascosaurus,
and] they also show a nutrient foramen beneath the lesser trochanter."
"However, the femora from Laño are more robust than that from Beausset,
with less anteriorly oriented heads."
Abelisaurid teeth were mentioned early by Astibia et al., who noted a
"large species (indicated by a few large teeth, 3-5 cm long)" which
"may belong to the same form as the above-mentioned femora."
Torices et al. (2015) assigned these to Morphotypes 1 and 2 of
Theropoda indet., but Isasmendi et al. (2020) have more recently called
them cf. Arcovenator.
References- Astibia, Buffetaut, Buscalioni, Cappetta, Corral, Estes,
Garcia-Garmilla, Jaeger, Jimenez-Fuentes, Le Loeuff, Mazin, Orue-Extebarria,
Pereda-Superbiola, Powell, Rage, Rordriguez-Lazaro, Sanz and Tong, 1990. The
fossil vertebrates from Laño (Basque Country, Spain); New evidence on
the composition and affinities of the Late Cretaceous continental faunas of
Europe. Terra Research. 2, 460-466.
Le Loeuff and Buffetaut, 1991. Tarascosaurus salluvicus nov. gen., nov.
sp., theropod dinosaur from the Upper Cretaceous of southern France. Géobios
- Paléontologie, Stratigraphie, Paléoécologie. 24, 585-594.
Torices, Currie, Canudo and Pereda-Suberbiola, 2015. Theropod
dinosaurs from the Upper Cretaceous of the South Pyrenees Basin of Spain. Acta
Palaeontologica Polonica. 60(3), 611-626.
Isasmendi, Torices, Canudo and Pereda-Suberbiola, 2020.
Paleobiodiversity of theropod dinosaurs from the Upper Cretaceous Laño
site, northern Iberian peninsula. The Society
of Vertebrate Paleontology 80th
Annual Meeting, Conference Program. 186-187.
unnamed Abelisauridae (Torices, 2002)
Late Campanian-Early Maastrichtian, Late Cretaceous
Montrebei, Tremp Formation, Spain
Material- (DPM-MON-T10; Morphotype 1) tooth (42x15x7 mm) (Torices, 2002)
Late Maastrichtian, Late Cretaceous
Blasi 2B and 3, Tremp Formation, Spain
(MPZ98/67; Morphotype 1) tooth (27.5x27x13 mm) (Torices, Currie, Canudo and
Pereda-Suberbiola, 2015)
(MPZ2004/5; Morphotype 1) tooth (18 mm) (Torices, Currie, Canudo and Pereda-Suberbiola,
2015)
(MPZ2004/8; Morphotype 1) tooth (28x12x7.5 mm) (Torices, Currie, Canudo and
Pereda-Suberbiola, 2015)
Comments- Torices et al. (2015) assigned these to Morphotype 1 of
Theropoda indet., but Isasmendi et al. (2020) have more recently called this morphotype cf. Arcovenator.
References- Torices, 2002. Los dinosaurios terópodos del Cretácico
Superior de la Cuenca de Tremp (Pirineos Sur-Centrales, Lleida). Coloquios de
Paleontología. 53, 139-146.
Torices, Currie, Canudo and Pereda-Suberbiola, 2015. Theropod dinosaurs from
the Upper Cretaceous of the South Pyrenees Basin of Spain. Acta Palaeontologica
Polonica. 60(3), 611-626.
unnamed Abelisauridae (Osi and Apesteguia, 2008)
Santonian, Late Cretaceous
Csehbanya Formation, Hungary
Material- (MTM V 2008.43.1) pedal ungual (Osi and Apesteguia, 2008)
(MTM coll.) fragmentary femur (Osi et al., 2012)
References- Osi and Apesteguia, 2008. Nonavian theropod dinosaur remains
from the Upper Cretaceous (Santonian) Csehbánya Formation (Iharkút,
Bakony Mountains, Western Hungary). EAVP 2008. 78-79.
Osi, Rabl, Makadl, Szentesi, Botfalval and Gulyas, 2012. The Late Cretaceous
continental vertebrate fauna from Iharkút (Western Hungary): A review.
In Godefroit (ed.). Bernissart Dinosaurs and Early Cretaceous Terrestrial Ecosystems.
Indiana University Press. 532-569.
Osi and Buffetaut, in press.
unnamed Abelisauridae (Kear, Rich, Vickers-Rich, Ali, Al-Mufarreh, Matari, Al-Massari, Nasser, Attia and Halawani, 2013)
Campanian-Early Maastrichtian, Late Cretaceous
Adaffa Formation, Aynunah Trough, Saudi Arabia
Material- (SGS 0061) tooth fragment (19.7x?x? mm)
(SGS 0090) lateral tooth (17.1x11.6x4.7 mm)
Comments- Collected between
2004 and 2008, Kear et al. (2013) state in these specimens "the distal
profile is clearly linear, consistent with the distinctive
maxillary-dentary teeth of abelisaurid ceratosaurians" and recover them
closest to Majungasaurus and Lameta AMNH 1753/1955 in most quantitative analyses.
Reference- Kear, Rich,
Vickers-Rich, Ali, Al-Mufarreh, Matari, Al-Massari, Nasser, Attia and
Halawani, 2013. First dinosaurs from Saudi Arabia. PLoS ONE. 8(12),
e84041.
unnamed Abelisauridae (Russell, 1996)
Cenomanian, Late Cretaceous
Kem Kem beds, Morocco
Material-
(CMN 41861) dentary fragment (teeth FABL 6-14 mm) (Russell, 1996)
(CMN 50403; bone taxon C) (~10.7 m) incomplete posterior dorsal vertebra
(158 mm) (Russell, 1996)
(CMN 50446) tooth (~24x~11x? mm) (Ibrahim, Sereno, Varricchio, Martill, Dutheil, Unwin, Baidder, Larsson,
Zouhri and Kaoukaya, 2020)
?(FSAC-KK 912) tooth (~17x~10x? mm) (Ibrahim, Sereno, Varricchio, Martill, Dutheil, Unwin, Baidder, Larsson,
Zouhri and Kaoukaya, 2020)
?(FSAC-KK 913) tooth (~22x~12x? mm) (Ibrahim, Sereno, Varricchio, Martill, Dutheil, Unwin, Baidder, Larsson,
Zouhri and Kaoukaya, 2020)
?(FSAC-KK 914) partial tooth (Ibrahim, Sereno, Varricchio, Martill, Dutheil, Unwin, Baidder, Larsson,
Zouhri and Kaoukaya, 2020)
?(FSAC-KK 915) tooth (~18x~8x? mm) (Ibrahim, Sereno, Varricchio, Martill, Dutheil, Unwin, Baidder, Larsson,
Zouhri and Kaoukaya, 2020)
(FSAC-KK-5015) (~2.7 m) axis (38 mm) (Smyth, Ibrahim, Kao and Martill, 2020)
?(FSAC-KK unnumbered) tooth (~16x~11x? mm) (Ibrahim, Sereno, Varricchio, Martill, Dutheil, Unwin, Baidder, Larsson,
Zouhri and Kaoukaya, 2020)
(GZG.V.19996) tooth (19x9.8x4.8 mm) (Richter, Mudroch and Buckley,
2013)
(GZG.V.19999) tooth (14.1x5.6x3.6 mm) (Richter, Mudroch and Buckley, 2013)
(MNHN MRS 348) tooth (48x17x13 mm) (Fanti, Cau, Martinelli and Contessi, 2014)
(MNHN MRS 783) tooth (28x14x6 mm) (Fanti, Cau, Martinelli and Contessi, 2014)
(MNHN MRS 1266) tooth (34x14x7 mm) (Fanti, Cau, Martinelli and Contessi, 2014)
(MNHN MRS 1838) tooth (36x19x9 mm) (Fanti, Cau, Martinelli and Contessi, 2014)
(MPUR NS 153/01) partial maxilla (Porchetti, Nicosia, Biava and Maganuco, 2011)
(MSNM V6053) tooth (~23x~10x? mm) (Ibrahim, Sereno, Varricchio, Martill, Dutheil, Unwin, Baidder, Larsson,
Zouhri and Kaoukaya, 2020)
(NMB-1672-R) tooth (21x9x4.5 mm) (Richter, Mudroch and Buckley, 2013)
(OLPH 025) (~7 m; ~1.85 tons) proximal femur (~770-920 mm) (Chiarenza and Cau, 2016) 1127
Comments- Russell (1996) described dentary fragment CMN 41861 as cf. Majungasaurus sp., but "shows only more broadly distributed abelisaurid features and is therefore probably not referable to Majungasaurus" as noted by Carrano and Sampson (2008). Richter et al. (2013) described three teeth as abelisaurid-like, one of which
(GZG.V.19999) was suggested to be noasaurid by Smyth et al. (2020) based on its high DSDI as in Masiakasaurus. Yet teeth with high DSDIs are also known in some Majungasaurus and Arcovenator specimens,
so it is retained as Abelisauridae here. Fanti et al. (2014) list
measurements of four "Abelisauridae indet." teeth housed in the MNHN
which have MRS numbers corresponding to the Gara Sbaa locality (e.g. Rebbachisaurus type) of the Kem Kem beds. Chiarenza and
Cau (2016) note the large and heart-shaped neural canal of Russell's
(1996) bone taxon C is similar to Libyan abelisaurid PRC.NF.1.21, so
propose they are closely related. Smyth et al. suggested the axis
FSAC-KK-5015 represents an abelisaurid outside
Majungasaurinae+Brachyrostra, perhaps a juvenile Rugops.
Ibrahim et al. (2020) figure CMN 50446 and MSNM V6053 as "?Abelisaurid
tooth", and several others as "Indeterminate theropod teeth" which seem
to be abelisaurid based on the low distal curvature in each, and
marginal undulations in FSAC-KK 912 and 915.
References- Russell, 1996. Isolated dinosaur bones from the Middle Cretaceous
of the Tafilalt, Morocco. Bulletin du Museum national d'Histoire naturelle.
18, 349-402.
Carrano and Sampson, 2008 (online 2007). The Phylogeny of Ceratosauria (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 6, 183-236.
Porchetti, Nicosia, Biava and Maganuco, 2011. New abelisaurid material from
the Upper Cretaceous (Cenomanian) of Morocco. Rivista Italiana di Paleontologia
e Stratigrafia. 117(3), 463-472.
Richter, Mudroch and Buckley, 2013. Isolated theropod teeth from the Kem Kem
Beds (Early Cenomanian) near Taouz, Morocco. Palaontologische Zeitschrift. 87,
291-309.
Fanti, Cau, Martinelli and Contessi, 2014. Integrating palaeoecology
and morphology in theropod diversity estimation: A case from the Aptian-Albian
of Tunisia. Palaeogeography, Palaeoclimatology, Palaeoecology. 410, 39-57.
Chiarenza and Cau, 2016. A large abelisaurid (Dinosauria, Theropoda) from Morocco
and comments on the Cenomanian theropods from North Africa. PeerJ. 4:e1754.
Ibrahim, Sereno, Varricchio, Martill, Dutheil, Unwin, Baidder, Larsson,
Zouhri and Kaoukaya, 2020. Geology and paleontology of the Upper
Cretaceous Kem Kem Group of eastern Morocco. ZooKeys. 928, 1-216.
Smyth, Ibrahim, Kao and Martill, 2020 (online 2019). Abelisauroid
cervical vertebrae from the Cretaceous Kem Kem beds of southern Morocco
and a review of Kem Kem abelisauroids. Cretaceous Research. 108, 104330.
unnamed Abelisauridae (Fanti, Cau, Martinelli and Contessi, 2014)
Late Aptian-Early Albian, Early Cretaceous
Chenini or Oum Ed Diab Member of the Ain el Guettar Formation, Dahar/Jeffara escarpment, Tunisia
Material- (MGGC 21889) dentary fragment
(MGGCTUN15; Morphotype 6) lateral tooth (24x15x11 mm)
(MGGCTUN16; Morphotype 8) (juvenile?) lateral tooth (18x13x6 mm)
(MGGCTUN27; Morphotype 6) lateral tooth (32x20x10 mm)
(MGGCTUN28; Morphotype 8) (juvenile?) lateral tooth (14x11x5 mm)
(MGGCTUN30; Morphotype 6) lateral tooth (31x19x9 mm)
(MGGCTUN32; Morphotype 6) lateral tooth (35x19x13 mm)
(MGGCTUN45; Morphotype 6) lateral tooth (18x16x10 mm)
(MGGCTUN47; Morphotype 6) lateral tooth (29x19x12 mm)
(MGGCTUN75; Morphotype 8) (juvenile?) lateral tooth (13x10x5 mm)
(MGGCTUN76; Morphotype 6) lateral tooth (24x14x8 mm)
(MGGCTUN78; Morphotype 8) (juvenile?) lateral tooth (20x14x7 mm)
(MGGCTUN79; Morphotype 6) lateral tooth (26x18x13 mm)
(MGGCTUN80; Morphotype 6) lateral tooth (16x9x6 mm)
(ONM TM 02) dentary fragment
Comments- Fanti et al. (2014)
referred MGGC 21889 "Based on the combination of distal paradental
laminae that are fused and ornamented by lingual furrows and
rugosities", with ONM TM 02 presumably placed there for the same
reasons although it is unillustrated and undescribed except to note it
only preserves four alveoli instead of five. They used a version
of Cau's theropod megamatrix to resolve the former specimen in
Abelisauridae.
Fanti et al. (2014) noted "diagnostic features of abelisaurid teeth are
observed in ... Morphotype" 6, such as "the crown displaying a
sub-triangular shape with the mesial side curved (with a typical
inflexion point near the midlenght of the crown) and the distal one
almost straight, 2.5-3 denticles per mm, and well developed blood
grooves." They further found that in Morphotype 8 "The posterior
carina also displays an inflection point where the curvature becomes
more pronounced distally, and blood grooves are pronounced, both
characters typically observed in abelisaurid teeth", but because "The
relatively dense serrations (4.5 denticles per mm in the mesial carina
and 4 per mm in the distal one) are comparable with those of
posteriormost lateral teeth of abelisaurids", they suggested "that
Morphotype 8 includes abelisaur posterior lateral teeth or possibly
juvenile lateral teeth." Both morphotypes emerged as sister taxa
in Abelisauridae when Fanti et al. analyzed them in Hendrickx's
theropod tooth matrix.
Reference- Fanti, Cau, Martinelli and Contessi, 2014. Integrating palaeoecology
and morphology in theropod diversity estimation: A case from the Aptian-Albian
of Tunisia. Palaeogeography, Palaeoclimatology, Palaeoecology. 410, 39-57.
unnamed Abelisauridae (Dalla Vecchia, 1995)
Hauterivian-Barremian, Early Cretaceous
Cabao Formation, Nalut, Libya
Material- (MPCM 13693) tooth (27.5x16.2x9.8 mm) (Dalla Vecchia, 1995)
?(PRC.NF.1.21) (~7-9 m; subadult) incomplete posterior dorsal vertebra
(~90 mm), proximal caudal centrum (~111 mm), proximal caudal neural
arch fragment, distal femur (~810-1013 mm), proximal tibia (Smith,
Tshakreen, Rasmussen and Lamanna, 2006; described in Smith, Lamanna,
Askar, Bergig, Tshakreen, Abugares and Rasmussen, 2010)
Comments- Dalla Vecchia (1995) described a theropod tooth
discovered in 1984 and supposedly from the Chicla Sandstone Formation
(also called the Kiklah Formation), but Le Loeuff et al. (2010) stated
"our own investigations in this area suggest that this material comes
rather from the Cabao Formation." While Dalla Vechhia only determined
the tooth was not Carcharodontosaurus,
Smith and Dalla Vecchia (2006) found it "possesses several features
common to abelisaurids but distinct from other Theropoda", including a
"mesial curvature profile [that] describes a strong curve beginning at
about the midpoint of the crown" and a "distal curvature profile [that]
exhibits almost no curvature." Furthermore, as in Rugops but not Majungasaurus
or Lameta specimens AMNH 1753/1955, "the denticles of MPCM 13693
exhibit no traces of the caudae/interdenticular sulci complex, which
might suggest that the specimen is derived from a more basal member of
the clade." Furthermore, quantitave analysis classified it with
the Lameta material, although the authors note the temporal and
geographic distances are too great to belong to a single taxon.
Discovered in August 2005 at locality PRC NF001, Smith et al. (2006)
first suggested in an abstract PRC.NF.1.21 "appears to be a previously
unrecorded taxon as evidenced by several autapomorphies" and "possesses
femoral and tibial characters very similar to Masiakasaurus and is thus
perhaps a large-bodied noasaurid." Smith and Lamanna (2009) in a
following abstract merely stated it was "referable to the widespread
ceratosaurian clade Abelisauroidea." Once fully described in
2010, Smith et al. (2010) suggested since the vertebrae "exhibit clear
neurocentral sutures" ... "despite its large size, the individual
represented by PRC.NF.1.21 was not fully mature at death." They
said it "may be referable to Abelisauridae because the poorly
differentiated lateral tibial condyle is located at the same
proximodistal level as is the medial condyle, but we cannot
conclusively refer the specimen to this more exclusive abelisauroid
clade at this time." Finally, Smith et al. stated "the proximally
bifurcate fibular crest of PRC.NF.1.21 appears unique to this specimen,
indicating that the Libyan form probably pertains to a previously
unrecognized genus or species." This and MPCM 13693 "might be
conspecific, although this cannot presently be confirmed."
References- Dalla Vecchia, 1995. Second record of a site with dinosaur
skeletal remains in Libya (northern Africa). Natura Nascosta. 11, 16-19.
Smith and Dalla Vecchia, 2006. An abelisaurid (Dinosauria: Theropoda) tooth
from the Lower Cretaceous Chicla formation of Libya. Journal of African Earth
Sciences. 46, 240-244.
Smith, Tshakreen, Rasmussen and Lamanna, 2006. New dinosaur discoveries from
the Early Cretaceous of Libya. Journal of Vertebrate Paleontology. 26(3), 126A.
Smith and Lamanna, 2009. Abelisauroid dinosaurs from the Early Cretaceous of
Libya. Geological Society of America, Abstracts with Programs. 41, 242.
Le Loeuff, Métais, Dutheil, Rubinos, Buffetaut, Ois Lafont, Cavin, Moreau,
Tong, Blanpied and Sbeta, 2010. An Early Cretaceous vertebrate assemblage from
the Cabao Formation of NW Libya. Geological Magazine. 147(5), 750-759.
Smith, Lamanna, Askar, Bergig, Tshakreen, Abugares and Rasmussen, 2010. A large
abelisauroid theropod dinosaur from the Early Cretaceous of Libya. Journal of
Paleontology. 84(5), 927-934.
unnamed abelisaurid (Stromer and Weiler, 1930)
Campanian, Late Cretaceous
layer 3-4a, Naga el Ledaia, Nubian Sandstone Formation, Egypt
Material- (HMN MB.R coll?; destroyed) proximal tibia (93x40 mm prox)
Comments-
Stromer and Weiler (1930) called this "Theropoda g. et sp. indet.",
noting closest similarities to abelisaurid HMN MB.R 3626 (Tibia 69 of
Janensch) from the Tendaguru Formation and Ceratosaurus,
but stated "If the mentioned similarities are sufficient to proof a
relationship with the Ceratosauridae remains questionable to me"
(translated). They distinguished it from HMN MB.R 3626 in "the
concave lower edge of the ... crista [cnemialis] and in a concavity at
its inner side, in the less stronger vaulting of the inner surface and
in larger width and very slight separation of the inner and outer
condylus." Carrano et al. (2002) believed it "may represent an
abelisauroid, but unfortunately this material was destroyed in World
War II." Buffetaut et al. (2005) stated it resembles abelisaurids
in having a "proximally ascending cnemial crest and a fibular condyle
confluent with the cnemial crest."
References- Stromer and Weiler, 1930. Ergebnisse der Forschungsreisen
Prof. E. Stromers in den Wüsten Ägyptens. VI. Beschreibung von Wirbeltier-Resten
aus dem nubischen Sandsteine Oberägyptens und aus ägyptischen Phosphaten
nebst Bemerkungen über die Geologie der Umgegend von Mahamîd in Oberägypten. Abhandlungen der Bayerischen Akademie
der Wissenschaften Mathematisch-naturwissenschaftliche Abteilung, Neue Folge.
7, 1-42.
Carrano, Sampson and Forster, 2002. The osteology of Masiakasaurus knopfleri,
a small abelisauroid (Dinosauria: Theropoda) from the Late Cretaceous of Madagascar.
Journal of Vertebrate Paleontology. 22(3), 510-534.
Buffetaut, Escuillie and Pohl, 2005. First theropod dinosaur from the Maastrichtian
phosphates of Morocco. Kaupia. 14, 3-8.
unnamed probable abelisaurid (Sallam, O'Connor, Kora, Sertich, Seiffert, Faris, Ouda, El-Dawoudi, Saber and El-Sayed, 2016)
Middle Campanian, Late Cretaceous
Baris, El Hindaw Member, Quseir Formation, Kharga Oasis, Egypt
Material- (MUVP 187) proximal fibula (25x5 mm prox)
Comments- Discovered in 2008 or
2010, Sallam et al. (2016) initially reported "A proximal left fibula
of a theropod dinosaur was also collected from the Baris area" and
listed it as Theropoda indet.. Salem et al. (2021) noted the
prominent m. iliofibularis tubercle is like abelisaurids and it has a
"wide, slightly deep [proximomedial] fossa that closely resembles the
condition in the abelisaurid Majungasaurus." They say it "exhibits some affinities with abelisaurids" and refer to it as "Cf. Abelisaurid" in theit table 3.
References- Sallam, O'Connor,
Kora, Sertich, Seiffert, Faris, Ouda, El-Dawoudi, Saber and El-Sayed,
2016. Vertebrate paleontological exploration of the Upper Cretaceous
succession in the Dakhla and Kharga Oases, Western Desert, Egypt.
Journal of African Earth Sciences. 117, 223-234.
Salem, O'Connor, Gorscak, El-Sayed, Sertich, Seiffert and Sallam, 2021.
Dinosaur remains from the Upper Cretaceous (Campanian) of the Western
Desert, Egypt. Cretaceous Research. 123, 104783.
unnamed Abelisauridae (Di Stefano, 1919)
Late Campanian, Late Cretaceous
Duwi Formation, Al Sharauna, Egypt
Material- (MGUP MEGA002; lost?) tooth (16.63x13.01x6.69 mm) (described by Smith and
Lamanna, 2006)
Late Campanian, Late Cretaceous
Duwi Formation, Al Sharauna, Gebel Duwi and/or Gebel Nakheil, Egypt
(MGUP MEGA001) partial lateral tooth (?x~22.7x? mm) (Di Patti, pers. comm. 6-2023)
(MGUP MEGA002 B) lateral tooth (~35.2x~25x? mm) (Di Patti, pers. comm. 6-2023)
(MGUP MEGA003) partial lateral tooth (?x~30x? mm) (Di Patti, pers. comm. 6-2023)
(MGUP coll.; lost) more than two lateral teeth
Comments- Gemmellaro (1921) referred several remains to Megalosaurus crenatissimus,
from three localities in eastern Egypt. The phosphate layers of Gebel
Duwi are the Late Campanian Duwi Formation (e.g. Salama et al., 2021).
Similarly, "Phosphate mines at Kosseir-el-Khadim, Gebel Nakheil, near
Kosseir [now Quseer], on the Red Sea" are also the Duwi Formation
(Valentine, 1985). The third locality has been mistaken in every
subsequent reference as "Sciarauna-el-Ghibli, presse Sibaiya (Valle del
Nilo)", but the 'c' is actually a poorly photocopied 'e', as indicated
by Di Stefano's (1919) original publication of the expedition which
states "The
richest and most extensive deposit is the one found on the hill of
Seiarauna, on which are the two villages called Seiarauna-el-Bahri,
i.e. Seiarauna at the river, and Seiarauna-el-Ghibli, i.e. upstream"
(translated). The latter is now known as Ash Sharawinah al Qibliyah,
El-Sharâwna el-Qiblîya or simply Al Sharauna among other variants, and is indeed directly across the Nile from Sebaiya (also called As Sibaiyyah and
Al Sebaaia West City). Smith and Lamanna (2005) stated the particular
tooth they described was from "near Idfu in the Nile Valley of Egypt
(Fig. 1, exact locality data are unknown; research in Palermo did not
reveal a more accurate location)", but Idfu (also called Edfu) is 30 km
to the southeast. Gemmellaro states "the majority of the specimens
came from [Seiarauna-el-Ghibli]. But a careful research recently
conducted by myself on the abundant material donated by Cortese has
allowed me to find all the types studied also in the locations of Gebel
Nakheil (Kosseir-el-Khadim and Uadi-el-Anz mines) and of Gebel Duwi"
(translated), suggesting no other localities were involved and perhaps
30 km was considered "near" by whoevever wrote that record.
Di Stefano (1919) first reported theropod material from Al Sharauna,
collected by him in 1912 and also donated to the MGUP by Cortese
between then and his publication- "In contact between the phosphates
and the alternating marls, bones and quite a few dinosaurian teeth (Mosasaurus, Platecarpus, Megalosaurus,
etc.) are found, which will be the subject of a special study." That
study was Gemmellaro (1921), who as noted above also discovered
theropod material in the Gebel Duwi and Gebel Nakheil collections.
Unfortunately, Gemmellaro never specifies which of the three localities
any of his discussed specimens is from, nor does he state how many
teeth were in the MGUP collections. Three teeth are figured, with
those in figures 12 and 13 typically abelisaurid with low crowns
serrated to the base both mesially and distally. Curvature and crown
height ratios are most similar to posterior dentary teeth of
Majungasaurus. Gemmellaro compares the teeth favorably to Majungasaurus
syntypes FSL 92.306a and b and a Maevarano tooth figured by Thevenin
1906 (MNHN coll.). A smaller, third tooth is illustrated as
figure 14,
said to be among "some which, although sharing some common features,
differ in the shape of the crown which is not backward curved but is
almost isosceles with the sides having a profile slightly convex or
straight." While Gemmellaro believed "such differences in shape
and in
dimensions between teeth has to be attributed to the diverse locations
they occupied in the jaws, and also to the different age of the
individuals whom the teeth belonged to", the mesiodistally narrower
crown suggests another taxon such as a noasaurid, and that tooth is
listed as Averostra indet. here. Gemmellaro described two unguals
and figured one. While he compares them favorably to crenatissimus
syntype FSL 92.290, the preserved section of the figured specimen is
less tapered in side view with no obvious vascular groove, and this
holds true as well for other Majungasaurus pedal unguals (e.g. FMNH PR 2434, MSNM V6418, V6419) as well as Masiakasaurus
(e.g. FMNH PR 2135) and indeed most theropods. Photos provided by
Di Patti (pers. comm. 6-2023) show the figured element (MGUP MEGA004)
is a sauropod pedal ungual while the other (MGUP MEGA005) compares well
to an ankylopollexian manual ungual I. They are referred to
Titanosauria indet. and Hadrosauroidea indet. here based on age.
Smith and Lamanna (2006) described one of Gemmellaro's teeth in detail
(MGUP MEGA002), although it is not one previously figured. They
identified it as abelisaurid based on "the distal curvature profile
exhibits almost no curvature and is tilted slightly toward the apex in
lateral view" (but this is not true of Gemmellaro's Figure 12), and as
closer to Majungasaurus and Lameta AMNH 1753/1955 than Rugops, Rugops?
sp. UCPC 10 or Bajo Barreal UNPSJB-PV 247 based on "mesial curvature
profile begins at a strong curve at about 1/3 the crown height basal to
the apex (but this is not true of Gemmellaro's Figure 12) and "the ...
interdenticular sulci complex forms an intermediate condition between
being absent and well developed" (unknown in Gemmellaro's figured
specimens). They find "The discriminant analysis (DA) correctly
identified 96.6% of the teeth in the sample and classified MGUP MEGA002
as Majungasaurus" but note "the assignment of the Egyptian tooth to the genus Majungasaurus
in particular is unlikely given that Africa and Madagascar were
separated by the Mozambique Channel throughout the Cretaceous", and
indeed the relatively low sample of abelisaurid teeth tested makes this
questionable. In fact he combination of stout shape plus concave
basal distal edge with a convex apical distal edge is unlike figured Majungasaurus teeth.
Interestingly, the MGUP only has three teeth in their collection now
(Di Patti, pers. comm. 6-2023), none of which are MGUP MEGA002 or match
the three illustrated by Gemmellaro. One (MGUP MEGA002 B) is
abelisaurid based on the slightly convex distoapical edge, although the
mesial edge is similar to Gemmellaro's Figure 12 instead of curving
abruptly at a third of its height. Marginal undulations are
present as in some other abelisaurids (Abelisaurus, Chenanisaurus, Majungasaurus, Skorpiovenator).
There are ~10 mesial serrations per 5 mm and ~11 distal
serrations. The other two teeth (MGUP MEGA001 and MGUP MEGA003)
are more partially preserved but of similar size and serration density
(~10 distal serrations per 5 mm in MEGA001; ~10.5 mesial serrations per
5 mm in MEGA003), and each shows marginal undulations as well, so
they are tentatively placed as Abelisauridae here. There are no
precise locality data for any of them past Maastrichtian of Egypt.
References-
Di Stefano, 1919. Osservazioni sul Cretaceo e sull'Eocene del Deserto
Arabico di el-Sibaiya, nella valle del Nilo. Bollettino del R. Comitato
Geologico d'Italia. Serie V(47), 1-39.
Gemmellaro, 1921. Rettili maëstrichtiani di Egitto. Giornale
di Scienze Naturali ed Economiche. 32, 339-351.
Valentine, 1985. Structure and tectonics of the southern Gebel Duwi
area, Eastern Desert of Egypt. Department of Geology and Geography,
University of Massachusetts. Contribution No. 53, 141 pp.
Smith and Lamanna, 2006. An abelisaurid from the Late Cretaceous of Egypt: Implications
for theropod biogeography. Naturwissenschaften. 93(5), 242-245.
Salama, Altoom, a Allam, Ajarem and Abd-Elhameed, 2021. Late Cretaceous anacoracid sharks (Squalicorax)
from Duwi Formation, Gebel Duwi, central Eastern Desert, Egypt:
Qualitative and quantitative analyses. Historical Biology. 33(11),
3056-3064.
unnamed Abelisauridae (Pochat-Cottilloux, Allain and Lasseron, 2022)
Aptian, Early Cretaceous
GAD 5, Gadoufaoua, Elrhaz Formation, Niger
Material- (MNHN F.GDF-M30) two tooth fragments
(MNHN F.GDF-M37) two partial teeth
Comments- Collected in 1970,
Pochat-Cottilloux et al. (2022) noted "some of this material was stored
at the MNHN in a separate box" and had a "lighter color and a better
quality of fossil preservation than other seen in other samples" which
"suggested a different depositional environment to that of other
samples so, as a result, this sample was separated from the rest" ...
"as "site GADb"." This is true for MNHN F.GDF-M30. The
authors state there are 3 serrations per mm on MNHN F.GDF-M30 and 4 per
mm on MNHN F.GDF-M37, in the latter the carinae "extend along the
entire length of both sides of the base" and in all specimens "the
shape of the denticles and their number are typical of theropod teeth
and could be assignable to Abelisauridae."
Reference- Pochat-Cottilloux,
Allain and Lasseron, 2022. Microvertebrate fauna from Gadoufaoua
(Niger, Aptian, Early Cretaceous). Comptes Rendus Palevol. 21(41),
901-926.
undescribed abelisaurid (Sertich, Manthi, Sampson, Loewen and Getty,
2006)
Maastrichtian, Late Cretaceous
Lapurr sandstone (= Turkana Grits), Turkana, Kenya
Material- (KNM-WT coll.) (<11 m) teeth and appendicular elements
Comments- Sertich et al. (2006) noted "dental and
appendicular materials suggest the presence of two abelisaurid theropod
taxa, one large-bodied and the other giant-sized", the latter of which
is catalogued under the nomen nudum "Titanovenator" here. The
museum collection is based on an azhdarchid vertebra found by the same
team from the same locality (O'Connor et al., 2011).
References- Sertich, Manthi, Sampson, Loewen and Getty, 2006. Rift Valley
dinosaurs: A new Late Cretaceous vertebrate fauna from Kenya. Journal of Vertebrate
Paleontology. 26(3), 124A.
O'Connor, Sertich and Manthi, 2011. A pterodactyloid pterosaur from the
Upper Cretaceous Lapurr sandstone, west Turkana, Kenya. Anais da
Academia Brasileira de Ciências. 83(1), 309-315.
unnamed abelisaurid (Janensch, 1925)
Late Tithonian, Late Jurassic
Upper Dinosaur Member of the Tendaguru Formation, Tanzania
Material- (HMN MB R 3621; = 68 of Janensch) femur (773 mm)
....(HMN MB R 3626; = 69 of Janensch) tibia (~610 mm)
Late Kimmeridgian, Late Jurassic
Middle Dinosaur Member of the Tendaguru Formation, Tanzania
Material- (MB R 3725; = 37 of Janensch) tibia (567 mm)
Comments- Janensch (1925) stated these were similar to Ceratosaurus
and thought they might be referrable to C. roechlingi, but Rauhut (2011)
notes they are more similar to abelisaurids. They are listed as Tetanurae indet. by Carrano (1998).
References- Janensch, 1925. Die Coelurosaurier und Theropoden der Tendaguru-Schichten
Deutsch-Ostafrikas. Palaeontographica. 1(supp. 7), 1-99.
Carrano, 1998. The evolution of dinosaur locomotion: Functional morphology,
biomechanics, and modern analogs. PhD Thesis, The University of Chicago. 424
pp.
Rauhut, 2011. Theropod dinosaurs from the Late Jurassic of Tendaguru (Tanzania).
Palaeontology. 86, 195-239.
unnamed Abelisauridae (O'Connor, Gottfried, Stevens, Roberts, Ngasala, Kapilima and
Chami, 2006)
Albian, Early Cretaceous
TZ-07, Namba Member of the Galula Formation, Tanzania
Material- (TNM 02088) lateral tooth (~19x~11.5x? mm)
(TNM coll.) eight lateral teeth, two anterior teeth
Comments- Discovered between
2002 and 2005, this material was described as theropod by O'Connor et
al. (2006) and referred to the Unit I of the Red Sandstone Group at the
time. Roberts et al. (2010) subsequently revised the stratigraphic
nomenclature, naming Unit I the Galula Formation. Salem et al. (2021)
refers the isolated teeth to Abelisauridae without comment, which seems
likely given their stratigraphy, locality, size and morphology
(particularly the nearly straight distal edge). Several of the teeth
were associated with the holotype of the titanosaur Shingopana,
"suggesting a possible scavenging event." Note the theropod
caudals from the same locality described by O'Connor et al. differ from
abelisaurids in their strong infradiapophyseal laminae and are here
tentatively referred to Megaraptora.
Teeth have ~13-16 serrations per 5 mm mesially and distally. O'Connor
et al. further indicate "Two teeth are D-shaped in cross-section,
indicating a position in the rostralmost portion of either the dentary
or premaxilla."
References- O'Connor, Gottfried, Stevens, Roberts, Ngasala, Kapilima and
Chami, 2006. A new vertebrate fauna from the Cretaceous Red Sandstone Group,
Rukwa Rift Basin, southwestern Tanzania. Journal of African Earth Sciences.
44, 277-288.
Roberts, O'Connor, Stevens, Gottfried, Jinnah, Ngasala, Choh and
Armstrong, 2010 (online 2009). Sedimentology and depositional
environments of the Red Sandstone Group, Rukwa Rift Basin, southwestern
Tanzania: New insight into Cretaceous and Paleogene terrestrial
ecosystems and tectonics in sub-equatorial Africa. Journal of African
Earth Sciences. 57, 179-212.
Salem, O'Connor, Gorscak, El-Sayed, Sertich, Seiffert and Sallam, 2021.
Dinosaur remains from the Upper Cretaceous (Campanian) of the Western
Desert, Egypt. Cretaceous Research. 123, 104783.
undescribed Abelisauridae (Bond and Bromley, 1970)
Early Cretaceous
Gokwe Formation, Zimbabwe
Material- (University of Zimbabwe coll.) at least two teeth (~48x~31x?, ~49x~34x? mm)
(Natural History Museum of Zimbabwe coll?) (medium-sized) material including many teeth and several vertebrae
Comments- Bond and Bromley
(1970) were the first to figure three theropod teeth, discovered
between 1962
and 1970, as their Plate 1A "Reptilian teeth of two types from the
Gokwe Formation." Listed as being " in collection of Geology
Department, University College of Rhodesia", the latter has since been
renamed the University of Zimbabwe. The first and third are
clearly abelisaurid, with broad bases, straight to convex distal edges
and fine serrations both mesially and distally (~6 per 5 mm on both
carinae). The second is narrower and more recurved, so referred
to Averostra indet. here.
Based on fieldwork in 1997, Munyikwa et al. (1998) initially reported
"Theropods are represented by several vertebrae, as well as numerous
shed teeth, the latter displaying relatively low, transversely broad
crowns." Woolley et al. (2015) note "Theropod remains, consisting
primarily of isolated teeth and vertebrae, suggest the presence of at
least one taxon of medium-bodied abelisauroid."
References-
Bond and Bromley, 1970. Sediments with the remains of dinosaurs near
Gokwe, Rhodesia. Palaeogeography, Palaeoclimatology, Palaeoecology.
8(4), 313-327.
Munyikwa, Sampson, Rogers, Forster, Curry and Curtice, 1998. Vertebrate
palaeontology and geology of the Gokwe Formation, Zimbabwe. Gondwana
10: Event Stratigraphy of Gondwana. Journal of African Earth Sciences.
27(1A), 142-143.
Woolley, Sertich, Forster, Munyikwa, Sampson, Curry Rogers
and Rogers, 2015. Titanosaurian and other vertebrate remains from the Cretaceous
Gokwe Formation, central Zimbabwe. Journal of Vertebrate Paleontology. Program
and Abstracts 2015, 241.
undescribed Abelisauridae (Rama, 1932)
Late Maastrichtian, Late Cretaceous
Kallamedu Formation, India
Material- cervical vertebra (76 mm) (Rama, 1932)
teeth (Goswami, Prasad, Benson, Verma and Flynn,
2012)
References- Rao, 1932. Reptilian vertebra from the south Indian Cretaceous. American Journal of Science. 5(141), 220-224.
Goswami, Prasad, Benson, Verma and Flynn, 2012. New vertebrates
from the Late Cretaceous Kallamedu Formation, Cauvery Basin, South India, including
a troodontid dinosaur, a gondwanatherian mammal, and a Simosuchus-like
notosuchian crocodyliform. Journal of Vertebrate Paleontology. Program and Abstracts
2012, 102.
unnamed Abelisauridae (Huene and Matley, 1933)
Late Maastrichtian, Late Cretaceous
Lameta Formation, India
Material- (AMNH 1733) premaxilla (Chatterjee, 1978)
(AMNH 1753) premaxilla, maxilla (Chatterjee, 1978)
(AMNH 1955) maxilla (Chatterjee, 1978)
(AMNH 1958) distal caudal vertebra (Chatterjee, 1978)
(AMNH 1960) anterior dentary tip (Chatterjee, 1978)
(AMNH 1960) proximal caudal neural arch (AMNH online)
(AMNH coll.) partial frontal, partial dentary (Dalman and Gishlick, 2011)
(GSI K19/579) tibia (600 mm) (Huene and Matley, 1933)
(GSI K 19/581) tibia (Huene and Matley, 1933)
(GSI K27/529; incorrectly labeled 527) dentary (260 mm) (Huene and Matley, 1933)
(GSI K27/532) distal caudal vertebra (110 mm) (Huene and Matley, 1933)
(GSI K27/538; incorrectly labeled 548) incomplete maxilla (Huene and Matley,
1933)
(GSI K27/539) metatarsal IV (250 mm) (Huene and Matley, 1933)
(GSI K27/544; incorrectly labeled 538) fragmentary maxilla (Huene and Matley,
1933)
(GSI K27/548) incomplete maxilla (~340 mm) (Huene and Matley, 1933)
(GSI K27/550) dentary (260 mm) (Huene and Matley, 1933)
(GSI K27/558; lost) femur (Huene and Matley, 1933)
(GSI K27/568; lost) tibia (540 mm) (Huene and Matley, 1933)
(GSI K27/570; lost) incomplete femur (~600 mm) (Huene and Matley, 1933)
(GSI K27/571) three sacral centra (centrum 70 mm) (Huene and Matley, 1933)
(GSI K27/575) squamosal fragment (Huene and Matley, 1933)
(GSI K27/577) jugal fragment (Huene and Matley, 1933)
(GSI K27/580) partial jugal (Huene and Matley, 1933)
(GSI K27/594) distal caudal vertebra (Huene and Matley, 1933)
(GSI K27/595; syntype of Coeluroides largus) proximal caudal vertebra
(Huene and Matley, 1933)
(GSI K27/596) caudal vertebra (Huene and Matley, 1933)
(GSI K27/618; lost ) distal femur (Huene and Matley, 1933)
(GSI K27/653) pedal phalanx III-2
(GSI K27/654) pedal phalanx II-1 (80 mm) (Huene and Matley, 1933)
(GSI K27/658) metatarsal III (254 mm) (Huene and Matley, 1933)
(GSI K27/705) distal caudal vertebra (Huene and Matley, 1933)
(GSI K27/709) dentary (Huene and Matley, 1933)
(GSI K27/710) premaxilla (80 mm) (Huene and Matley, 1933)
(ISI R 163) lacrimal, jugal, posterior dentary, angular (Chatterjee and Rudra,
1996)
six teeth (Mather and Srivastava, 1987)
Comments- Cranial elements showing external sculpturing, caudal vertebrae
resembling Majungasaurus, and robust hindlimb elements are here referred
to Abelisauridae. Jugal GSI K27/580 is
distinct from GSI K27/577 in the large posterior notch in the ascending process
(shared with Carnotaurus) and different pattern of rugosities. Sacral
vertebrae GSI K27/571 resemble Carnotaurus by having transversely narrow
centra with only slight intercentral expansion. Proximal caudal neural arch
AMNH 1960 resembles Majungasaurus more than Aucasaurus or Carnotaurus
in the laterally oriented transverse processes. It differs from Aucasaurus
and Carnotaurus in the lack of hyposphene-hypantrum articulations. AMNH
1958, GSI K27/532, 589 (though note Carrano et al., 2011 listed it as noasaurid),
594, 596 and 705 have low ridgelike transverse processes and short rounded prezygopophyses,
unlike noasaurids, but like Majungasaurus, Ornithomimoides, O? barasimlensis
and Dryptosauroides. This material is likely referrable to Rahiolisaurus, Rajasaurus,
Indosuchus and/or additional unrecognized Lameta abelisaurid taxa. Chatterjee
(1978) and Chatterjee and Rudra (1996) have referred some of it to Indosuchus,
but this cannot be confirmed until more material of the latter is discovered.
Dalman and Gishlick (2011) reexamined the AMNH material, noting some previously
unreported elements, and found that it resolves as a basal abelisaurid while
Indosuchus was carnotaurine. Mathur and Srivastava (1987) referred teeth
to Majungasaurus, but these are more likely to belong to a separate taxon,
as the teeth of the AMNH abelisaurid specimens strongly resemble those of Majungasaurus.
References- Huene and Matley, 1933. The Cretaceous Saurischia and Ornithischia
of the central provinces of India. Palaeontologica Indica. 21, 1-74.
Chatterjee, 1978. Indosuchus and Indosaurus, Cretaceous carnosaurs
from India. Journal of Paleontology. 52(3), 570-580.
Mathur and Srivastava, 1987. Dinosaur teeth from Lameta Group (Upper Cretaceous)
of Kheda District, Gujarat. Journal of the Geological Society of India. 29,
554-566.
Chatterjee and Rudra, 1996. KT events in India: Impact, rifing, volcanism and
dinosaur extinction. In Novas and Molnar (eds.). Proceedings of the Gondwanan
Dinosaur Symposium. Memiors of the Queensland Museum. 39(3), 489-532.
Novas, Agnolin and Bandyopadhyay, 2004. Cretaceous theropods from India: A review
of specimens described by Huene and Matley (1933). Revista del Museo Argentino
del Ciencias Naturales. 6(1), 67-103.
Dalman and Gishlick, 2011. Theropod material from Lameta, India, in the collection
of the American Museum of Natural History and its bearing on the diagnosis and
phylogenetic and taxonomic status of Indosuchus raptorius. Journal of
Vertebrate Paleontology. Program and Abstracts 2011, 95.
undescribed Abelisauridae (Gallina, Apesteguia, Haluza and Canale, 2014)
Late Berriasian-Valanginian, Early Cretaceous
Bajada Colorada Formation, Neuquen, Argentina
Material- (MMCh-PV-66) (~7-8 m) tibia
(MMCH-PV-68-1) tooth
Reference- 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.
undescribed abelisaurid (Canale, Carballido, Otero, Canudo and Garrido,
2014)
Albian-Cenomanian, Early Cretaceous-Late Cretaceous
Bayo Overo Member of the Cerro Barcino Formation, Chubut, Argentina
Material- tooth
Reference- Canale, Carballido, Otero, Canudo and Garrido, 2014. Carcharodontosaurid
teeth associated with titanosaur carcasses from the Early Cretaceous (Albian)
of the Chubut Group, Chubut Province, Patagonia, Argentina. Jornadas Argentinas
de Paleontologia de Vertebrados. Ameghiniana. 51(6) suplemento, 6.
undescribed abelisaurid (Fuentes, 2015)
Cenomanian-Early Coniacian, Late Cretaceous
Rio Neuquen Subgroup, Neuquen, Argentina
Material- (Museo Municipal "Argentino Urquiza" coll.) cranial
elements, vertebrae, cervical ribs, dorsal ribs, appendicular elements
Reference- Fuentes, 2015. Trabajos de limpieza y consolidacion de un
Abelisauridae procedente de Rincon de los Sauces, Neuquen. XXIX Jornadas Argentinas
de Paleontología de Vertebrados, resumenes. Ameghiniana. 52(4) suplemento,
18.
unnamed Abelisauridae (Lamanna, Martinez and Smith, 2002)
Mid Cenomanian-Turonian, Late Cretaceous
Lower Bajo Barreal Formation, Chubut, Argentina
Material- (MPM-99) incomplete tenth cervical vertebra, incomplete first
dorsal vertebra, mid dorsal centrum, ~eighth caudal vertebra, two mid caudal
vertebrae (Martinez, Novas and Ambrosio, 2004)
(UNPSJB-PV 221) tooth (14.0x10.8x4.9 mm) (Casal, Candeiro, Martinez, Ivany and Ibiricu, 2009)
(UNPSJB-PV 224) tooth (14.5x12.2x6.2 mm) (Casal, Candeiro, Martinez, Ivany and Ibiricu, 2009)
(UNPSJB-PV 225) tooth (12.9x9.9x4.8 mm) (Casal, Candeiro, Martinez, Ivany and Ibiricu, 2009)
(UNPSJB-PV 226) tooth (15.5x?x5.7 mm) (Casal, Candeiro, Martinez, Ivany and Ibiricu, 2009)
(UNPSJB-PV 229) tooth (22.3x12.5x7.1 mm) (Casal, Candeiro, Martinez, Ivany and Ibiricu, 2009)
(UNPSJB-PV 230) tooth (25.5x15.7x7.3 mm) (Casal, Candeiro, Martinez, Ivany and Ibiricu, 2009)
(UNPSJB-PV 231) tooth (22.2x11.9x6.3 mm) (Casal, Candeiro, Martinez, Ivany and Ibiricu, 2009)
(UNPSJB-PV 233) tooth (13.6x9.7x4.7 mm) (Casal, Candeiro, Martinez, Ivany and Ibiricu, 2009)
(UNPSJB-PV 247) (~8-10 m) maxilla (Lamanna, Martinez and Smith, 2002)
(UNPSJB-PV 981) tooth (13.4x9.1x4.6 mm) (Casal, Candeiro, Martinez, Ivany and Ibiricu, 2009)
(UNPSJB-PV 982) tooth (15.3x7.8x7.1 mm) (Casal, Candeiro, Martinez, Ivany and Ibiricu, 2009)
?(UNPSJB-PV 983) tooth (11.7x6.6x3.3 mm) (Casal, Candeiro, Martinez, Ivany and Ibiricu, 2009)
(UNPSJB-PV 984) tooth (11.9x9.0x3.6 mm) (Casal, Candeiro, Martinez, Ivany and Ibiricu, 2009)
(UNPSJB-PV 985) tooth (113.x7.6x4.0 mm) (Casal, Candeiro, Martinez, Ivany and Ibiricu, 2009)
(UNPSJB-PV 986) tooth (9.8x5.9x4.9 mm) (Casal, Candeiro, Martinez, Ivany and Ibiricu, 2009)
(UNPSJB-PV 1003) posterior skull, dentaries, teeth, two dorsal
ribs, partial sacrum, twenty-three caudal vertebrae, fifteen chevrons, scapula,
tarsus, incomplete pes including metatarsal III (Lamanna, Casal and Martinez, 2012)
(UNPSJB-PV 1067) (~6 m) tooth, mid-posterior cervical vertebra, dorsal
rib?, distal tibia (Ibiricu, Casal, Martinez, Alvarez and Poropat, 2020)
Comments- UNPSJB-PV 247 resembles Rugops very closely in the distribution
of external grooves and pits, elevated dental lamina, and fine striae on the
latter (Sereno et al., 2004). It may be referrable to Xenotarsosaurus,
from the same formation, though the latter was smaller and seemingly an adult.
Martinez et al. (2004) describe MPM-99 as an abelisaurid that
differs from both Carnotaurus and Majungasaurus but is more similar
to the former. Comparisons with Xenotarsosaurus from the same formation
are limited to the poorly preserved second dorsal of the latter, which differs
in minor ways that may be due to positional variation.
Casal
et al. (2009) refer multiple teeth to Abelisauroidea, with figured
specimens (except UNPSJB-PV 983) being abelisaurid based on the lack of
distal concavity.
Lamanna et al. (2012) referred UNPSJB-PV 1003 to basal Abelisauridae
based on- minimal cranial ornamentation; fenestra between lacrimal, postorbital
and frontal; dorsoventrally thin frontal; lack of awl-like, proximally-directed
projection at lateral end of proximal caudal transverse process; mediolaterally
slender metatarsal III. They note it cannot be compared to the contemporaneous
Xenotarsosaurus, but that the first caudal vertebra does differ from
MPM-99 from that formation. It may be diagnostic based on- rounded prominence bordered posteriorly
by mediolaterally-oriented groove on posterolateral part of dorsal surface of
frontal; low, acute posteromedial tuberosity on dorsal surface of frontal bordering
posterior median fossa; proximal caudal transverse processes abruptly expand
proximodistally well medial to their lateral extremes.
Ibiricu et al. (2020) refer UNPSJB-PV 1067 to Abelisauroidea indet.,
"although based on the low and subtriangular shaped facet for the
astragalar ascending process it is probably referable to Abelisauridae"
and distinguish it from Xenotarsosaurus
based on "the morphology of the facet for the reception of the
ascending process of the astragalus and the development and projection
of the medial malleolus."
References- Martinez, Maure, Oliva and Luna, 1993. Un maxilar de Theropoda
(Abelisauria) de la Formacion Bajo Barreal, Cretacico Tardio, Chubut, Argentina.
Ameghiniana. 30(1), 109-110.
Lamanna, Martinez and Smith, 2002. A definitive abelisaurid theropod dinosaur
from the early Late Cretaceous of Patagonia. Journal of Vertebrate Paleontology.
22(1), 58-69.
Martinez, Novas and Ambrosio, 2004. Abelisaurid remains (Theropoda,
Ceratosauria) from Southern Patagonia. Ameghiniana. 41(4), 577-585.
Sereno, Wilson and Conrad, 2004. New dinosaurs link southern landmasses in the
Mid-Cretaceous. Proceedings of the Royal Society of London B. 271(1546), 1325-1330.
Casal, Candeiro, Martinez, Ivany and Ibiricu, 2009. Theropod teeth
(Dinosauria: Saurischia) from the Bajo Barreal Formation, Upper
Cretaceous, Chubut Province, Argentina. Geobios. 42, 553-560.
Lamanna, Casal and Martinez, 2012. A new abelisaurid (Theropoda:
Ceratosauria) skeleton from the Upper Cretaceous Bajo Barreal Formation of Chubut
Province, Argentina. Journal of Vertebrate Paleontology. Program and Abstracts
2012, 124.
Ibiricu, Casal, Martinez, Alvarez and Poropat, 2020 (online 2019). New
materials and an overview of Cretaceous vertebrates from the Chubut
Group of the Golfo San Jorge Basin, central Patagonia, Argentina.
Journal of South American Earth Sciences. 98, 102460.
undescribed abelisaurid (Gonzalez Riga and Calvo, 1999)
Late Turonian-Early Coniacian, Late Cretaceous
Loma del Lindero, Portezuelo Formation of Rio Neuquen Subgroup, Neuquen, Argentina
Material- (MRS-PV coll.)
Comments- Gonzalez Riga and Calvo (1999) state "The Loma del
Lindero locality, close to Rincón de los Sauces city, has yielded 200
disarticulated bones belonging to two Titanosauridae specimens and one
Abelisauridae theropod." This material was excavated from 1998-2001,
with the titanosaur later being described as Muyelensaurus pecheni
(Calvo et al., 2007), and its description merely stating "At Loma del
Lindero locality, numerous titanosaur remains (~ 300 bones) associated
with a turtle specimen and theropods have been found." Note Loma del
Lindero ended up being in the Rio Neuquen Subgroup, not the Rio
Colorado as implied by Gonzalez Riga and Calvo.
References- Gonzalez Riga and Calvo, 1999. Unusual caudal series of Titanosauridae
of the Late Cretaceous in the Rio Colorado Formation, border between the Neuquen and Mendoza Provinces,
Argentina. VII International Symposium on Mesozoic Terrestrial Ecosystems, abstracts.
29-30.
Calvo, González Riga and Porfiri, 2007. A new titanosaur
sauropod from the Late Cretaceous of Neuquén, Patagonia, Argentina. Arquivos
do Museu Nacional, Rio de Janeiro. 65(4), 485-504.
unnamed abelisaurid (Novas and Bandyopadhyay, 2001)
Cenomanian-Early Coniacian, Late Cretaceous
Rio Limay Subgroup, Neuquen?, Argentina
Material- (MCA 56) cranial elements, distal caudal vertebrae, hindlimb
elements, pedal unguals IV (65 mm)
Comments- This has cranial sculpturing.
Reference- Novas and Bandyopadhyay, 2001. Abelisaurid pedal unguals from
the Late Cretaceous of India. VII International Symposium on Mesozoic Terrestrial
Ecosystems, Asociacion Paleontologica Argentina, Publicacion Especial 7. 145-149.
unnamed abelisaurid (Sereno, Martinez, Wilson, Varricchio, Alcober and Larsson, 2009)
Late Coniacian, Late Cretaceous
Plottier Formation of the Rio Neuquen Subgroup, Mendoza, Argentina
Holotype- (MCNA-PV-3137 in part; holotype of Aerosteon riocoloradensis in part) maxillary
or dentary tooth (33.4x16.5x8.4 mm)
Comments- Sereno et al. (2009) reported "an isolated crown of relatively small size was found at the site" of the Aerosteon
holotype and referred it to the taxon. Novas et al. (2013) stated
"this tooth is very different from that of the remaining megaraptorans
in being blade-like and transverselly compressed, and in having
chisel-like denticles of subequal length in both mesial and distal
carinae. Because its morphology is closer to an abelisaurid kind of
tooth, and the fact that it was not found in direct association with
cranial bones of Aerosteon,
we opt to consider the association of this isolated dental piece with
this taxon as dubious." Hendrickx et al. (2020) tested this and
recovered the tooth as an abelisaurid in cladistic analyses based on "a
mesial carina centrally positioned on the mesial surface of the crown
and extending close to the cervix, a weakly convex distal margin of the
crown, symmetrically convex labial and lingual surfaces of the crown,
apically inclined distal denticles, interdenticular sulci between both
mesial and distal denticles, and an irregularly textured surface of the
enamel."
References-
Sereno, Martinez, Wilson, Varricchio, Alcober and Larsson, 2009 (online 2008). Evidence for
avian intrathoracic air sacs in a new predatory dinosaur from Argentina. PLoS
ONE. 3(9), e3303.
Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution of the carnivorous
dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research.
45, 174-215.
Hendrickx, Tschopp and Ezcurra, 2020 (online 2019). Taxonomic
identification of isolated theropod teeth: The case of the shed tooth
crown associated with Aerosteon (Theropoda: Megaraptora) and the dentition of Abelisauridae. Cretaceous Research. 108, 104312.
unnamed Abelisauridae (Porfiri and Calvo, 2006)
Santonian, Late Cretaceous
Bajo de la Carpa Formation of the Rio Colorado Subgroup, Neuquen, Argentina
Material- (MAU-Pv-N-496/6) lateral tooth (?x5.5x2.2 mm) (Filippi, Martinelli
and Carrido, 2015)
(MAU-Pv-N-498) lateral tooth (21.9x9.6x4.3 mm) (Filippi, Martinelli and Carrido, 2015)
(MAU-Pv-N-505) anterior tooth (?x8.6x4.7 mm) (Filippi, Martinelli and Carrido, 2015)
(MAU-Pv-N-507) lateral tooth (?x?x3.3 mm) (Filippi, Martinelli and Carrido, 2015)
(MAU-Pv-N-508) lateral tooth (?x7.2x3.5 mm) (Filippi, Martinelli and Carrido, 2015)
(MAU-Pv-N-512) anterior tooth (?x8.9x4.3 mm) (Filippi, Martinelli and Carrido, 2015)
(MAU-Pv-LI-547) two teeth, hyoid fragment, incopmplete synsacrum,
several caudal transverse processes, incomplete furcula, ilia (one
incomplete; 510 mm), distal pubes (Mendez, Filippi and Garrido, 2015;
described in Méndez, Filippi, Gianechini and Juárez Valieri, 2018)
(MAU-Pv-LI-549) dorsal centrum (Mendez, Filippi and Garrido, 2015)
(MAU-Pv-LI-550) dorsal centrum (Mendez, Filippi and Garrido, 2015)
(MUCPv 1125) six sacral vertebrae, pelvises (Porfiri and Calvo, 2006)
Santonian, Late Cretaceous
Bajo de la Carpa Formation of the Rio Colorado Subgroup, Rio Negro, Argentina
(MACN-PV-RN 1012) incomplete ~fifth caudal vertebra (Ezcurra and Mendez, 2009)
(MPCA 249) tooth (~15.53x9.07x4.94 mm) (Gianechini, Lio and Apesteguía, 2011)
(MPCA 251) tooth (~20.79x~10.52x~5.68 mm) (Gianechini, Lio and Apesteguía, 2011)
Comments- Porfiri and Calvo (2006) briefly described a pelvis as a carnotaurine
based on- elongated and low ilium with straight dorsal border; acute ventral
notch on preacetabular process; posterior border of postacetabular process with
marked concavity. Yet their concept of Carnotaurinae differs from that used
here in including Genusaurus, Majungasaurus and Rajasaurus.
Ezcurra and Mendez (2009) described a caudal which they referred to Carnotaurinae
based on a character shared with Carnotaurus and Aucasaurus- transverse
processes oriented at angle of 45° or more with regard to horizontal axis
of the caudal.
Gianechini et al. (2011) describe two teeth with characaters that "are
similar to those present in abelisaurid teeth, considering principally
the straight distal border of the crown, the degree of labiolingual
compression, and the morphology of the denticles (chisel-like with
rounded apex) . The latter character allows extends tentatively the
assignment of MPCA 249 and 251 to Abelisauridae."
Filippi et al. (2015) described several teeth as Abelisauroidea
indet..
Mendez et al. (2015) mention a fragmentary skeleton which was later
described by Mendez et al. (2018). The latter assigned it to
Brachyrostra based on "a marked centrodiapophyseal lamina on the
ventral surface of the transverse processes in the anterior caudal
vertebrae."
References- Porfiri and Calvo, 2006. A new record of Carnotaurinae (Theropoda:
Abelisauridae) from the Upper Cretaceous of Neuquen, Patagonia. Journal of Vertebrate
Paleontology. 26(3), 111A-112A.
Ezcurra and Mendez, 2009. First report of a derived abelisaurid theropod from
the Bajo de la Carpa Formation (Late Cretaceous), Patagonia, Argentina. Bulletin
of Geosciences. 84(3), 547-554.
Gianechini, Lio and Apesteguía, 2011. Isolated archosaurian teeth from
"La Bonita" locality (Late Cretaceous, Santonian-Campanian), Río Negro
Province, Argentina. Historia Natural, tercera serie. 1, 5-16.
Filippi, Martinelli and Garrido, 2015. Una nueva asociación de dientes
vertebrados para la Formación Bajo de la Carpa (Santoniense, Cretácico
Superior) en Rincón de los Sauces, Neuquén, Argentina. Revista
Española de Paleontología. 30(2), 223-238.
Mendez, Filippi and Garrido, 2015. Nuevos hallazgos de dinosaurios teropodos
provenientes del sitto la invernada (Formacion Fajo de la Carpa), Rincon de
los Sauces, Neuquen. XXIX Jornadas Argentinas de Paleontología de Vertebrados,
resumenes. Ameghiniana. 52(4) suplemento, 28-29.
Méndez, Filippi, Gianechini and Juárez Valieri, 2018 (online 2017). New
brachyrostran remains (Theropoda, Abelisauridae) from La Invernada
fossil site (Bajo de la Carpa Formation, Upper Cretaceous), northern
Patagonia, Argentina. Cretaceous Research. 83, 120-126.
unnamed Abelisauridae (Coria and Arcucci, 2004)
Early Campanian, Late Cretaceous
Anacleto Formation of Rio Colorado Subgroup, Rio Negro, Argentina
Material- (MCF-PVPH-421) premaxillary tooth, six maxillary teeth (Coria and Arcucci, 2004)
(MPCN-PV 69) (~3-5 m) (subadult or adult) incomplete premaxilla, three
partial ?dorsal vertebrae, five partial fused sacral centra, proximal
humeri (~250 mm), distal pubis, incomplete pedal ungual, fragments
(Gianechini, Apesteguia, Landini, Finotti, Valieri and Zandonai, 201)
Comments- MCF-PVPH-421 are
teeth associated with the Auca Mahuevo titanosaur nesting site Layer
4. Coria and Arcucci (2004) described them as Theropoda indet.,
but noted a resemblence to Aucasaurus and concluded they were from
(translated) "an abelisaurian or other theropod with a dental
convergence with them."
References- Coria and Arcucci, 2004. Nuevos dinosaurios terópodos de Auca Mahuevo,
provincia del Neuquén (Cretácico tardío, Argentina). Ameghiniana. 41,
597-603.
Gianechini, Apesteguia, Landini, Finotti, Valieri and Zandonai,
2015. New abelisaurid remains from the Anacleto Formation (Upper Cretaceous),
Patagonia, Argentina. Cretaceous Research. 54, 1-16.
undescribed abelisaurid (Gasparini, Sterli, Parras, O'Gorman, Salgado,
Varela and Pol, 2015)
Maastrichtian, Late Cretaceous
La Colonia Formation, Chubut, Argentina
Material- (MPEF-PV 10826) specimen including maxilla, nasal, postorbital,
metatarsals including III, pedal phalanges, pedal unguals
Comments- Gasparini et al. (2015) referred this to Abelisauridae based
on- maxillary and nasal ornamentation; pedal ungual with lateral groove bifurcated
proximally; ventral side of pedal ungual with very conspicuous circular depression.
They distinguished it from the contemporaneous Carnotaurus based on-
ornamentation is slight elongated grooves; dorsal surface of postorbital anteroposteriorly
elongated and less curved.
Reference- Gasparini, Sterli, Parras, O'Gorman, Salgado, Varela and Pol,
2015. Late Cretaceous reptilian biota of the La Colonia Formation, central Patagonia,
Argentina: Occurrences, preservation and paleoenvironments. Cretaceous Research.
54, 154-168.
unnamed abelisaurid (Sales, Martinelli, Francischini, Rubert, Marconato, Soares and Schultz, 2017)
Santonian-Maastrichtian, Late Cretaceous
Morro do Cambambe, Bauru or Parecis Group, Brazil
Material- (UFRGS-PV-038-K) tooth (8.9x5.8x? mm)
Comments- This was discovered
in the 1990s. Sales et al. (2017) identified it as an
abelisaurid, finding it to be closest in morphology to Rugops, Kryptops and Majungasaurus in Hendrickx's dental analysis.
Reference- Sales, Martinelli,
Francischini, Rubert, Marconato, Soares and Schultz, 2017. New dinosaur
remains and the tetrapod fauna from the Upper Cretaceous of Mato Grosso
State, central Brazil. Historical Biology. 30(5), 661-676.
unnamed Abelisauridae (Bertini, 1996)
Late Santonian-Campanian, Late Cretaceous
Alfredo Marcondes, Adamantina Formation of the Bauru Group, Brazil
Material-
(UFRJ-DG 378-Rd) tooth (Candeiro, Martinelli, Avilla and Rich, 2006)
Late Santonian-Campanian, Late Cretaceous
Florida Paulista, Adamantina Formation of the Bauru Group, Brazil
(UFRJ-DG 371-Rd) tooth (Candeiro, Martinelli, Avilla and Rich, 2006)
(UFRJ-DG 374-Rd) tooth (Candeiro, Martinelli, Avilla and Rich, 2006)
Late Santonian-Campanian, Late Cretaceous
Prata, Adamantina Formation of the Bauru Group, Brazil
(MMR/UFU-PV 0006) tooth (Candeiro, Martinelli, Avilla and Rich,
2006)
Late Santonian-Campanian, Late Cretaceous
Santa Irene Farm, Adamantina Formation of the Bauru Group, Brazil
(MPMA-12-00D1-97) tooth (29.1x15x7 mm) (Tavares, Branco and Santucci, 2014)
(MPMA-12-00D3-97) tooth (28.2x11x5.5 mm) (Tavares, Branco and Santucci, 2014)
(MPMA-12-00D4-97) tooth (25x10.9x5.5 mm) (Tavares, Branco and Santucci, 2014)
(MPMA-12-00D5-97) tooth (26.2x11.6x5.7 mm) (Tavares, Branco and Santucci, 2014)
(MPMA-12-00D7-97) tooth (18.3x11.2x5.5mm) (Tavares, Branco and Santucci, 2014)
(MPMA-12-00D10-97) tooth (10.8x5.4x3.7 mm) (Tavares, Branco and Santucci, 2014)
(MPMA-12-00D11-97) tooth (15.7x7.5x4.5 mm) (Tavares, Branco and Santucci, 2014)
(MPMA-12-00D12-97) tooth (13.6x7.7x4.8 mm) (Tavares, Branco and Santucci, 2014)
?(MPMA-12-00D13-97) tooth (12.6x7.4x3 mm) (Tavares, Branco and Santucci, 2014)
?(MPMA-12-00D14-97) tooth (15.7x8.4x3.6 mm) (Tavares, Branco and Santucci, 2014)
Late Santonian-Campanian, Late Cretaceous
Santo Anastacio, Adamantina Formation of the Bauru Group, Brazil
(DGM 927-R) partial ilium (Brum, Machado, Campos and Kellner,
2016)
(MCT 1857-R) distal femur (~504 mm) (Brum, Machado, Campos and Kellner, 2016)
(URC 44-R) premaxilla, tooth (Bertini, 1996)
Comments- Tavares et al. (2014) described eight teeth as abelisaurid
and two as dromaeosaurid (MPMA-12-00D13-97 and 14-97), but in their morphometric
analyses, these were closer to each other than they were to sampled abelisaurids
and dromaeosaurids. This and the absence of any clear dromaeosaurid characters
(e.g. 8-shaped basal section, high DSDI) suggests all of the described teeth
may belong to one abelisaur taxon. Brum et al. (2021) found MPMA-12-00D13-97 and 14-97 fell out within Raptorex and closest to Guanlong respectively in their morphometric analysis.
References- Bertini, 1996. Evidencias de Abelisauridae (Carnosauria:
Saurischia) do Neocretaceo da Bacia do Parana. Boletim do 4o Simposio
sobre o Cretáceo do Brasil. 267-271.
Candeiro, Martinelli, Avilla and Rich, 2006. Tetrapods from
the Upper Cretaceous (Turonian-Maastrichtian) Bauru Group of Brazil: A reappraisal.
Cretaceous Research. 27(6), 923-946.
Candeiro, 2014. Descripcion de los caracteres de un premaxilar (Theropoda, Abelisauridae)
de la formacion Adamantina (Grupo Bauru), Sao Paulo, Brasil. Jornadas Argentinas
de Paleontologia de Vertebrados. Ameghiniana. 51(6) suplemento, 6-7.
Tavares, Branco and Santucci, 2014. Theropod teeth from the Adamantina Formation
(Bauru Group, Upper Cretaceous), Monte Alto, São Paulo, Brazil. Cretaceous
Research. 50, 59-71.
Brum, Machado, Campos and Kellner, 2016. Morphology and internal structure of
two new abelisaurid remains (Theropoda, Dinosauria) from the Adamantina Formation
(Turonian-Maastrichtian), Bauru Group, Paraná Basin, Brazil. Cretaceous
Research. 60, 287-296.
Brum, Pegas, Bandeira, Souza, Campos and Kellner, 2021. A new
unenlagiine (Theropoda, Dromaeosauridae) from the Upper Cretaceous of
Brazil. Papers in Palaeontology. 7(4), 2075-2099.
unnamed Abelisauridae (Candeiro, 2002)
Late Maastrichtian, Late Cretaceous
National Road BR-050, Serra da Galga Formation of the Bauru Group, Brazil
Material-
(CPP 893) (~4-5 m) incomplete ~fifth dorsal vertebra (80 mm) (Novas, Carvalho, Ribeiro
and Mendez, 2008)
Late Maastrichtian, Late Cretaceous
Peirópolis, Serra da Galga Formation of the Bauru Group, Brazil
(CPP 002, 020-021, 121, 123, 129b, 129c, 131, 132, 134-136, 144,
150, 154, 158, 161/1, 198, 205-207, 211, 242, 372, 375/2, 446, 451/1, 452/1,
463, 476-478) thirty-two teeth (Candeiro, 2002)
(MCT 1783-R) tibia (Machado, Campos and Kellner, 2011)
Late Maastrichtian, Late Cretaceous
Serra do Veadinho outcrop 1, Serra da Galga Formation of the Bauru Group, Brazil
?(CPP 692) pedal phalanx III-2 (60 mm) (Novas, Carvalho, Ribeiro and Mendez,
2008)
Late Maastrichtian, Late Cretaceous
Serra do Veadinho outcrop 2, Serra da Galga Formation of the Bauru Group, Brazil
(CPP 174) (~3-4 m) distal femur (~570 mm) (Novas, Carvalho,
Ribeiro and Mendez, 2008)
Comments- In 2021 the Serra da Galga and Ponte Alta Members of the Marilia Formation were recognized as the Serra da Galga Formation.
References- Candeiro, 2002. Dentes de Theropoda da Formacao Marylia (Santoniano-Maastrichtiano),
Bacia Bauru, Regiao de Peiropolis, Uberaba, Minas Gerais, Brasil. Masters thesis,
Universidade Federal do Rio de Janeiro, Rio de Janeiro. 136 pp.
Candeiro, Bergqvist, Novas and Currie, 2004. Theropod teeth from the Marilia
Formation (Upper Maastrichtian), Minas Gerais state, Brazil. Journal of Vertebrate
Paleontology. 24(3), 204A.
Candeiro, Martinelli, Avilla and Rich, 2006. Tetrapods from the Upper Cretaceous
(Turonian-Maastrichtian) Bauru Group of Brazil: A reappraisal. Cretaceous Research.
27(6), 923-946.
Novas, Carvalho, Ribeiro and Mendez, 2008. First abelisaurid
bone remains from the Maastrichtian Marília Formation, Bauru Basin, Brazil.
Cretaceous Research. 29(4), 625-635.
Machado, Campos and Kellner, 2011. A new abelisaurid tibia from the Late Cretaceous
of Brazil. Journal of Vertebrate Paleontology. Program and Abstracts 2011, 149.
Candeiro, Currie and Bergqvist, 2012. Theropod teeth from the Marília
Formation (Late Maastrichtian) at the paleontological site of Peirópolis
in Minas Gerais State, Brazil. Revista Brasileira de Geociências. 42(2),
323-330.
unnamed abelisaurid (Mendez, Novas and Iori, 2014)
Maastrichtian, Late Cretaceous
Sao Jose do Rio Preto Formation, Brazil
Material- (MPMA 08-0069-13) fibula (400 mm)
Reference- Mendez, Novas and Iori, 2014. New record of abelisauroid theropods
from the Bauru Group (Upper Cretaceous), Sao Paulo state, Brazil. Revista Brasileira
de Paleontologia. 17(1), 23-32.
unnamed Abelisauridae (Salinas, Marshall and Sepulveda, 1991)
Santonian-Maastrichtian, Late Cretaceous
Vinita Formation, Chile
Material- (SGO.PV.329c) two incomplete teeth
Comments- These were identified as coelurosaurian by Salinas et al. (1991),
but Soto-Acuna et al. (2015) reidentified them as abelisaurid.
References- Salinas, Marshall and Sepulveda, 1991. Vertebrados continentales
del Paleozoico y Mesozoico de Chile. Actas del VI Congreso Geológico
Chileno. 310-313.
Rubilar-Rogers, 2003. Registro de dinosaurios en Chile. Boletín Museo
Nacional de Historia Natural. 52, 137-150.
Rubilar-Rogers, Otero, Yury-Yanez, Vargas and Gutstein, 2012. An overview of
the dinosaur fossil record from Chile. Journal of South American Earth Sciences.
37, 242-255.
Soto-Acuna, Otero, Rubilar-Rogers and Vargas, 2015. Arcosaurios no avianos de
Chile. Publicacion Ocasional del Museo Nacional de Historia Natural, Chile.
63, 209-263.
unnamed Abelisauridae (Langston, 1953)
Maastrichtian, Late Cretaceous
Ortega, Columbia
Material- (UCMP 39649a) two incomplete lateral teeth
Comments- Initially identified as carnosaurian (Langston, 1953), these
were described as abelisaurid by Ezcurra (2009).
References- Langston, 1953. Cretaceous terrestrial vertebrates from Colombia,
South America. Bulletin of the Geological Society of America. 64, 1519.
Ezcurra, 2009. Theropod remains from the latest Cretaceous of Colombia and their
implications on the palaeozoogeography of western Gondwana. Cretaceous Research.
30, 1339-1344.
Chenanisaurus Longrich, Pereda-Suberbiola, Jalil, Khaldoune and Jourani, 2017
C. barbaricus Longrich, Pereda-Suberbiola, Jalil, Khaldoune and Jourani, 2017
Late Maastrichtian, Late Cretaceous
Couche III Phosphates, Morocco
Holotype- (OCP DEK-GE 772) (~7-8 m) anterior dentary (~430 mm)
Paratypes- ?(OCP DEK-GE 457) premaxillary tooth (44.1x20.6x14.3 mm)
?(OCP DEK-GE 458) premaxillary tooth (42.6x20x15.2 mm)
?(WDC-CCPM-005) maxillary tooth (37x20x10 mm) (Buffetaut, Escuillie and Pohl, 2005)
Diagnosis- (after Longrich et al., 2017) proportionately deep mandible;
dentary bowed in lateral view; lateral groove and associated foramina
located high on dentary (also in Genyodectes); anterodorsal margin of dentary downturned in
lateral view; massive symphysis, with anterior margin vertical in
lateral view; tip of jaws broad, with dentaries meeting at an obtuse
angle.
Comments- Discovered prior to
November 2016, Longrich et al. described the holotype mandible and
referred three teeth, one of which had been previously described as a
possible abelisaurid by Buffetaut et al. (2005). The authors
state "referral of the isolated teeth to
Chenanisaurus is made on the basis of provenance, size, and the general
resemblance of the teeth to those of the holotype."
Longrich et al. added Chenanisaurus to Tortosa et al.'s ceratosaur
analysis and recovered it in a trichotomy with Kryptops and Rugops plus
derived abelisaurids.
References- Buffetaut, Escuillie and Pohl, 2005. First theropod dinosaur
from the Maastrichtian phosphates of Morocco. Kaupia. 14, 3-8.
Longrich, Pereda-Suberbiola, Jalil, Khaldoune and Jourani, 2017. An
abelisaurid from the latest Cretaceous (Late Maastrichtian) of Morocco,
north Africa. Cretaceous Research. 76, 40-52.
Kryptops Sereno and Brusatte, 2008
K. palaios Sereno and Brusatte, 2008
Aptian-Albian, Early Cretaceous
Elrhaz Formation, Niger
Holotype- (MNN GAD1; in part) incomplete maxilla (~250 mm)
Diagnosis- (after Sereno and Brusatte, 2008) a deep secondary wall in
the anteroventral corner of the antorbital fossa that completely obscures the
antorbital fossa and that has a scalloped and fluted dorsal margin; external
texture on the maxilla, which is composed of short linear grooves.
Comments-
This was mentioned by Sereno et al. (2004) as an undescribed
abelisaurid from Gadoufaoua. It was later described in detail and named
by Sereno and Brusatte (2008). Carrano et al. (2012) noticed the
postcrania (MNN GAD1; in part- three dorsals, two ribs, sacrum and
pelves) was incongruous for an abelisaurid, found in situ 15 meters
away from the maxilla, and suggested it was carcharodontosaurid
instead. While traditionally a basal abelisaurid, in Farke and
Sertich's (2013) analysis it cannot be placed more exactly than the Rugops plus derived Abelisauridae clade using only the maxilla.
References- Sereno, Wilson and Conrad, 2004. New dinosaurs link southern
landmasses in the Mid-Cretaceous. Proceedings: Biological Sciences. 71(1546),
1325-1330.
Brusatte and Sereno, 2006. Basal abelisaurid and carcharodontosaurid theropods
from the Elrhaz Formation (Aptian-Albian) of Niger. Journal of Vertebrate Paleontology.
27(3), 46A.
Sereno and Brusatte, 2008. Basal abelisaurid and carcharodontosaurid theropods
from the Lower Cretaceous Elrhaz Formation of Niger. Acta Palaeontologica Polonica.
53(1), 15-46.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Farke and Sertich, 2013. An abelisauroid theropod dinosaur from the Turonian
of Madagascar. PLoS ONE. 8(4), e62047.
Spectrovenator Zaher, Pol, Navarro, Delcourt and Carvalho, 2020
S. ragei Zaher, Pol, Navarro, Delcourt and Carvalho, 2020
Barremian-Aptian, Early Cretaceous
Quirico Formation, Brazil
Holotype- (MZSP-PV 833) skull
(~254 mm), mandibles (260.98 mm), hyoid, anterior cervical vertebrae
including proatlas, atlanatal intercentrum, atlantal neurapophyses,
cervical ribs, posterior dorsal vertebrae, dorsal ribs, sacrum,
proximal caudal vertebrae, distal caudal vertebrae, chevrons, pelvis,
hindlimbs including femora, tibiae, fibulae, astragalcananea,
metatarsal I, metatarsal II, pmetatarsal III, metatarsal IV, pedal
phalanges including pedal ungual II, metatarsal V
Diagnosis- (after Zaher et al.,
2020) posterior ramus of maxilla ornamented with vertically oriented
grooves except for the smooth region anterior to the maxilla-jugal
suture; lacrimal lateral surface rugose except for smooth ventral
region; ventrally bowed posterior process of jugal; nuchal crest with
thin and smooth dorsal margin; straight ventral margin of dentaries
with deep sulcus on lateral surface; dorsal margin of surangular
slightly convex; longitudinal ridge along posteroventral end of
mandibular ramus.
Comments Discovered around 2010 beneath the holotype of Tapuiasaurus. Zaher et al. use Rauhut and Carrano's ceratosaur analysis to recover Spectrovenator as an abelisaurid more derived than Eoabelisaurus but diverging before Rugops.
Reference- Zaher, Pol, Navarro,
Delcourt and Carvalho, 2020. An Early Cretaceous theropod dinosaur from
Brazil sheds light on the cranial evolution of the Abelisauridae.
Comptes Rendus Palevol. 19(6), 101-115.
undescribed basal abelisaurid (Canale, Haluza and Novas, 2015)
Early Cenomanian, Late Cretaceous
Candelaros Formation of Rio Limay Group, Neuquen, Argentina
Material- (MMCh-PV 69) (14 year old adult, ~4 m, ~240 kg) incomplete
frontal, two dorsal rib fragments, two fused sacral centra, incomplete pelvis,
partial femur, distal metatarsal II, incomplete metatarsal III
Comments- Canale et al. (2016) added the specimen to a version of Carrano
and Sampson's ceratosaur matrix and found it to be sister to Majungasaurinae+Brachyrostra,
more closely related to the latter than Rugops.
References- Canale, Haluza and Novas, 2015. Histologia osea de un nuevo
ejempar de Abelisauridae (Dinosauria, Theropoda) del Cenomaniano de la Provincia
de Neuquen, Argentina. XXIX Jornadas Argentinas de Paleontología de Vertebrados,
resumenes. Ameghiniana. 52(4) suplemento, 10.
Canale, Cerda, Novas and Haluza, 2016. Small-sized abelisaurid (Theropoda: Ceratosauria)
remains from the Upper Cretaceous of northwest Patagonia, Argentina. Cretaceous
Research. 62, 18-28.
Kurupi Iori, Ismael de Araújo-Júnior, Simionato Tavares, da Silva Marinho and Martinelli, 2021
K. itaata Iori, Ismael de Araújo-Júnior, Simionato Tavares, da Silva Marinho and Martinelli, 2021
Maastrichtian, Late Cretaceous
Gaviões's Paleontological Site, Marilia Formation of the Bauru Group, Brazil
Holotype- (MPMA 27-0001-02) (~5 m adult) incomplete
first caudal vertebra (88 mm), partial fifth caudal vertebra,
incomplete seventh caudal vertebra (85 mm), partial ilium fused to
pubic fragment and incomplete ischia, two fragments
Diagnosis- (after Iori et al.,
2021) first caudal vertebra with anterolaterally projected
triangular process placed below and at mid-length of the anterior edge
of the transverse process; first caudal vertebra with a notch at the
anterodistal portion of the transverse process (possibly to receive the
iliac blade), between a shelf-like process and the anterodistal corner
of the transverse process; first caudal vertebra with a very concave
and broad anterior centrodiapophyseal fossa; proximal caudal vertebrae
(at least c1 and c7) with a finger-like process, anterodorsally
projected, located on the dorsal surface of the transverse process,
near the anterior base of the neural spine; increased inclination (from
~20 to ~35 degrees) of the transverse processes along the first third
of the caudal series; lack of infradiapophyseal fossa in the first
caudal; presence of reduced anterior and posterior centrodiapophyseal
laminae in the seventh caudal; lack of ischial tuberosity.
Comments- The pelvic elements
were found in 2002, with the other holotype material discovered in
2006, 2009, 2013 and 2014. Mendez et al. (2014) initially described the
pelvis as Abelisauroidea indet., but Iori et al. (2021)
described the caudals and named it as a new taxon Kurupi itaata.
They added it to Rauhut and Carrano's ceratosaur analysis and recovered
it in a polytomy with abelisaurids more derived than Spectrovenator.
References-
Mendez, Novas and Iori, 2014. New record of abelisauroid theropods from the
Bauru Group (Upper Cretaceous), Sao Paulo state, Brazil. Revista Brasileira
de Paleontologia. 17(1), 23-32.
Iori, Ismael de Araújo-Júnior, Simionato Tavares, da Silva Marinho and
Martinelli, 2021. New theropod dinosaur from the Late Cretaceous
of Brazil improves abelisaurid diversity. Journal of South American
Earth Sciences. 112(1), 103551.
Rugops Sereno, Wilson and Conrad, 2004
R. primus Sereno, Wilson and Conrad, 2004
Cenomanian, Late Cretaceous
Echkar Formation, Niger
Holotype- (MNN IGU1) partial skull including premaxilla, incomplete maxilla,
nasal, lacrimal, prefrontal, frontal, parietal, prootic and teeth
Diagnosis- (after Sereno et al., 2004) small fenestra in the skull roof
between the prefrontal, frontal, postorbital and lacrimal; row of seven small
invaginated depressions on the dorsal surface of each nasal.
Comments- This taxon is placed basal to other resolved abelisaurids (Majungasaurus, Rajasaurus, Abelisaurus, Carnotaurus) by Sereno et al.
(2004). It is very similar to an unnamed maxilla (UNPSJB-PV247) described by
Lamanna et al. (2002) from the Bajo Barreal Formation of Argentina. The nasal
fossae are intruded by vascular grooves, and may have served as a base for sensory
or display structures.
References- Lamanna, Martinez and Smith, 2002. A definitive abelisaurid
theropod dinosaur from the early Late Cretaceous of Patagonia. Journal of Vertebrate
Paleontology. 22(1), 58-69.
Sereno, Conrad and Wilson, 2002. Abelisaurid theropods from Africa: Phylogenetic
and biogeographic implications. Journal of Vertebrate Paleontology. 22(3), 106A.
Sereno, Wilson and Conrad, 2004. New dinosaurs link southern landmasses in the
Mid-Cretaceous. Proceedings: Biological Sciences. 271(1546), 1325-1330.
R. sp. (Sereno, Wilson and Conrad, 2004)
Cenomanian, Late Cretaceous
Kem Kem beds, Morocco
Material-
(MPUR NS 153/02) partial maxilla (Porchetti, Nicosia, Biava and Maganuco, 2011)
?(UCPC 10) partial maxilla (Mahler, 2005)
Comments- Sereno et al. (2004)
were the first to mention UCPC 10, stating "an abelisaurid maxilla, in
addition, has come to light recently from the same rocks in Morocco
(Mahler 2004)", citing an in press paper "Record of Abelisauridae
(Dinosauria: Therapoda) from Cenomanian Morocco." The latter was
published with a different title the next year, describing the maxilla
in detail as Abelisauridae gen. et sp. indet.. Smyth et al.
(2020) believed Mahler's characters "to distinguish UCPC 10 from Rugops
are equivocal" and "with limited differentiation of the material and
the extent of intraspecific variation in other abelisaurids" referred
it to Rugops sp..
Ibrahim et al. (2020) disagreed, as Bajo Barreal maxilla UNPSJB-PV 247
shares the same characters. Yet just as Kem Kem teeth are
referred to Carcharodontosaurus despite no known dental characters distinguishing it from Mapusaurus or Giganotosaurus, I support Smyth et al.'s referral to a coeval taxon here.
Porchetti et al. (2011) described two maxillae as Abelisauridae gen et
sp. indet., one of which was considered by Smyth et al. (2020) to be
"likely referable to Rugops
sp.." The other (MPUR NS 153/01) differs in several characters
which Porchetti et al. considered taxonomic, but Smyth et al. noted
some "such as simple, and reduced ornamentation; lack of alveolar
bulges and interalveolar depressions; and absence of a posterior
groove" and its small size could be explainable by ontogeny as
well. It is provisionally retained as Abelisauridae indet. here.
References- Sereno, Wilson and Conrad, 2004. New dinosaurs link southern
landmasses in the Mid-Cretaceous. Proceedings of the Royal Society of London
B. 271(1546), 1325-1330.
Mahler, 2005. Record of Abelisauridae (Dinosauria: Theropoda) from the Cenomanian
of Morocco. Journal of Vertebrate Paleontology. 25(1), 236-239.
Porchetti, Nicosia, Biava and Maganuco, 2011. New abelisaurid material from
the Upper Cretaceous (Cenomanian) of Morocco. Rivista Italiana di Paleontologia
e Stratigrafia. 117(3), 463-472.
Ibrahim, Sereno, Varricchio, Martill, Dutheil, Unwin, Baidder, Larsson,
Zouhri and Kaoukaya, 2020. Geology and paleontology of the Upper
Cretaceous Kem Kem Group of eastern Morocco. ZooKeys. 928, 1-216.
Smyth, Ibrahim, Kao and Martill, 2020 (online 2019). Abelisauroid
cervical vertebrae from the Cretaceous Kem Kem beds of southern Morocco
and a review of Kem Kem abelisauroids. Cretaceous Research. 108, 104330.
Abelisauridae sensu Novas, 1997
Definition- (Abelisaurus comahuensis + Carnotaurus sastrei
+ Indosaurus matleyi + Indosuchus raptorius + Majungasaurus
crenatissimus + Xenotarsosaurus bonapartei)
Tralkasaurus Cerroni, Motta, Agnolín, Aranciaga Rolando, Brissón Egli and Novas, 2020
= "Tralkasaurus" Cerroni, Motta, Agnolín, Aranciaga Rolando, Brissón Egli and Novas, 2019
T. cuyi Cerroni, Motta, Agnolín, Aranciaga Rolando, Brissón Egli and Novas, 2020
= "Tralkasaurus cuyi" Cerroni, Motta, Agnolín, Aranciaga Rolando, Brissón Egli and Novas, 2019
Middle Cenomanian-Early Turonian, Late Cretaceous
Huincul Formation of Rio Limay Subgroup, Rio Negro, Argentina
Holotype- (MPCA-Pv 815)
incomplete maxilla, cervical ribs, partial mid-posterior dorsal
vertebra, two mid-posterior dorsal transverse processes, two incomplete
sacral centra, two partial proximal caudal vertebrae, partial proximal
caudal transverse process, caudal centrum, fragmentary ilium, pubes
Diagnosis- (after Cerroni et
al., 2020) excavation of antorbital fossa widely exposed over entire
dorsal portion of maxillary body; internal and external margins of
antorbital fossa anterodorsally directed; dorsal parapophysis
dorsoventrally narrow and rod-like; proximally restricted dorsal
paradiapophyseal lamina.
Comments- This specimen was
announced by Motta et al. (2018) who stated it was recovered in "a
recent exploration", and referred to it contradictorally as
"Abelisauridae indet." and one of the specimens which "show distinct
autapomorphic traits and are different from previously known
theropods."
This paper was initially only available as a journal
pre-proof posted on November 30 2019 which included a blank section
under "Zoobank registration:", which according to ICZN Article 8.5.3 (a work must "be
registered in the Official Register of Zoological Nomenclature
(ZooBank) (see Article 78.2.4) and contain evidence in the work itself
that such registration has occurred"), made "Tralkasaurus cuyi" Cerroni et
al. 2019 a nomen nudum until the issue was published in March 2020.
Cerroni et al. added Tralkasaurus to the Tortosa et al. ceratosaur
matrix and recovered it in an abelisaurid polytomy with basal
majungasaurines (Arcovenator) and basal brachyrostrans (Xenotarsosaurus, Dahalokely, Rahiolisaurus, Ilokelesia).
References- Motta, Brissón
Egli, Aranciaga-Rolando, Rozadilla, Gentil, Lio, Cerroni, Garcia Marsà,
Agnolín, D´Angelo, Álvarez-Herrera, Alsina and Novas, 2018. New
vertebrate remains from the Huincul Formation (Cenomanian-Turonian;
Upper Cretaceous) in Río Negro, Argentina. XXXII Jornadas Argentinas de
Paleontologia de Vertebrados. R26.
Cerroni, Motta, Agnolín, Aranciaga Rolando, Brissón Egli and Novas,
2020. A new abelisaurid from the Huincul Formation
(Cenomanian-Turonian; Upper Cretaceous) of Río Negro province,
Argentina. Journal of South American Earth Sciences. 98, 102445.
unnamed Abelisauridae (Hendrickx and Mateus, 2014)
Late Kimmeridgian-Tithonian, Late Jurassic
Lourinha Formation, Portugal
Material- (ML 327) lateral tooth (31.8x20.1x10.7 mm)
(ML 966) lateral tooth (46.4x23.7x13 mm)
Reference- Hendrickx and Mateus, 2014. Abelisauridae (Dinosauria: Theropoda)
from the Late Jurassic of Portugal and dentition-based phylogeny as a contribution
for the identification of isolated theropod teeth. Zootaxa. 3759(1), 1-74.
Majungasaurinae Tortosa, Buffetaut, Vialle,
Dutour, Turini and Cheylan, 2014
Definition- (Majungasaurus crenatissimus <- Carnotaurus
sastrei) (after Tortosa, Buffetaut, Vialle, Dutour, Turini and Cheylan,
2014)
= "Rajasaurinae" Malkani, 2010
= "Majungasaurinae" Tortosa, Buffetaut, Vialle, Dutour, Turini and
Cheylan, 2013 online
= Majungasaurini Tortosa, Buffetaut, Vialle,
Dutour, Turini and Cheylan, 2014 vide Delcourt, 2018
Definition- (Majungasaurus crenatissimus <- Carnotaurus
sastrei) (modified after Delcourt, 2018)
Comments- Malkani (2010) proposed Rajasaurinae for Rajasaurus, apparently
within his new invalid family "Vitakrisauridae." However, it was not
given a diagnosis (ICZN Article 13.1.1) or explicitly stated to be new (ICZN
16.1), so is a nomen nudum. It could be used for the Rajasaurus+Majungasaurus+Indosaurus
clade that appears in some recent analyses. Tortosa et al. (2014) also
found this group, and proposed to name it Majungasaurinae.
Delcourt (2018) suggested Majungasaurinae "should be considered a tribe
and called Majungasaurini because is inserted in the subfamily
Carnotaurinae" in Wang et al.'s topology, as the latter has Majungasaurus closer to Carnotaurus than Abelisaurus. If this prooves to be a consensus topology, that might have merit.
References- Malkani, 2010. New Pakisaurus (Pakisauridae, Titanosauria,
Sauropoda) remains, and Cretaceous Tertiary (K-T) boundary from Pakistan. Sindh
University Research Journal (Science Series). 42(1), 39-64.
Tortosa, Buffetaut, Vialle, Dutour, Turini and Cheylan, 2014. A new abelisaurid
dinosaur from the Late Cretaceous of southern France: Palaeobiogeographical
implications. Annales de Paléontologie. 100(1), 63-86.
Delcourt, 2018. Ceratosaur palaeobiology: New insights on evolution and
ecology of the southern rulers. Scientific Reports. 8:9730.
Arcovenator Tortosa, Buffetaut,
Vialle, Dutour, Turini and Cheylan, 2014
= "Arcovenator" Tortosa, Buffetaut, Vialle, Dutour, Turini and
Cheylan, 2013 online
A. escotae Tortosa, Buffetaut, Vialle, Dutour, Turini and Cheylan,
2014
= "Arcovenator escotae" Tortosa, Buffetaut, Vialle, Dutour, Turini
and Cheylan, 2013 online
Late Campanian, Late Cretaceous
Lower Argiles Rutilantes Formation, Var, France
Holotype- (MHNA-PV-2011.12.1) postorbital, braincase
....(MHNA-PV-2011.12.2) squamosal
....(MHNA-PV-2011.12.3) tibia (510 mm)
....(MHNA-PV-2011.12.4) fibula (490 mm)
....(MHNA-PV-2011.12.5) proximal caudal vertebra
....(MHNA-PV-2011.12.15) tooth
Paratypes- (MHNA-PV-2011.12.20) tooth
(MHNA-PV-2011.12.187) tooth
(MHNA-PV-2011.12.198) proximal caudal vertebra
(MHNA-PV-2011.12.213) proximal caudal vertebra
(MHNA-PV-2011.12.297) tooth
Diagnosis- (after Tortosa et al., in press) frontals with slight median
thickening; median dorsal foramen between frontals; low depression with small
fenestra in skull roof between frontal, postorbital and lacrimal; postorbital
forming strong, rugose, dorsolaterally developed supraorbital brow; thin bony
expansion between ventral and posterior rami of postorbital; lateral tuberosity
on ventral ramus of postorbital; paroccipital processes bounded dorsally and
ventrally by two horizontal bars, forming depression lateral to foramen magnum;
basipterygoid process laterally directed; slightly proximally anterodorsally
curved fibular crest; lateral tibial condyle more developed posteriorly than
medial condyle.
Comments- The type materials were discovered in 2011 and described by
Tortosa et al. (2014) as a new taxon of abelisaurid. Their phylogenetic analysis
is shown as recovering Arcovenator in a clade with Majungasaurus,
Indosaurus, Rajasaurus and Rahiolisaurus which they name Majungasaurinae, yet the matrix is misprinted and when fixed actually
finds the new genus as a majungasaurine or a basal abelisaurid.
Although electronic publication of names was allowed by the ICZN in 2013, Tortosa
et al. did not register their name with Zoobank (ICZN 8.5.3), making it unofficial
until physical published in 2014.
Reference- Tortosa, Buffetaut, Vialle, Dutour, Turini and Cheylan, 2014 (online 2013).
A new abelisaurid dinosaur from the Late Cretaceous of southern France: Palaeobiogeographical
implications. Annales de Paléontologie. 100(1), 63-86.
unnamed majungasaurine (Allain and Pereda Suberbiola, 2003)
Late Campanian, Late Cretaceous
Begudian beds, Bouches-du-Rhone, France
Material- (La Boucharde taxon) tibia (~400 mm)
Comments- Assigned to Neoceratosauria indet. by Allain and Suberbiola
(2003), this was identified as abelisaurid by Carrano and Sampson (2008), which
was confirmed in the ceratosaur analysis of Tortosa et al. (2014). Most recently,
Filippi et al. (2016) used a version of that matrix to recover it as a majungasaurine
sister to Arcovenator.
References- Allain and Pereda Suberbiola, 2003. Dinosaurs of France.
Comptes Rendus Palevol. 2, 27-44.
Carrano and Sampson, 2008 (online 2007). The Phylogeny of Ceratosauria (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 6, 183-236.
Tortosa, Buffetaut, Vialle, Dutour, Turini and Cheylan, 2014. A new abelisaurid
dinosaur from the Late Cretaceous of southern France: Palaeobiogeographical
implications. Annales de Paléontologie. 100(1), 63-86.
Filippi, Mendez, Juarez Valieri and Garrido, 2016. A new brachyrostran with
hypertrophied axial structures reveals an unexpected radiation of latest Cretaceous
abelisaurids. Cretaceous Research. 61, 209-219.
Genusaurus Accarie, Beaudoin,
Dejax, Fries, Michard and Taquet, 1995
G. sisteronis Accarie, Beaudoin, Dejax, Fries, Michard and Taquet,
1995
Middle Albian, Early Cretaceous
Bevons Beds, France
Holotype- (MNHN, Bev.1) (3.16 m) seven dorsal centra, sacral centrum,
incomplete ilium, proximal pubis, femur (380 mm), proximal tibia, proximal fibula,
tarsal
Comments- This has been recently recovered as both a noasaurid and an
abelisaurid.
Reference- Accarie, Beaudoin, Dejax, Fries, Michard and Taquet, 1995.
Découverte d'un Dinosaure théropode nouveau (Genusaurus sisteronis
n. g., n. sp.) dans l'Albien marin de Sisteron (Alpes de Haute-Provence, France)
et extension au Crétacé inférieur de la lignée cératosaurienne.
Comptes Rendus de l'Académie des Sciences à Paris, série
IIa. 320, 327-344.
Indosaurus Huene, 1931 vide Huene,
1932
I. matleyi Huene, 1931 vide Huene, 1932
Maastrichtian, Late Cretaceous
Lameta Formation, India
Holotype- (GSI K27/565) frontals, parietals, braincase
Diagnosis- (after Novas et al., 2004) Provisionally indeterminate relative
to Indosuchus raptorius (pending publication of Dalman and Gishlick's
2011 analysis).
Comments- Though traditionally thought to be more Allosaurus-
or Carnotaurus-like than the tyrannosaurid- or Abelisaurus-like
Indosuchus, Novas et al. (2004) found that poor preservation made the
two taxa impossible to distinguish. In particular, there is no evidence of frontal
horns or sagittal crest shape differences. Other minor differences may be individual
variation, based on the high amount observed in Majungasaurus. Dalman
and Gishlick (2011) however noted in an abstract that the genera resolved in
different positions in their phylogenetic analysis.
References- Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre
Entwicklung und Geschichte. Monographien zur Geologie und Palaeontologie. 4(1),
viii + 361 pp.
Huene and Matley, 1933. The Cretaceous Saurischia and Ornithischia of the central
provinces of India. Palaeontologia Indica. 21, 1-74.
Novas and Bandyopaphyay, 1999. New approaches on the Cretaceous theropods from
India. VII International Symposium on Mesozoic Terrestrial Ecosystems, abstracts.
46-47.
Novas, Agnolin and Bandyopadhyay, 2004. Cretaceous theropods from India: A review
of specimens described by Huene and Matley (1933). Revista del Museo Argentino
del Ciencias Naturales. 6(1), 67-103.
Dalman and Gishlick, 2011. Theropod material from Lameta, India, in the collection
of the American Museum of Natural History and its bearing on the diagnosis and
phylogenetic and taxonomic status of Indosuchus raptorius. Journal of
Vertebrate Paleontology. Program and Abstracts 2011, 95.
Indosuchus Huene, 1931 vide
Huene, 1932
I. raptorius Huene, 1931 vide Huene, 1932
Maastrichtian, Late Cretaceous
Lameta Formation, India
Lectotype- (GSI K27/685) frontals, parietals,
Paralectotypes- (GSI K20/350) (subadult?) partial lacrimals, frontals,
parietals
(GSI K20/690) skull roof
Diagnosis- (after Novas et al., 2004) frontonasal suture placed anteriorly
compared to lacrimal.
References- Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre
Entwicklung und Geschichte. Monographien zur Geologie und Palaeontologie. 4(1),
viii + 361 pp.
Huene and Matley, 1933. The Cretaceous Saurischia and Ornithischia of the central
provinces of India. Palaeontologia Indica. 21, 1-74.
Novas, Agnolin and Bandyopadhyay, 2004. Cretaceous theropods from India: A review
of specimens described by Huene and Matley (1933). Revista del Museo Argentino
del Ciencias Naturales. 6(1), 67-103.
Dalman and Gishlick, 2011. Theropod material from Lameta, India, in the collection
of the American Museum of Natural History and its bearing on the diagnosis and
phylogenetic and taxonomic status of Indosuchus raptorius. Journal of
Vertebrate Paleontology. Program and Abstracts 2011, 95.
Majungasaurus Lavocat,
1955
= Majungatholus Sues and Taquet, 1979
M. crenatissimus (Deperet, 1896) Lavocat, 1955
= Megalosaurus crenatissimus Deperet, 1896
= Majungatholus atopus Sues and Taquet, 1979
Maastrichtian, Late Cretaceous
Maevarano Formation, Madagascar
Neotype- (MNHN.MAJ 1) (subadult) incomplete dentary, two teeth (Lavocat,
1955)
Referred- ?(FSL 92.289; syntype of Megalosaurus crenatissimus)
incomplete caudal vertebra (Deperet, 1896)
?(FSL 92.290; syntype of Megalosaurus crenatissimus) partial pedal ungual
(Deperet, 1896)
(FSL 92.306a; syntype of Megalosaurus crenatissimus) posterior dentary
tooth (Deperet, 1896)
(FSL 92.306b; syntype of Megalosaurus crenatissimus) fourth premaxillary
tooth (Deperet, 1896)
?(FSL 92.343; syntype of Megalosaurus crenatissimus) sacral centrum (Deperet,
1896)
?(FSL 92.???; syntype of Megalosaurus crenatissimus) sacral centrum (Deperet,
1896)
(MNHN.MAJ 4; holotype of Majungatholus atopus) lacrimal fragment, frontals,
parietals, mesethmoid, laterosphenoids (Sues and Taquet, 1979)
(MNHN 1911a-d) anterior premaxillary tooth, middle maxillary or dentary tooth,
posterior maxillary or dentary tooth, posterior dentary tooth (Piveteau, 1926)
?(MNHN coll.; lost?) vertebra, limb bone fragments (Boule, 1896)
?(MNHN coll.; lost) teeth (Boule, 1900)
?(MNHN coll.; lost) tooth, vertebrae including a caudal vertebra, fragmentary
limb bones (Thevenin, 1906)
?(MNHN coll.; lost) teeth, bones (Lavocat, 1955)
(UA Bv 532) pedal ungual I (Krause, Sampson, Carrano and O'Connor, 2007)
(UA Bv 1658) pedal ungual I (Krause, Sampson, Carrano and O'Connor, 2007)
(UA Bv 1260) pedal phalanx II-1 (Krause, Sampson, Carrano and O'Connor, 2007)
(UA Bv 1265) pedal phalanx III-1 (Krause, Sampson, Carrano and O'Connor, 2007)
(UA coll.) numerous teeth, three fragmentary femora or tibiae (Ravoavy, 1991)
Early Maastrichtian(?), Late Cretaceous
Masorobe Member of Maevarano Formation, Madagascar
Referred- teeth (Krause, Sampson, Carrano and O'Connor, 2007)
Middle Maastrichtian, Late Cretaceous
Anembalemba Member of Maevarano Formation, Madagascar
Referred- (FMNH PR 2008) premaxilla (72 mm) (Sampson, Krause, Dodson
and Forster, 1996)
(FMNH PR 2099) (subadult) partial skull (Sampson, Witmer, Forster, Krause, O'Connor,
Dodson and Ravoavy, 1998)
(FMNH PR 2100) incomplete skull (570 mm), mandibles, fifth caudal vertebra (84.9
mm), sixth caudal vertebra (95.7 mm), seventh caudal vertebra (98.1 mm), eighth
caudal vertebra (98.1 mm), ninth caudal vertebra (97 mm), tenth caudal vertebra
(~93.1 mm), eleventh caudal vertebra (96 mm), twelfth caudal vertebra (96.7
mm), thirteenth caudal vertebra (95.6 mm), fourteenth caudal vertebra (94.2
mm), fifteenth caudal vertebra (91.4 mm), sixteenth caudal vertebra (91.5 mm),
seventeenth caudal vertebra (90.4 mm), eighteenth caudal vertebra (91.1 mm),
nineteenth caudal vertebra (90.9 mm), twentieth caudal vertebra (88.7 mm), twenty-first
caudal vertebra (86.6 mm), twenty-second caudal vertebra (83.5 mm), twenty-third
caudal vertebra (81.6 mm), twenty-fourth caudal vertebra (79.2 mm), twenty-fifth
caudal vertebra (79 mm), twenty-sixth caudal vertebra (76.6 mm), twenty-seventh
caudal vertebra (74.6 mm), twenty-eighth caudal vertebra (70.8 mm), twenty-ninth
caudal vertebra (67.5 mm), distal caudal vertebra (36.5 mm), eighteen chevrons
(Sampson, Witmer, Forster, Krause, O'Connor, Dodson and Ravoavy, 1998)
....(UA 9089) four proximal caudal vertebrae (Ravoavy, 1991)
(FMNH PR 2198) (juvenile) lateral tooth (Carrano et al., 2002)
(FMNH PR 2226) (juvenile) anterior tooth (Carrano et al., 2002)
(FMNH PR 2228) (juvenile) lateral tooth (Carrano et al., 2002)
(FMNH PR 2278) (adult) premaxillae, maxillae, jugal, quadratojugal, ectopterygoid,
quadrate, surangular, angular, prearticular, articular, two fragmentary cervical
neural arches, three fragmentary dorsal vertebrae, one fragmentary caudal vertebra,
scapulocoracoid, partial ilium, femur, tibiae (one fragmentary), fibulae (one
partial; ~406 mm), astragalocalcaneum, metatarsal II (198.9 mm), phalanx II-1,
metatarsal III (250 mm), metatarsal IV (207.7 mm), phalanx IV-2, phalanx IV-3
(Krause, Sampson, Carrano and O'Connor, 2007)
(FMNH PR 2293) axis (Krause, Sampson, Carrano and O'Connor, 2007)
(FMNH PR 2294) five caudal vertebrae, two chevrons (Krause, Sampson, Carrano
and O'Connor, 2007)
(FMNH PR 2295) third cervical vertebra (Krause, Sampson, Carrano and O'Connor,
2007)
(FMNH PR 2423) humerus (Krause, Sampson, Carrano and O'Connor, 2007)
(FMNH PR 2424) tibia (Krause, Sampson, Carrano and O'Connor, 2007)
(FMNH PR 2425) astragalocalcaneum (Krause, Sampson, Carrano and O'Connor, 2007)
(FMNH PR 2426) pedal phalanx II-1 (Krause, Sampson, Carrano and O'Connor, 2007)
(FMNH PR 2427) pedal phalanx II-2 (Krause, Sampson, Carrano and O'Connor, 2007)
(FMNH PR 2428) distal pedal phalanx II-1, pedal ungual II, phalanx III-2 (Krause,
Sampson, Carrano and O'Connor, 2007)
(FMNH PR 2429) pedal phalanx III-1 (Krause, Sampson, Carrano and O'Connor, 2007)
(FMNH PR 2430) pedal phalanx IV-1 (Krause, Sampson, Carrano and O'Connor, 2007)
(FMNH PR 2431) pedal phalanx IV-3 (Krause, Sampson, Carrano and O'Connor, 2007)
(FMNH PR 2432) pedal phalanx IV-4 (Krause, Sampson, Carrano and O'Connor, 2007)
(FMNH PR 2433) pedal phalanx IV-4 (Krause, Sampson, Carrano and O'Connor, 2007)
(FMNH PR 2434) pedal ungual IV (Krause, Sampson, Carrano and O'Connor, 2007)
(MSNM V3342) lateral tooth (Fanti and Therrien, 2007)
(MSNM V3360) anterolateral tooth (Fanti and Therrien, 2007)
(MSNM V3363) anterolateral tooth (Fanti and Therrien, 2007)
(MSNM V3368) anterolateral tooth (Fanti and Therrien, 2007)
(MSNM V5255) pedal phalanx III-1 (60 mm) (Maganuco et al., 2008)
(MSNM V5256) pedal phalanx III-2 (36 mm) (Maganuco et al., 2008)
(MSNM V5267) pedal ungual II (42 mm) (Maganuco et al., 2008)
(MSNM V5276) pedal ungual IV (52 mm) (Maganuco et al., 2008)
(MSNM V5368) anterior premaxillary tooth (Fanti and Therrien, 2007)
(MSNM V5509) pedal ungual IV (51 mm) (Maganuco et al., 2008)
(MSNM V5510) pedal phalanx III-1 (74 mm) (Maganuco et al., 2008)
(MSNM V5518) anterolateral tooth (Fanti and Therrien, 2007)
(MSNM V6418) pedal ungual III (~57 mm) (Maganuco et al., 2008)
(MSNM V6419) pedal ungual I (29 mm) (Maganuco et al., 2008)
(MSNM V6420) pedal ungual IV (61 mm) (Maganuco et al., 2008)
(MSNM V6421) pedal ungual II (56 mm) (Maganuco et al., 2008)
(MSNM V coll.) 123 lateral teeth (Fanti and Therrien, 2007)
(MSNM V coll.) 21 anterolateral teeth (16-35 mm) (Fanti and Therrien, 2007)
(MSNM V coll.) 19 anterior premaxillary teeth (15-30 mm) (Fanti and Therrien,
2007)
(UA 8678) (subadult) incomplete skull, splenial, prearticular, surangular, atlantal
neurapophysis, axis (69.8 mm), third cervical vertebra (59.9 mm), fourth cervical
vertebra (61.7 mm), fifth cervical vertebra (63.2 mm), sixth cervical vertebra
(~65.5 mm), seventh cervical vertebra (~60.1 mm), eighth cervical vertebra (~56.1
mm), ninth cervical vertebra (57.9 mm), tenth cervical vertebra (58.4 mm), thirteen
cervical ribs, first dorsal vertebra (59.3 mm), second dorsal vertebra (~51.9
mm), third dorsal vertebra (55.2 mm), fourth dorsal vertebra (~51.6 mm), fifth
dorsal vertebra (58.9 mm), sixth dorsal vertebra (59.7 mm), seventh dorsal neural
arch, eighth dorsal vertebra (67.4 mm), ninth dorsal vertebra (67.1 mm), tenth
dorsal neural arch, eleventh dorsal vertebra (65.6 mm), twelfth dorsal vertebra,
thirteenth dorsal neural arch, fourteen dorsal ribs, second sacral vertebra,
third sacral vertebra, fourth sacral neural arch, first caudal vertebra (71.3
mm), second caudal vertebra (69.6 mm), third caudal vertebra (70.8 mm), fourth
caudal vertebra (73.2 mm), fifth caudal vertebra (75.2 mm), mid caudal vertebra,
first chevron, ilia (585 mm) (Sampson, Witmer, Forster, Krause, O'Connor, Dodson
and Ravoavy, 1998)
(UA 8709) incomplete skull, mandibles (Krause, Sampson, Carrano and O'Connor,
2007)
(UA 8716) premaxilla (Krause, Sampson, Carrano and O'Connor, 2007)
(UA 8717) premaxillae (Krause, Sampson, Carrano and O'Connor, 2007)
(UA 8718) partial lacrimal (Krause, Sampson, Carrano and O'Connor, 2007)
(UA 8719) partial skull roof (Krause, Sampson, Carrano and O'Connor, 2007)
(UA 8782) distal quadrate (Krause, Sampson, Carrano and O'Connor, 2007)
(UA 9031) humerus (Krause, Sampson, Carrano and O'Connor, 2007)
(UA 9032) tibia (Krause, Sampson, Carrano and O'Connor, 2007)
(UA 9033) astragalocalcaneum (Krause, Sampson, Carrano and O'Connor, 2007)
(UA 9034) metatarsal II (Krause, Sampson, Carrano and O'Connor, 2007)
(UA 9035) metatarsal IV (Krause, Sampson, Carrano and O'Connor, 2007)
(UA 9036) pedal phalanx II-1 (88 mm) (Krause, Sampson, Carrano and O'Connor,
2007)
(UA 9037) pedal phalanx II-2 (45 mm) (Krause, Sampson, Carrano and O'Connor,
2007)
(UA 9038) pedal ungual II (Krause, Sampson, Carrano and O'Connor, 2007)
(UA 9039) pedal phalanx III-1 (Krause, Sampson, Carrano and O'Connor, 2007)
(UA 9040) pedal phalanx IV-1 (Krause, Sampson, Carrano and O'Connor, 2007)
(UA 9041) pedal phalanx IV-2 (Krause, Sampson, Carrano and O'Connor, 2007)
(UA 9042) pedal phalanx III-2 (Krause, Sampson, Carrano and O'Connor, 2007)
(UA 9043) pedal ungual IV (Krause, Sampson, Carrano and O'Connor, 2007)
(UA 9077) tibia, fibula (406.4 mm) (Krause, Sampson, Carrano and O'Connor, 2007)
(UA 9078) fibula (Krause, Sampson, Carrano and O'Connor, 2007)
(UA 9079) metatarsal III (Krause, Sampson, Carrano and O'Connor, 2007)
(UA 9081) pedal phalanx III-1 or III-2 (Krause, Sampson, Carrano and O'Connor,
2007)
(UA 9082) astragalus (Krause, Sampson, Carrano and O'Connor, 2007)
(UA coll.) gastralia (O'Connor, 2007)
(FMNH and UA coll.) thousands of isolated teeth (Krause, Sampson, Carrano and
O'Connor, 2007)
specimen including scapulocoracoid, humerus, radius, ulna, metacarpal I, phalanx
I-1, metacarpal II, phalanx II-1, phalanx II-2, metacarpal III, phalanx III-1
fused with III-2, metacarpal IV fused with phalanx IV-1 (Burch and Carrano,
2008)
Late Maastrichtian, Late Cretaceous
Miadana Member of Maevarano Formation, Madagascar
Referred- teeth (Krause, Sampson, Carrano and O'Connor, 2007)
Late Cretaceous(?)
Madagascar
Referred- ?(MNHN.MAJ 242) premaxillary tooth (Smith, 2007)
Diagnosis- (after Krause et al., 2007) dorsoventrally deep, fused nasals
that are strongly pneumatized via large, bilateral foramina; nasal processes
of left and right premaxillae separated by a thin lamina of nasal; maxilla bearing
17 alveoli; frontals rounded rostrally rather than forming a double notch; frontals
with a sculptured, median cornual process in adults that is variably pneumatic;
pronounced median fossa on sagittal (frontoparietal) crest; dentary bearing
17 alveoli, virtually no extension caudal to last alveolus, and ventral position
of lateral sulcus; teeth bearing weakly developed interdenticular sulci; long,
falciform atlantal epipophysis; vertebral centra in cranial dorsal series with
dorsoventrally elongate articular surfaces; cranial and caudal borders of midcervical
transverse processes parallel in lateral view; dorsal and caudal vertebral neural
spines dorsally expanded (transversely and craniocaudally); cervical ribs pneumatized
via multiple, enlarged foramina on medial surface of shaft, and accessory foramina
on cranial and caudal surfaces of capitulotubercular web.
Comments- Deperet (1986) never specified a holotype from his syntypes
for Megalosaurus crenatissimus, which came from different localities.
Lavocat (1955) described a dentary (MNHN.MAJ 1) he believed to be from the same
species, but which he thought indicated generic difference from Megalosaurus.
He designated this dentary the type specimen of Majungasaurus, making
the new combination Majungasaurus crenatissimus. This was an improper
procedure, as Deperet's syntypes remain the type specimens of crenatissimus
regardless of which genus they are assigned to. Sampson et al. (1996) and Krause
et al. (2007) claimed Lavocat implicitly designated the dentary the neotype
of Majungasaurus crenatissimus, but the ICZN does not allow neotype designation
without petition unless the type material is lost (Article 75). So Sampson et
al.'s and Krause et al.'s use of the dentary as the type specimen for Majungasaurus
crenatissimus was incorrect. Subsequently, Carrano et al. (2009) petitioned
the ICZN to designate the dentary as the neotype because "the original
Depret type specimens are indeed indeterminate as to genus and species",
oddly citing Krause et al. who stated the two syntype teeth (FSL 92.306a-b)
are identical to those in the dentary, and that Majungasaurus teeth are
diagnostic among abelisaurids. The ICZN decided the case in 2011, declaring
the dentary to be the neotype, despite the fact unknown to them that two of
the syntypes are seemingly considered to be diagnostic by the petitioners themselves.
Smith (2007) also found one of the syntype teeth to be confidently referrable
to Majungasaurus.
An additional nomenclatural issue involves the holotype of Majungatholus
atopus, which was originally identified as a pachycephalosaur (Sues and
Taquet, 1979). Though a few authors expressed doubt at this identification (Rage,
1988; Giffen, 1989), it was not disproven until 1996 when the skull of FMNH
PR 2100 was found (Sampson et al., 1998), showing a Majungatholus dome
on an abelisaurid skull. Sampson et al. believed Deperet's and Lavocat's material
to be undiagnostic, making the diagnostic Majungatholus specimen the
holotype of the species. This also resulted in Majungatholus atopus being
the valid name for this theropod from 1998 until 2007. 2007 saw the publication
of an extensive monograph on the Maevarano abelisaurid (Sampson and Krause,
eds.) which determined both Deperet's and Lavocat's material could be distinguished
from other abelisaurids, making Majungasaurus crenatissimus the valid
name after all.
Carrano et al. (2011) reidentified three supposed Masiakasaurus teeth
from Carrano et al. (2002) as juvenile Majungasaurus- FMNH PR 2198, 2226
and 2228.
Several specimens from outside Madagascar have been referred to Majungasaurus,
but these are all fragmentary and more likely to be other abelisaurid taxa,
as dental apomorphies of Majungasaurus were only discovered in 2007.
Such specimens include teeth and two unguals from Egypt (Gemmellaro, 1921; the unguals being titanosaurian and hadrosauroid),
a tooth from India (Gemmerallo, 1921), teeth from the Lameta Formation of India
(Mathur and Srivastava, 1987), and dentary fragment CMN 41861 from the Kem Kem beds
of Morocco (Russell, 1996).
References- Deperet, 1896. Note on the sauropod and theropod dinosaurs
from the Upper Cretaceous of Madagascar. Bulletin de la Societe Geologique de
France, 3rd series. 24, 176.
Boule, 1896. Note préliminaire sur les débris de dinosauriens
envoyés au Muséum par M. Bastard. Bulletin du Muséum d’Histoire
Naturelle de Paris. 2, 347-351.
Boule, 1900. Note sur quelques fossiles de Madagascar parvenus récemment
au laboratoire de Paléontologie. Bulletin du Muséum National d’Histoire
Naturelle à Paris. 6, 201.
Thevenin, 1906. Note sur des fossils de Madagascar, recueillis par le Dr. Decorse.
Bulletin du Muséum National d’Histoire Naturelle à Paris.
12, 334-336.
Thevenin, 1907. Dinosauriens (Paléontologie de Madagascar IV). Annales
de Paléontologie. 2, 121-136.
Gemmellaro, 1921. Rettili maëstrichtiani di Egitto. Giornale
di Scienze Naturali ed Economiche. 32, 339-351.
Piveteau, 1926. Contribution à l’étude des formations lagunaires
du Nord-Ouest de Madagascar. Bulletin de la Société Géologique
de France. (4) 26, 33-38.
Lavocat, 1955. Etude des gisements de Dinosauriens de la région de Majunga
(Madagascar). Travaux du Bureau Géologique. 69, 1-19.
Lavocat, 1955. Sur une portion de mandibule de Théropode provenant du
Crétacé supérieur de Madagascar. Bulletin du Muséum
National d’Histoire Naturelle à Paris. 27, 256-259.
Lavocat, 1955. Les recherches de reptiles fossils à Madagascar. Le Naturaliste
Malgache. 7(2), 203-207.
Lavocat, 1957. Sur les couches à dinosauriens de Madagascar. CCTA and
Service Géologique de Madagascar, Comptes Rendus. Comités régionaux
Centre, Est et Sud Conférence de Tananarive, Avril 1957, Geology, Second
Volume. 363-364.
Russell, Russell, Taquet and Thomas, 1976. Nouvelles récoltes de Vertébrés
dans les terrains continentaux du Crétacé supérieur de
la région de Majunga (Madagascar). Comptes Rendu Sommaire Des Séances
et Bulletin de la Société Géologique de France. 5, 205-208.
Sues and Taquet, 1979. A pachycephalosaurid dinosaur from Madagascar and a Laurasia-Gondwanaland
connection in the Cretaceous. Nature. 279, 633-635.
Sues, 1980. A pachycephalosaurid dinosaur from the Upper Cretaceous of
Madagascar and its paleobiogeographical implications. Journal of
Paleontology. 54(5), 954-962.
Mathur and Srivastava, 1987. Dinosaur teeth from Lameta Group (Upper Cretaceous)
of Kheda District, Gujarat. Journal of the Geological Society of India. 29,
554-566.
Rage, 1988. Gondwana, Tethys, and terrestrial vertebrates during the Mesozoic
and Cainozoic. In Audley-Charles and Hallam (eds.). Gondwana and Tethys. Geological
Society Special Publication 37. 255-273
Giffin, 1989. Pachycephalosaur paleoneurology (Archosauria: Ornithischia). Journal
of Vertebrate Paleontology. 9(1), 67-77.
Ravoavy, 1991. Identification et mise en catalogue des vertébrés
fossiles récoltes dans le Crétacé supérieur continental
de la région de Berivotra (Majunga) fouille 1987. Université d’Antananarivo
Mémoire de Recherche. Part II, 55-104.
Russell, 1996. Isolated dinosaur bones from the Middle Cretaceous of the Tafilalt,
Morocco. Bulletin du Muse'um national d'Histoire naturelle (4e se'r.) 18, 349-402.
Sampson, Krause, Dodson and Forster, 1996. The premaxilla of Majungasaurus
(Dinosauria: Theropoda) with implications for Gondwanan Paleobiography. Journal
of Vertebrate Paleontology. 16(4), 601-605.
O'Connor, Suny and Sampson, 1998. The vertebral column of Majungatholus atopus
(Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. Journal of
Vertebrate Paleontology. 18(3), 67A.
Sampson, Witmer, Forster and Krause, 1998. The evolution and biogeography of
Gondwanan theropod dinosaurs: New information from the Late Cretaceous of Madagascar.
Journal of African Earth Sciences. Special Abstracts Issue, Gondwana 10: Event
Stratigraphy of Gondwana. 27(1A), 167-168.
Sampson, Witmer, Forster, Krause, O'Connor, Dodson and Ravoavy, 1998. Predatory
dinosaur remains from Madagascar: Implications for the Cretaceous biogeography
of Gondwana. Science, 280, 1048-1051.
Carrano, Sampson and Forster, 2002. The osteology of Masiakasaurus knopfleri,
a small abelisauroid (Dinosauria: Theropoda) from the Late Cretaceous of Madagascar.
Journal of Vertebrate Paleontology. 22(3), 510-534.
Witmer, Ridely and Sampson, 2004. The ear region, cerebral endocast, and cephalic
sinuses of the abelisaurid theropod dinosaur Majungatholus. Journal of
Vertebrate Paleontology. 24(3), 27A.
Fanti, 2005. Stratigraphy and paleontology of the Cretaceous layers of Berivotra
(Mahajanga, Madagascar): Paleobiogeographic implications. Masters thesis. University
of Bologna. 375 pp.
Carrano, 2007. The appendicular skeleton of Majungasaurus crenatissimus
(Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. In Sampson
and Krause (eds.). Majungasaurus crenatissimus (Theropoda: Abelisauridae)
from the Late Cretaceous of Madagascar. SVP Memoir 8, 164-179.
Fanti and Therrien, 2007. Theropod tooth assemblages from the Late Cretaceous
Maevarano Formation and the possible presence of dromaeosaurids in Madagascar.
Acta Palaeontologica Polonica. 52(1), 155-166.
Farke and O'Connor, 2007. Pathology in Majungasaurus crenatissimus (Theropoda:
Abelisauridae) from the Late Cretaceous of Madagascar. In Sampson and Krause
(eds.). Majungasaurus crenatissimus (Theropoda: Abelisauridae) from the
Late Cretaceous of Madagascar. SVP Memoir 8, 180-184.
Krause, Sampson, Carrano and O'Connor, 2007. Overview of the history of discovery,
taxonomy, phylogeny, and biogeography of Majungasaurus crenatissumus
(Theropoda: Abelisauridae) form the Late Cretaceous of Madagascar. In Sampson
and Krause (eds.). Majungasaurus crenatissimus (Theropoda: Abelisauridae)
from the Late Cretaceous of Madagascar. SVP Memoir 8, 1-20.
O'Connor, 2007. The postcranial axial skeleton of Majungasaurus crenatissimus
(Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. In Sampson
and Krause (eds.). Majungasaurus crenatissimus (Theropoda: Abelisauridae)
from the Late Cretaceous of Madagascar. SVP Memoir 8, 127-162.
Rogers, Krause, Curry Rogers, Rasoamiaramanana and Rahanarisoa, 2007. Paleoenivronment
and paleoecology of Majungasaurus crenatissimus (Theropoda: Abelisauridae)
from the Late Cretaceous of Madagascar. In Sampson and Krause (eds.). Majungasaurus
crenatissimus (Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar.
SVP Memoir 8, 21-31.
Sampson and Witmer, 2007. Craniofacial anatomy of Majungasaurus crenatissimus
(Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. In Sampson
and Krause (eds.). Majungasaurus crenatissimus (Theropoda: Abelisauridae)
from the Late Cretaceous of Madagascar. SVP Memoir 8, 32-102.
Smith, 2007. Dental morphology and variation in Majungasaurus crenatissimus
(Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. In Sampson
and Krause (eds.). Majungasaurus crenatissimus (Theropoda: Abelisauridae)
from the Late Cretaceous of Madagascar. SVP Memoir 8, 103-126.
Burch and Carrano, 2008. Abelisaurid forelimb evolution: New evidence from Majungasaurus
crenatissimus (Abelisauridae: Theropoda) from the late Cretaceous of Madagascar.
Journal of Vertebrate Paleontology. 22(3), 58A.
Hieronymus and Witmer, 2008. The facial skin of Majungasaurus crenatissimus
(Abelisauridae: Saurischia): Pronounced dermal metaplasia as the cause of rugosity
in abelisaurid skulls. Journal of Vertebrate Paleontology. 28(3), 90A.
Maganuco, Cau and Pasini, 2008. New information on the abelisaurid pedal elements
from the Late Cretaceous of NW Madagascar (Mahajanga Basin). Atti Della Societa
Italiana de Scienze Naturale Museo Civico de Storia Naturale in Milano. 149
(II), 239-252.
Carrano, Krause, O'Connor and Sampson, 2009. Case 3487 Megalosaurus crenatissimus
Depéret, 1896 (currently Majungasaurus crenatissimus; Dinosauria,
Theropoda): Proposed replacement of the holotype by a neotype. Bulletin of Zoological
Nomenclature. 66(3), 261-264.
Carrano, Loewen and Sertich, 2011. New materials of Masiakasaurus knopfleri
Sampson, Carrano, and Forster, 2001, and implications for the morphology of
the Noasauridae (Theropoda: Ceratosauria). Smithsonian Contributions to Paleobiology.
95, 53 pp.
ICZN, 2011. Opinion 2269 (Case 3487) Megalosaurus crenatissimus Depéret,
1896 (currently Majungasaurus crenatissimus; Dinosauria, Theropoda):
Designation of a neotype. Bulletin of Zoological Nomenclature. 68(1), 89-90.
Tsuihiji and O'Connor, 2012. Reconstruction of muscular and pneumatic systems
in the neck and anterior trunk of Abelisauridae: Insights from Majungasaurus
crenatissimus (Dinosauria: Theropoda). Journal of Vertebrate Paleontology.
Program and Abstracts 2012, 185.
Ratsimbaholison, O'Connor and Felice, 2013. Ontogenetic trends in the craniomandibular
skeleton of Majungasaurus crenatissimus and derivation of the abelisaurid
skull morphotype. Journal of Vertebrate Paleontology. Program and Abstracts
2013, 196.
Ratsimbaholison, Felice and O'Connor, 2014. Craniomandibular ontogeny in nonavian
theropod dinosaurs: Insights from the abelisaurid Majungasaurus crenatissimus.
Journal of Vertebrate Paleontology. Program and Abstracts 2014, 211.
M? sp. indet. (Curry, 1997)
Coniacian?, Late Cretaceous
Ankazomihaboka Sandstones, Madagascar
Reference- Curry, 1997. Vertebrate fossils from the Upper Cretaceous Ankazomihaboka
Sandstones, Mahajanga Basin, Madagascar. Journal of Vertebrate Paleontology.
17(3), 40A.
Rajasaurus Wilson, Sereno, Srivastava,
Bhatt, Khosla and Sahni, 2003
= "Rajasaurus" Badam, 2003
R. narmadensis Wilson, Sereno, Srivastava, Bhatt, Khosla and Sahni,
2003
= "Rajasaurus narmadensis" Badam, 2003
Maastrichtian, Late Cretaceous
Lameta Formation, India
Holotype- (GSI No. 21141/1-33) (7-9 m) braincase, cervical centrum, partial
dorsal vertebrae, sacrum, partial caudal vertebrae, partial scapula, partial
ilia, proximal pubis, femora (one distal), distal tibia, proximal fibula, metatarsals
II, metatarsal IV
Referred- (paralectotypes of Lametasaurus indicus; lost) sacrum
(610 mm), ilia (960 mm), tibia (580 mm) (Matley, 1923)
(in cranial reconstruction online) premaxilla, maxilla, lacrimal, jugal, postorbital,
quadratojugal, dentary, teeth
Diagnosis- (from Wilson et al., 2003) median nasofrontal prominence,
with the frontals forming only the posterior rim of the prominence; anteroposteriorly
elongate supertemporal fenestrae, with length approx. 150% transverse breadth
of frontal; robust ilium with transversed ridge separating the brevis fossa
from the acetabulum.
Comments- Collected in 1983 by Suresh Srivastava of the Geological Survey
of India (GSI) and Ashok Sahni, a paleontologist at Panjab University. Matley
(1923) described a sacrum, ilium, tibia and scutes as the thyreophoran Lametasaurus
indicus. The postcrania were later recognized as theropod by Chakravarti
(1935). The scutes may be crocodilian, sauropod or thyreophoran. Walker (1964)
made them the lectotype of Lametasaurus, leaving the theropod elements
without a name. Wilson et al. note they may belong to Rajasaurus.
References- Matley, 1923. Note on an armored dinosaur from the Lameta
beds of Jubbulpore. Records of the Geological Survey of India. 55, 105-109.
Chakravarti, 1935. Is Lametasaurus indicus an armored dinosaur? American
Journal of Science. 30(5), 138-141.
Walker, 1964. Triassic reptiles from the Elgin area: Ornithosuchus and the origin
of carnosaurs. Philosophical Transactions of the Royal Society of London B.
248, 53-134.
Badam, 2003. Scientists discover new carnivorous dinosaur species in India.
Associated Press, 14 August.
Wilson, Sereno, Srivastava, Bhatt, Khosla and Sahni, 2003. A new abelisaurid
(Dinosauria, Theropoda) from the Lameta Formation (Cretaceous, Maastrichtian)
of India. Contributions from the Museum of Paleontology. 31(1), 1-42.
Brachyrostra Canale, Scanferla, Agnolin and
Novas, 2009
Definition- (Carnotaurus sastrei <- Majungasaurus crenatissimus)
(Canale, Scanferla, Agnolin and Novas, 2009)
References- Canale, Scanferla, Agnolin and Novas, 2009. New carnivorous
dinosaur from the Late Cretaceous of NW Patagonia and the evolution of abelisaurid
theropods. Naturwissenschaften. 96, 409-414.
Thanos Delcourt and Iori, 2020
= "Thanos" Delcourt and Iori, online 2018
T. simonattoi Delcourt and Iori, 2020
= "Thanos simonattoi" Delcourt and Iori, online 2018
Santonian, Late Cretaceous
Sao Jose do Rio Preto Formation of the Bauru Group, Brazil
Holotype- (MPMA 08-0016-95) (~5-6 m) (subadult or adult) incomplete axis (82.62 mm)
Diagnosis- (after Delcourt and Iori, 2018) well-developed keel on ventral surface of axis becoming wider and deeper posteriorly.
Other diagnoses- Delcourt and
Iori (2018) list "lateral small foramina separated by a relative wide
wall", referring to the axial pleurocoels. However, pneumatic
features like this can be incredibly variable in number and size, even
between sides of the same specimen. For instance, Tyrannosaurus
specimen FMNH PR2081 has two pairs on each side, but the ventral one is
much larger on the left side, while MOR 555 has the dorsal pleurocoel
split and only small foramina in fossae ventrally. Even Thanos'
pleurocoels show a 20% difference in separation between sides, and so
this difference between the single known axes of Thanos and Carnotaurus is rejected as taxonomically valid.
They also listed "well-developed and deep prezygapophyseal
spinodiapophyseal fossae", stating "the prezygapophyseal
spinodiapophyseal fossae in Thanos
are deeper than in any ceratosaur, including all abelisauroids."
Yet abelisaurids do not have prezygapophyseal spinodiapophyseal fossae,
which would require a spinodiapophyseal lamina to exist. Delcourt
and Iori label a hollow the prezygapophyseal spinodiapophyseal fossa in
their figure 2D, but this is an internal feature (probably the
centroprezygapophyseal fossa or prezygapophyseal centrodiapophyseal
fossa based on CT-scanned noasaurid cervical DGM929-R) as the
prezygapophysis and diapophysis are unpreserved.
Comments- Discovered in 1995,
the anterior portion of this axis was described by Mendez et al. (2014)
as Abelisauroidea indet., although they noted due to its larger size it
"possibly would be referable to Abelisauridae." In 2014 the
posterior portion was discovered and described by Delcourt and Iori
(2020) as a new taxon of abelisaurid. Note that while the
electronic version was posted on November 16 2018, there is no mention
of ZooBank in the paper and the journal's website lists it under
"Latest Articles" ("articles accepted for publication in this journal
but not yet published in a volume/issue"). Thus according to ICZN
Article 8.5.3 (a work must "be registered in
the Official Register of Zoological Nomenclature (ZooBank) (see Article
78.2.4) and contain evidence in the work itself that such registration
has occurred"), "Thanos simonattoi" Delcourt and Iori, 2018 was a nomen
nudum until the physical volume was published in August 8 2020.
Of Thanos' supposed three autapomorphies, one is questionable due to high
interspecific variation in theropods and another is invalid (see
above). The third is that the ventral median keel is stronger than Majungasaurus and Carnotaurus, particularly posteriorly. As Carnotaurus
is the only other brachyrostran with a preserved axis, Thanos cannot
be distinguished from any other member of the clade, though it is
probably not a junior synonym considering it's from a unique
formation. Thus while technically diagnostic and probably a new
species, the usefullness of giving a name to a single element based on
a single difference which can only be compared to a single close
relative is questionable.
Delcourt and Iori (2018) added Thanos to the Tortosa et al.
ceratosaur matrix and recovered it as a brachyrostran in a polytomy
with all taxa except Rahiolisaurus plus Dahalokely.
References- Mendez, Novas and Iori, 2014. New record of abelisauroid theropods
from the Bauru Group (Upper Cretaceous), Sao Paulo state, Brazil. Revista Brasileira
de Paleontologia. 17(1), 23-32.
Delcourt and Iori, 2020 (online 2018). A new Abelisauridae (Dinosauria: Theropoda)
from São José do Rio Preto Formation, Upper Cretaceous of Brazil and
comments on the Bauru Group fauna. Historical Biology. 32(7), 917-924.
Xenotarsosaurus Martinez,
Gimenez, Rodriguez and Bochatey, 1987
X. bonapartei Martinez, Gimenez, Rodriguez and Bochatey, 1987
Cenomanian?, Late Cretaceous
Bajo Barreal Formation, Chubut, Argentina
Syntypes- (PVL 612) (4.8 m) femur (611 mm), tibia (592 mm), fibula, astragalocalcaneum
(134 mm wide, 96 mm tall)
....(UNPSJB PV 184) two anterior dorsal vertebrae
References- Martínez, Gimenez, Rodríguez and Bochatey,
1986. Xenotarsosaurus bonapartei nov. gen. et sp. (Carnosauria, Abelisauridae),
a new theropod from the Bajo Barreal Formation, Chubut, Argentina. Actas del
Congreso Argentino de Paleontología y Bioestratigrafía. 4, 3-31.
Coria and Rodríguez, 1993. Sobre Xenotarsosaurus bonapartei Martínez,
Giménez, Rodríguez, y Bochatey, 1986; un problematico Neoceratosauria
(Novas, 1989) del Cretacico de Chubut. Ameghiniana. 30(3), 326-327.
Dahalokely Farke and Sertich,
2013
D. tokana Farke and Sertich, 2013
Turonian, Late Cretaceous
Ambolafaltsy Formation, Madagascar
Holotype- (UA 9855) (~3.5 m subadult) fifth cervical vertebra (64.5 mm),
first dorsal vertebra (44 mm), second dorsal vertebra (48.7 mm), sixth dorsal
vertebra (52.1 mm), seventh dorsal vertebra (55.1 mm), eighth dorsal vertebra
(55.2 mm), incomplete ninth dorsal vertebra, second dorsal rib (339.9 mm), seven
dorsal rib fragments
Diagnosis- (after Farke and Sertich, 2013) mid-cervical vertebrae have
prezygoepipophyseal lamina with prominent convexity at midpoint nearly equal
in length to centrum and prominent notches at either end separating convexity
from epipophyses; prezygapophyses and centroprezygapophyseal lamina nearly vertical
and linear in lateral view in dorsals 1 and 2, with cranial margin of prezygapophyses
and cranial face of centrum nearly co-planar; postzygapophyses on dorsal 2 strongly
concave; infraprezygapophyseal fossa divided through dorsal six.
Comments- The specimen was discovered in 2007 and announced by Farke
and Sertich (2009) as an abelisaurid. It was later described by Farke and Sertich
(2013), who found it to be a basal noasaurid. Only one extra step is needed
for it to be a majungasaurine though, and two more for it to be a non-abelisaurian
abelisauroid. The Tortosa et al. matrix recovered it as a non-furileusaurian
brachyrostran.
References- Farke and Sertich, 2009. A medium-sized theropod from the
Late Cretaceous (Turonian or Coniacian) of the Ambilobe Basin, Northernmost
Madagascar. Journal of Vertebrate Paleontology. 29(3), 94A.
Farke and Sertich, 2013. An abelisauroid theropod dinosaur from the Turonian
of Madagascar. PLoS ONE. 8(4), e62047.
Rahiolisaurus Novas,
Chaterjee, Rudra and Datta, 2010
= "Rahiolisaurus" Novas, Chaterjee, Rudra and Datta vide Mendez, Novas
and Chatterjee, online 2010
R. gujaratensis Novas, Chaterjee, Rudra and Datta, 2010
= "Rahiolisaurus gujaratensis" Novas, Chaterjee, Rudra and Datta vide
Mendez, Novas and Chatterjee, online 2010
Late Maastrichtian, Late Cretaceous
Lameta Formation, India
Holotype- (ISIR 550) (~8 m adult) ilium fused to proximal pubis
....(ISIR 554) incomplete pubis (~750-800 mm)
....(ISIR 557) femur (770 mm)
Paratypes- (ISIR 401) premaxilla
(ISIR 402-403)
(ISIR 404-407b) fused first to fifth sacral vertebrae
(ISIR 408-409) two mid and distal caudal vertebrae
(ISIR 410) proximal caudal vertebra
(ISIR 411-414) four mid and distal caudal vertebrae
(ISIR 415) posterior dorsal centrum
(ISIR 416-429) fourteen mid and distal caudal vertebrae
(ISIR 430-431)
(ISIR 432) incomplete scapula
(ISIR 433)
(ISIR 435)
(ISIR 436) ilium (800 mm) fused to proximal pubis
(ISIR 437) ilium fused to proximal pubis
(ISIR 438) tibia (430 mm)
(ISIR 439) fused distal ischia
(ISIR 440) femur
(ISIR 441) femur
(ISIR 442) femur
(ISIR 443) femur
(ISIR 444) femur
(ISIR 445) tibia (600 mm)
(ISIR 446) tibia
(ISIR 447) tibiotarsus
(ISIR 449) tibia
(ISIR 450) metatarsal
(ISIR 451-454)
(ISIR 457) tibia
(ISIR 458) tibia
(ISIR 464) pubis
(ISIR 465) incomplete scapulocoracoid
(ISIR 474)
(ISIR 475) metatarsal IV
(ISIR 486-503)
(ISIR 504) posterior dorsal centrum
(ISIR 505) posterior dorsal centrum
(ISIR 506) posterior dorsal centrum
(ISIR 506-507) fused first and second sacral vertebrae
(ISIR 508) posterior dorsal centrum
(ISIR 509) posterior dorsal centrum
(ISIR 510) posterior dorsal centrum
(ISIR 511-514) fused first to fifth sacral vertebrae
(ISIR 515) sixth cervical vertebra
(ISIR 516-517) fused fifth and sixth sacral vertebrae
(ISIR 518-545) twenty-eight mid and distal caudal vertebrae
(ISIR 546) chevron
(ISIR 547) proximal scapula
(ISIR 549) metatarsal
(ISIR 551) ilium fused to proximal pubis
(ISIR 552) pubis
(ISIR 553) pubis
(ISIR 555) fibula
(ISIR 556) femur
(ISIR 558) femur
(ISIR 559) femur
(ISIR 560) femur
(ISIR 561) femur
(ISIR 562) tibia
(ISIR 563) tibia
(ISIR 564) tibia
(ISIR 565) tibia
(ISIR 566-568)
(ISIR 569) metatarsal II
(ISIR 570) tibiotarsus
(ISIR 571) tibiotarsus
(ISIR 572)
(ISIR 573) metatarsal I(?), fused metatarsals II and III
(ISIR 574)
(ISIR 575) metatarsal
(ISIR 576) pedal phalanx
(ISIR 577) pedal phalanx
(ISIR 578) pedal phalanx
(ISIR 579) pedal phalanx
(ISIR 580)
(ISIR 581) metatarsal
(ISIR 582) pedal phalanx
(ISIR 583-584)
(ISIR 585) pedal phalanx
(ISIR 586) metatarsal
(ISIR 587) pedal phalanx
(ISIR 588) pedal phalanx
(ISIR 589) pedal phalanx
(ISIR 590) pedal phalanx
(ISIR 591) pedal phalanx
(ISIR 592) pedal phalanx
(ISIR 593) pedal phalanx
(ISIR 594) pedal phalanx
(ISIR 595) pedal phalanx
(ISIR 596-602)
(ISIR 634) ilium fused to proximal pubis
(ISIR 645) proximal scapula
(ISIR 649)
(ISIR 657) proximal humerus (~250 mm) (Mendez, Novas and Chatterjee, 2010)
(ISIR 658) axis
....(ISIR 659) third cervical vertebra
....(ISIR 660) fourth cervical vertebra
Diagnosis- (after Novas et al., 2010) premaxillary interdental plates
fused and lacking vertical ridges; dental foramina absent; premaxillary teeth
with teardrop-shaped cross section; a faint mesial keel but a rounded distal
edge; iliac blade with deep caudal notch on postacetabular process; metatarsal
I rod-like; metatarsal II strongly narrow proximally.
Comments- Chatterjee and Rudra (1996) mentioned and schematically illustrated
new abelisaurid material discovered in 1995 and 1997 which they referred to
Indosuchus. Though they described it as having an unreduced forelimb,
the humerus (ISIR 434) has been reassigned to Titanosauria by Novas et al. Similarly,
the supposed radius (ISIR 549) is a broken metatarsal of Rahiolisaurus.
Mendez et al. (2010) described a proximal humerus belonging to the taxon which
is reduced as in other abelisaurs.
The name "Rahiolisaurus gujaratensis" was first listed in the bibliography
of Mendez et al. (2010), which was published online before (February 3) the
official description was published (June 28). However, Mendez et al.'s paper
was not published on paper until the September issue, making the nomen nudum
online only.
References- Chatterjee and Rudra, 1996. KT events in India: Impact, rifting,
volcanism and dinosaur extinction. In Novas and Molnar (eds.). Proceedings of
the Gondwanan Dinosaur Symposium: Memiors of the Queensland Museum. 39(3), 489-532.
Mendez, Novas and Chatterjee, 2010. An abelisaurid humerus from the Upper Cretaceous
of India. Paläontologische Zeitschrift. 84(3), 421-425.
Novas, Chaterjee, Rudra and Datta, 2010. Rahiolisaurus gujaratensis,
n. gen. n. sp., a new abelisaurid theropod from the Late Cretaceous of India.
In Bandyopadhyay (ed). New aspects of Mesozoic biodiversity. Springer. Lecture
Notes in Earth Science. 132, 185 pp.
"Bayosaurus" Coria, Currie
and Carabajal, 2006
"B. pubica" Gasparini, Salgado and Coria, 2007
Turonian, Late Cretaceous
Lisandro Formation of the Rio Limay Subgroup, Neuquen, Argentina
Material- (MCF-PVPH-237; Cerro Bayo Mesa taxon) eleventh dorsal vertebra
(63 mm), anterior sacrum (91, 110, 62 mm), incomplete ilium, proximal pubes,
proximal ischia
Comments- Coria et al. (2006) do not officially name this specimen, but
the name "Bayosaurus" can be found in large font in their cladogram
(figure 6), while the specimen number MCF-PVPH-237 is lacking. Thus, one can
infer they intended to name the specimen "Bayosaurus", but the name
was left in the figure accidentally. This is confirmed by Coria (pers. comm.
to Auditore, 2007). Gasparini et al. (2007) reference "Bayosaurus pubica",
citing Coria's chapter in the same volume, yet Coria never uses that name. This
is apparently another prepublication mistake. Initially identified merely as
an abelisauroid, Tortosa et al. (2014)
recovered it as an abelisaurid, and Filippi et al. (2016;
as the Cerro Bayo Mesa taxon) found it to be a brachyrostran closer to furileusaurians
than Xenotarsosaurus, Dahalokely and Rahiolisaurus, but
outside Carnotaurini.
References- Coria, Currie and Carabajal, 2006. A new abelisauroid theropod
from Northwestern Patagonia. Canadian Journal of Earth Sciences. 43, 1283-1289.
Gasparini, Salgado and Coria, 2007. Reptilian faunal succession in the Mesozoic
of Patagonia: An updated overview. In Gasparini, Salgado and Coria (eds.). Patagonian
Mesozoic Reptiles. Indiana University Press, Bloomington, Indiana. 335-358.
Tortosa, Buffetaut, Vialle, Dutour, Turini and Cheylan, 2014. A new abelisaurid
dinosaur from the Late Cretaceous of southern France: Palaeobiogeographical
implications. Annales de Paléontologie. 100(1), 63-86.
Filippi, Mendez, Juarez Valieri and Garrido, 2016. A new brachyrostran with
hypertrophied axial structures reveals an unexpected radiation of latest Cretaceous
abelisaurids. Cretaceous Research. 61, 209-219.
Guemesia Agnolín, Cerroni, Scanferla, Goswami, Paulina-Carabajal, Halliday, Cuff and Reuil, 2022
G. ochoai Agnolín, Cerroni, Scanferla, Goswami, Paulina-Carabajal, Halliday, Cuff and Reuil, 2022
Campanian-Maastrichtian, Late Cretaceous
Los Blanquitos Formation, Salta, Argentina
Holotype- (IBIGEO-P 103) (subadult or adult) incomplete braincase
Diagnosis- (after Agnolin et
al., 2022) frontals with median row of superficial and large pits;
orbital region of each frontal highly pneumatized by the presence of a
passage (probably pneumatic and vascular) which connects the orbital
margin with the dorsal surface; supratemporal fossa subdivided by a low
ridge at the orbital level; dorsal margin of paroccipital process
(below the posterolateral alae of the parietals) convex, forming a
rounded protuberance (also in Majungasaurus, Viavenator, Llukalkan and Ekrixinatosaurus);
subdivided anterior tympanic recess; robust and well-defined
otosphenoidal crest markedly anteroposteriorly elongate (running from
crista antotica to base of the paroccipital process) (also in Majungasaurus and Viavenator); cranial nerves IX-XII open within a well-marked paracondylar recess; exit foramina for orbitocerebral vein (also in Carnotaurus);
basipterygoid process elongate with distal end posteroventrally
oriented and projected below the basal tubera level; distance between
basipterygoid process and basal tuber shorter than the transverse
distance between basal tubera; basal tubera delicate and not thickened,
distally separated from each other by notch in the basituberal web;
basisphenoid recess small and subcircular, medially does not reach neck
of occipital condyle; foramen magnum transversely wider than
dorsoventrally tall (also in Spectrovenator and Arcovenator).
Comments- The holotype was
discovered in 2015 (Agnolin, pers. comm. 2-22-2002). Note the
article was first published online on February 10 2022, despite the
issue being listed as 2021. Agnolin et
al. (2022) used Filippi et al.'s ceratosaur analysis to recover Guemesia as a brachyrostran in a polytomy with Ekrixinatosaurus, Ilokelesia, Llukalkan, Niebla, Skorpiovenator and carnotaurins.
Reference- Agnolín, Cerroni,
Scanferla, Goswami, Paulina-Carabajal, Halliday, Cuff and Reuil, 2022 (as 2021).
First definitive abelisaurid theropod from the Late Cretaceous of
northwestern Argentina. Journal of Vertebrate Paleontology. 41(4), e2002348.
unnamed possible brachyrostran (Salem, Lamanna, O'Connor, El-Qot, Shaker, Thabet, El-Sayed and Sallam, 2021)
Early Cenomanian, Late Cretaceous
Gebel el Dist, Baharija Formation, Egypt
Material- (MUVP 477) (~5.8 m) tenth cervical vertebra (67.00 mm)
Comments- Discovered in 2016,
Salem et al. (2021) added this to Tortosa et al.'s ceratosaur matrix
and recovered it as an abelisaurid based on "(1) long axis of
diapophysis forms angle of 65° to midsagittal plane; (2) dorsal surface
of neural arch clearly delimited from lateral surface of diapophysis;
(3) deep spinoprezygapophyseal and spinopostzygapophyseal fossae; and
(4) well-developed epipophyses, comparable to those observed in the
tenth cervicals of Majungasaurus and MPM 99 (but smaller than those of Carnotaurus and Ekrixinatosaurus novasi)."
The resulting Salem et al. (2022) publication officially describing the
specimen did not use character 1. While it was a brachyrostran
(in a polytomy with Ekrixinatosaurus, Ilokelesia
and the derived clade) in majority rule trees, a posteriori pruning of
taxa was not done to determine if this is real signal or merely an
artifact. Examination of the 2022 supplementary data should resolve this.
References- Salem, Lamanna,
O'Connor, El-Qot, Shaker, Thabet, El-Sayed and Sallam, 2021. First
definitive record of Abelisauridae from the Bahariya Formation,
Bahariya Oasis, Western Desert of Egypt increases diversity of
large-bodied theropods in the MIddle Cretaceous of northeastern Africa.
The Society of
Vertebrate Paleontology Virtual Meeting Conference Program, 81st Annual
Meeting. 225-226.
Salem, Lamanna,
O'Connor, El-Qot, Shaker, Thabet, El-Sayed and Sallam, 2022. First
definitive record of Abelisauridae (Theropoda: Ceratosauria) from the
Cretaceous Bahariya Formation, Bahariya Oasis, Western Desert of Egypt.
Royal Society Open Science. 9: 220106.
Ilokelesia Coria and
Salgado, 2000
= "Ilokelesia" Coria and Salgado vide Coria, 1999
I. aguadagradensis Coria and Salgado, 2000
Late Cenomanian, Late Cretaceous
Middle Cenomanian-Early Turonian, Late Cretaceous
Huincul Formation of Rio Limay Subgroup, Neuquen, Argentina
Holotype- (~5 m) postorbital, quadrate, occipital condyle, partial third
cervical vertebra, fourth cervical vertebra, posterior dorsal vertebra, five
mid caudal vertebrae, eight chevrons, eight pedal phalanges, two pedal unguals
Diagnosis- (after Coria and Salgado, 2000) quadrate with reduced lateral
condyle; cervical vertebrae with very reduced diapopostzygopophyseal laminae;
dorsal vertebrae with ventrally concave infraparapophyseal laminae and with
ventrally oriented parapophyses; dorsal vertebrae lacking pleurocoels; mid caudal
vertebrae with distally expanded transverse processes bearing anteriorly and
posteriorly projecting processes; distal edge of caudal transverse processes
exhibiting a gently sigmoid profile that is convex anteriorly and concave posteriorly.
Comments- Note issue 15 of GAIA is listed as December 1998 but was not
actually published until October 2000.
References- Coria, 1999. Ornithopod dinosaurs from the Neuquén
Group, Patagonia, Argentina: Phylogeny and biostratigraphy. In Tomida, Rich
and Vickers-Rich (eds). Proceedings of the Second Gondwanan Dinosaur Symposium,
National Science Museum Monographs 15. 47-60.
Coria and Salgado, 1999. A primitive abelisaur theropod from the Rio Limay Formation
(Upper Cretaceous) of Patagonia. Journal of Vertebrate Paleontology. 19(3),
39A.
Coria and Salgado, 2000 (as 1998). A basal Abelisauria Novas 1992 (Theropoda-Ceratosauria)
from the Cretaceous of Patagonia, Argentina. Gaia. 15, 89-102.
Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution of the carnivorous
dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research.
45, 174-215.
Ekrixinatosaurus Calvo,
Rubilar-Roger and Moreno, 2004
= "Ekrixinatosaurus" Juarez Valieri, Fiorelli and Cruz, 2004
E. novasi Calvo, Rubilar-Roger and Moreno, 2004
= "Ekrixinatosaurus novasi" Juarez Valieri, Fiorelli and Cruz, 2004
Early Cenomanian, Late Cretaceous
Candelaros Formation of Rio Limay Group, Neuquen, Argentina
Holotype- (MUCPv-294) (7-8 m, ~1.4 tons) (skull ~850 mm) partial maxillae,
postorbital, squamosal, frontals, parietals, parasphenoid, occiput, incomplete
dentaries, teeth, anterior cervical vertebra, mid-posterior cervical vertebra,
dorsal centrum, dorsal ribs, sacrum, caudal vertebrae, chevrons, ilia, pubis,
proximal ischia, femora (776 mm), tibiae (694 mm), proximal fibula, astragalus,
calcaneum, metatarsals, phalanx II-1 (118 mm), phalanx III-1 (128 mm), phalanges,
pedal ungual
Diagnosis- (after Calvo et al., 2004) fenestra between postorbital and
frontal; posteriorly directed protuberance on contact of parietal and paroccipital
process; anteroposteriorly compressed cervical vertebrae; cervical neural spines
as tall as epipophyses; mid-posterior cervical centrum with flat venter; two
wide foramina on mid-posterior cervical vertebrae; small prespinal depression
with pneumatic excavation connected to neural canal in mid-posterior cervical
vertebrae; small prespinal lamina on mid-cervical vertebrae; tibia with swelling
at midshaft.
References- Juárez Valieri, Fiorelli and Cruz, 2004. Quilmesaurus
curriei Coria, 2001. Su validez taxonómica y relaciones filogenéticas.
XX Jornadas Argentinas de Paleontología de Vertebrados, Resúmenes.
36-37.
Calvo, Rubilar-Roger and Moreno, 2004. A new Abelisauridae (Dinosauria: Theropoda)
from northwest Patagonia. Ameghiniana. 41(4), 555-563.
Juarez Valieri, Porfiri and Calvo, 2011. New information on Ekrixinatosaurus
novasi Calvo et al 2004, a giant and massively-constructed abelisauroid
from the "Middle Cretaceous" of Patagonia. In Calvo, Porfiri, Gonzalez
Riga and Dos Santos (eds.). 2. Paleontología y Dinosaurios desde América
Latina. 161-169.
Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution of the carnivorous
dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research.
45, 174-215.
Skorpiovenator Canale,
Scanferla, Agnolin and Novas, 2009
= "Skorpiovenator" Canale, Scanferla, Agnolin and Novas, online 2008
S. bustingorryi Canale, Scanferla, Agnolin and Novas, 2009
= "Skorpiovenator bustingorryi" Canale, Scanferla, Agnolin and Novas,
online 2008
Middle Cenomanian-Early Turonian, Late Cretaceous
Huincul Formation of Rio Limay Subgroup, Neuquen, Argentina
Holotype- (MMCH-PV 48) (~6 m, ~1.2 tons) skull, mandible, ten cervical
vertebrae, cervical ribs, eleven dorsal vertebrae, dorsal ribs, few gastralia,
sacrum, caudal vertebrae 1-12, three distal caudal vertebrae, chevron, ulna,
ilium, proximal pubis, proximal ischium, femora, tibiae, fibula, astragali,
calcaneum, metatarsal II, phalanx II-1, phalanx II-2, metatarsal III, phalanx
III-1, phalanges III-2, phalanges III-3, pedal ungual III, metatarsal IV, phalanx
IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4
Diagnosis- (after Canale et al., 2009) ascending process of maxilla homogeneously
wide anteroposteriorly; maxillary horizontal ramus dorsoventrally deep with
subparallel dorsal and ventral margins; maxilla/jugal contact subvertical; 19
maxillary teeth; lacrimal anteriorly projected and with well-developed suborbital
process; quadratojugal with pronounced posterodorsal notch; dentary with posteroventral
process bifurcated to receive anterior end of angular; angular with anterior
end dorsoventrally deep to fit between splenial and prearticular.
Comments- Canale et al.'s paper was published online in December 2008
before being officially published on paper in March 2009.
References- Canale, Scanferla, Agnolin and Novas, 2009. New carnivorous
dinosaur from the Late Cretaceous of NW Patagonia and the evolution of abelisaurid
theropods. Naturwissenschaften. 96, 409-414.
Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution of the carnivorous
dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research.
45, 174-215.
Canale and Novas, 2015. New information about the anatomy and phylogenetic relationships
of Skorpiovenator bustingorryi (Theropoda, Ceratosauria) from the Upper
Cretaceous of Neuquen Province, Patagonia, Argentina. Journal of Vertebrate
Paleontology. Program and Abstracts 2015, 102.
Furileusauria Filippi, Mendez, Juarez Valieri
and Garrido, 2016
Definition- (Carnotaurus sastrei <- Ilokelesia aguadagrandensis,
Skorpiovenator bustingoryi, Majungasaurus crenatissimus) (Filippi,
Mendez, Juarez Valieri and Garrido, 2016)
Other definition- (Carnotaurus sastrei <- Skorpiovenator bustingorryi) (modified after Delcourt, 2018)
= Carnotaurinae sensu Hendrickx, Hartman and Mateus, 2015
Definition- (Carnotaurus sastrei <- Skorpiovenator bustingorryi)
= Furileusauria sensu Delcourt, 2018
Definition- (Carnotaurus sastrei <- Skorpiovenator bustingorryi)
(modified)
References- Filippi, Mendez, Juarez Valieri and Garrido, 2016. A new brachyrostran
with hypertrophied axial structures reveals an unexpected radiation of latest
Cretaceous abelisaurids. Cretaceous Research. 61, 209-219.
Delcourt, 2018. Ceratosaur palaeobiology: New insights on evolution and
ecology of the southern rulers. Scientific Reports. 8:9730.
Elemgasem Baiano, Pol, Bellardini, Windholz, Cerda, Garrido and Coria, 2022
E. nubilus Baiano, Pol, Bellardini, Windholz, Cerda, Garrido and Coria, 2022
Turonian-Coniacian, Late Cretaceous
Sierra del Portezuelo, Portezuelo Formation, Neuquen, Argentina
Holotype- (MCF-PVPH-380) (8
year old subadult) mid cervical epipophysis, two incomplete mid caudal
vertebrae (57, 55 mm), three fused mid caudal vertebrae (two partial;
57 mm) fused to chevron fragment, five distal caudal centra (three
partial; 52, 46 mm), two fused distal caudal centra (one partial; 52
mm), distal femur, proximal tibial fragment, proximal fibulae,
incomplete astragalocalcaneum, distal metatarsals II, proximal pedal
ungual II, distal tarsal III fused with proximal metatarsal III, distal
metatarsal III fragment, phalanges III-2 (one partial; 57 mm), proximal
pedal ungual III, proximal metatarsal IV, distal metatarsals IV,
phalanges IV-2 (one incomplete, one fragmentary), phalanges IV-3 (one
partial)
Diagnosis- (after Baiano et
al., 2022) distal caudal vertebrae with oval centrum articular
surfaces, being transversely wider than dorsoventrally high; distal
caudal centra with longitudinal median ventral groove; marked rugosity
formed by ridges and tubercles on lateral surface of proximal fibula;
proximally high wall of astragalocalcaneum separating tibia from fibula
observed laterally; proximodistally deep lateral surface of calcaneum.
Comments- The holotype was discovered in 2002. Note the fused caudals are pathological.
Baiano et al. (2022) added it to Baiano's ceratosaur matrix to recover it in a polytomy with other furileusaurians. "When Elemgasem
is constrained in more basal positions within Brachyrostra, the MPTs
are 1 step longer, whereas when it is constrained as a majungasaurine
the MPTs are 4 steps longer. Four extra steps were also required to
place Elemgasem as a non-abelisaurid abelisauroid. Finally, constraining Elemgasem within Noasauridae required MPTs of 5 extra steps."
References- Baiano, Pol, Bellardini, Windholz, Cerda, Garrido and Coria, 2022. Elemgasem nubilus:
A new brachyrostran abelisaurid (Theropoda, Ceratosauria) from the
Portezuelo Formation (Upper Cretaceous) of Patagonia, Argentina. Papers
in Palaeontology. 8(5), e1462.
Llukalkan Gianechini, Méndez, Filippi, Paulina-Carabajal, Juárez-Valieri and Garrido, 2021
L. aliocranianus Gianechini, Méndez, Filippi, Paulina-Carabajal, Juárez-Valieri and Garrido, 2021
Santonian, Late Cretaceous
Bajo de la Carpa Formation of the Rio Colorado Subgroup, Neuquen, Argentina
Holotype- (MAU-Pv-LI-581) (subadult?) incomplete skull (392 mm), prearticular
Diagnosis- (after Gianechini et
al., 2021) T-shaped lacrimal with anterior process longer than half of
the ventral process' length (ontogenetic?); slender ventral lacrimal
process without posterior suborbital process (ontogenetic?); presence
of a notch followed by a bulge on the anteromedial border of the
supratemporal fossa; caudal tympanic recess located just posterior to
the columellar recess; dorsoventrally expanded paroccipital process
with distal end posterodorsally oriented and posteriorly extended
beyond the occipital condyle; large foramina for the caudal middle
cerebral veins that are widely separated from the supraoccipital crest
and clearly observed in posterior view of the braincase; slender and
anteroventrally projected basipterygoid process.
Comments- This was discovered in 2015. Gianechini et al. (2021) note that "The holotype of Llukalkan shows similarities with Viavenator,
including squamosals and frontals lacking cornual processes, and the
general morphology of the endocranial cavities" and was discovered in
the same locality. However they retained it as a valid taxon
based on braincase differences and the small difference in size (~82%
the size of Viavenator's holotype), although lacrimal morphology suggests it maybe a subadult.
Gianechini et al. used a version of Tortosa et al.'s ceratosaur analysis to recover Llukalkan as a furileusaurian in a polytomy with Pycnonemosaurus, Quilmesaurus (neither comparable due to lacking cranial elements), Viavenator and carnotaurins.
Reference- Gianechini, Méndez,
Filippi, Paulina-Carabajal, Juárez-Valieri and Garrido, 2021. A new
furileusaurian abelisaurid from La Invernada (Upper Cretaceous,
Santonian, Bajo De La Carpa Formation), northern Patagonia, Argentina.
Journal of Vertebrate Paleontology. e1877151.
Pycnonemosaurus Kellner and
Campos, 2002
P. nevesi Kellner and Campos, 2002
Campanian-Maastrichtian, Late Cretaceous
Parecis Group, Brazil
Holotype- (DGM 859-R) (subadult) nine teeth(?), rib fragments, two incomplete
proximal caudal vertebrae (148, 144 mm), four proximal caudal centra (155, 154,
149, 155 mm), three proximal caudal transverse processes, incomplete pubis,
incomplete tibia (870 mm), distal fibula
Diagnosis- (after Delcourt, 2017) distal caudal transverse process hook-shaped
with anterodistal expansion being short and bowed; pubis with small rounded
foot and ventrally bowed anterodistal end; well-developed lateral malleolus
of tibia that is ventrally expanded.
Other diagnoses- Kellner and Campos (2002) listed "moderate distal
expansion of caudal transverse processes" and "relatively small pubic
foot", both of which were elaborated upon by Delcourt (2017). Kellner and
Campos also listed "hatchet-shaped cnemial crest", but Delcourt noted
this is present in many abelisaurids.
Comments- Initially described as being from the Adamantina Formation
of the Bauru Group, Bittencourt and Langer (2011) reassigned the locality to
the later Parecis Group. Delcourt (2017) questioned the referral of teeth to
the genus, as they "were found isolated from any dentigerous element (i.e.
premaxilla, maxilla and dentary) and the Pycnonemosaurus holotype is
based on caudal vertebrae and hind limb elements." Kellner and Campos (2002)
noted "several unidentified incomplete bones" were present in the
holotype, which were identified by Delcourt as a sauropod mid-distal caudal
neural spine similar to Aeolosaurus, titanosaurian distal caudal centrum,
abraded transverse process and large pneumatic element. Delcourt updated this
taxon's scores in Tortosa et al.'s ceratosaur analysis and found it to be in
a polytomy with other furileusaurians.
References- Bittencourt and Kellner, 2002. Abelisauria (Theropoda, Dinosauria)
teeth from Brazil. Boletim do Museu Nacional, Nova Série. 63, 1-8.
Kellner and Campos, 2002. On a theropod dinosaur (Abelisauria) from the continental
Cretaceous of Brazil. Arquivos do Museu Nacional. 60(3), 163-170.
Bittencourt and Langer, 2011. Mesozoic dinosaurs from Brazil and their biogeographic
implications. Anais da Academia Brasileira de Ciências. 83(1), 23-60.
Delcourt, 2017. Revised morphology of Pycnonemosaurus nevesi Kellner
& Campos, 2002 (Theropoda: Abelisauridae) and its phylogenetic relationships.
Zootaxa. 4276(1), 1-45.
Quilmesaurus Coria, 2001
Q. curriei Coria, 2001
Late Campanian, Late Cretaceous
Allen Formation, Rio Negro, Argentina
Holotype- (MPCA-PV-100) (6+ year old subadult) distal femur, tibia (520 mm)
Comments- Juarez Valieri et al. (2004) assigned Quilmesaurus to
Abelisauridae based on the marked distal expansion of the cnemial crest and
the asymmetrical distal end of the tibia, and to Carnotaurinae because of the
downturned distal part of the cnemial crest. More recently, it has been recovered
as a basal furileusaurian in the Tortosa et al. matrix.
Reference- Coria, 2001. New theropod from the Late Cretaceous of Patagonia.
In Tanke and Carpenter (eds.). Mesozoic Vertebrate Life: New Research inspired
by the Paleontology of Philip J. Currie. Indiana University Press, Bloomington
& Indianapolis, Indiana. 3-9.
Juárez Valieri, Fiorelli and Cruz, 2004. Quilmesaurus curriei
Coria, 2001. Su validez taxonómica y relaciones filogenéticas.
XX Jornadas Argentinas de Paleontología de Vertebrados (La Plata), Resúmenes:
36-37.
Baiano and Cerda, 2017. Bone microstructure of Quilmesaurus curriei
(Theropoda; Abelisauridae). Reunión de Comunicaciones de la Asociación
Paleontológica Argentina 2017. Publicacion electronica asociacion
Paleontologica Argentina. 18(2), R10.
Viavenator Filippi, Mendez, Juarez
Valieri and Garrido, 2016
V. exxoni Filippi, Mendez, Juarez Valieri and Garrido, 2016
Santonian, Late Cretaceous
Bajo de la Carpa Formation of the Rio Colorado Subgroup, Neuquen, Argentina
Holotype- (MAU-Pv-LI-530) (skull ~478 mm) postorbitals, squamosal, braincase, incomplete
tooth, tooth fragments, hyoid, atlas, third cervical vertebra, fourth cervical
vertebra, fifth cervical vertebra, seventh cervical vertebra, eighth cervical
vertebra, ninth cervical vertebra, tenth cervical vertebra, cervical ribs, second
dorsal vertebra, fourth dorsal vertebra, fifth dorsal vertebra, seventh dorsal
vertebra, eighth dorsal vertebra, ninth dorsal vertebra, tenth dorsal vertebra,
dorsal ribs, gastralial fragments, five proximal caudal vertebrae, five mid
caudal vertebrae, two distal caudal vertebrae, chevron, scapulocoracoid, distal
ischium
Diagnosis- (after Filippi et al., 2016) no frontal dorsal prominences;
dorsal edge of postorbital does not expand, with jugal process anteriorly projected
and anteroposteriorly narrow; parietal depression transversally compressed;
small paraoccipital processes with ventral edge located above level of dorsal
occipital condyle edge; basioccipital-opisthotic complex, about two and a half
times width of occipital condyle and almost twice height of occipital condyle
in posterior view; high and well developed crest below the occipital condyle,
diverging laterally toward the basal tubera, defining the ventral subcondylar
recess; basisphenoid recess highly developed and deeply excavated, with subcircular
contour opening ventrally, with well-defined borders, and its major axis transversely
oriented; basipterygoid processes positioned horizontally relative to cranial
roof, located on level slightly dorsal to basal tubera; anterior surface of
cervical centra with poorly developed articular condyle; anterior projection
of cervical epipophyses, well developed between fourth and seventh; mid and
posterior cervical centra with slightly convex lateral and ventral surfaces;
hyposphene-hypanthrum articulations present from second dorsal; interspinous
accessory articulation system developed in mid and posterior dorsal vertebrae;
pair of pneumatic foramina within proximal caudal prespinal fossa; proximal
and mid caudal vertebrae with distal edge of transverse process convex, and
strong development of anterior projection; distal scapula posteriorly recurved.
Comments- The skull length is estimated from being ~22% larger than Llukalkan based on six measurements.
Filippi et al. (2016) added this to Tortosa et al.'s ceratosaur
analysis and found it to be a basal furileusaurian.
References- Filippi, Mendez, Juarez Valieri and Garrido, 2016. A new
brachyrostran with hypertrophied axial structures reveals an unexpected radiation
of latest Cretaceous abelisaurids. Cretaceous Research. 61, 209-219.
Paulina-Carabajal and Filippi, 2018 (online 2017). Neuroanatomy of the abelisaurid theropod
Viavenator: The most complete reconstruction of a cranial endocast and
inner ear for a South American representative of the clade. Cretaceous Research. 83, 84-94.
Carnotaurini Coria, Chiappe and Dingus, 2002
Definition- (Carnotaurus sastrei + Aucasaurus garridoi)
(Coria, Chiappe and Dingus, 2002)
Niebla Aranciaga Rolando, Cerroni, Garcia Marsà, Agnolín, Motta, Rozadilla, Brisson Eglí and Novas, 2020
N. antiqua Aranciaga Rolando, Cerroni, Garcia Marsà, Agnolín, Motta, Rozadilla, Brisson Eglí and Novas, 2020
Early Maastrichtian, Late Cretaceous
Allen Formation of the Neuquen Group, Rio Negro, Argentina
Holotype- (MPCN-PV-796) (~4-4.5
m; 9+ year old adult) incomplete braincase, partial dentary, teeth, two
mid-posterior dorsal centra, incomplete dorsal ribs, scapulocoracoid (~63.7 mm)
Diagnosis- (after Aranciaga
Rolando et al., 2021) strongly thickened and heavily sculptured
frontals; frontals with deep longitudinal groove surrounding the
supraorbital swelling; frontals with short bony platform for contact
with the nasal; anteroposteriorly elongate supratemporal fenestra;
parietal with posterolaterally oriented alae; nuchal wedge of
supraoccipital subequal in dorsoventral depth to the height of the
occipital condyle; associated groove of the postemporal foramen
completely enclosed by parietal and otoccipital; bony plate below the
occipital condyle subrectangular in contour and dorsoventrally taller
than transversely wide; forebrain with large olfactory tract and bulbs;
paired laminae over the mesethmoid.
Comments- The holotype was discovered in 2017 (Aranciaga Rolando pers. comm. 2020). Aranciaga Rolando et al. (2021) entered Niebla
into Filippi et al.'s ceratosaur analysis and recovered it in a
polytomy with other carnotaurins (abelisaurines in their terminology).
Reference- Aranciaga Rolando,
Cerroni, Garcia Marsà, Agnolín, Motta, Rozadilla, Brisson Eglí and
Novas, 2021 (online 2020). A new medium-sized abelisaurid (Theropoda,
Dinosauria) from the Late Cretaceous (Maastrichtian) Allen Formation of
northern Patagonia, Argentina. Journal of South American Earth
Sciences. 105, 102915.
Carnotaurinae Sereno, 1998
Definition- (Carnotaurus sastrei <- Abelisaurus comahuensis)
(Sereno et al., 2004; modified from Sereno, 1998)
Other definitions- (Carnotaurus sastrei <- Skorpiovenator
bustingorryi) (Hendrickx, Hartman and Mateus, 2015)
References- Hendrickx, Hartman and Mateus, 2015. An overview of non-avian
theropod discoveries and classification. PalArch's Journal of Vertebrate Palaeontology.
12(1), 1-73.
Carnotaurus Bonaparte, 1985
C. sastrei Bonaparte, 1985
Maastrichtian, Late Cretaceous
La Colonia Formation, Chubut, Argentina
Holotype- (MACN-CH 894) (~7.75 m, 1.5 tons) almost complete skeleton including
skull (596 mm), corpus, ceratobranchials, atlas (45 mm), axis (118 mm), third
cervical vertebra (100 mm), fourth cervical vertebra (110 mm), fifth cervical
vertebra (119 mm), sixth cervical vertebra (120 mm), seventh cervical vertebra
(110 mm), eighth cervical vertebra (108 mm), ninth cervical vertebra (104 mm),
tenth cervical vertebra (98 mm), cervical ribs 1-10, first dorsal vertebra (100
mm), second dorsal vertebra (101 mm), third dorsal vertebra (103 mm), fourth
dorsal vertebra (108 mm), fifth dorsal vertebra (101 mm), sixth dorsal vertebra
(117 mm), seventh dorsal vertebra (123 mm), eighth dorsal vertebra (122 mm),
ninth dorsal vertebra (120 mm), tenth dorsal vertebra (116 mm), eleventh dorsal
vertebra (120 mm), dorsal ribs 1-11, gastralia, first sacral vertebra (132 mm),
second sacral vertebra (115 mm), third sacral vertebra (112 mm), fourth sacral
vertebra (~98 mm), fifth sacral vertebra (~71 mm), sixth sacral vertebra (118
mm), seventh sacral vertebra (124 mm), first caudal vertebra (~128 mm), second
caudal vertebra (122 mm), third caudal vertebra (120 mm), fourth caudal vertebra
(136 mm), fifth caudal neural arch, sixth caudal neural arch, twelfth caudal
centrum, fragments of chevrons, scapulocoracoids (905, 900 mm), clavicle, sternal
plates (180 mm long), humeri (285, 284 mm), radii (73, 80 mm), ulnae (78, 85
mm), metacarpal I (30 mm), metacarpals II (37, 36 mm), partial phalanx II-1,
metacarpals III (29, 31 mm), partial phalanges III-1 (~40 mm), manual ungual
III, metacarpals IV (84 mm), two pairs of carpals or manual phalanges, ilia
(970 mm), pubes (880 mm), ischia, femora (1.03 m), proximal tibiae, skin impressions
Comments- Though originally reported as being from the Albian-Cenomanian
Gorro Frigio Formation, it is actually from much later sediments (Lamanna et
al., 2002).
References- Bonaparte, 1985. A horned Cretaceous carnosaur from Patagonia.
National Geographic Research. 1, 149-151.
Bonaparte, Novas and Coria, 1990. Carnotaurus sastrei Bonaparte, the
horned, lightly built carnosaur from the Middle Cretaceous of Patagonia. Natural
History Museum of Los Angeles County Contributions in Science. 416, 41 pp.
Bonaparte, 1991. The Gondwanian theropod families Abelisauridae and Noasauridae.
Historical Biology. 5, 1-25.
Lamanna, Martinez and Smith, 2002. A definitive abelisaurid theropod dinosaur
from the early Late Cretaceous of Patagonia. Journal of Vertebrate Paleontology.
22(1), 58-69.
Mazzetta, Cisilino and Blanco, 2004. Distribución de tensiones durante la mordida en la mandíbula de Carnotaurus sastrei Bonaparte, 1985 (Theropoda: Abelisauridae). Ameghiniana. 41(4), 605-617.
Senter and Parrish, 2006. Forelimb function in the theropod dinosaur Carnotaurus
sastrei, and its behavioral implications. PaleoBios. 26(3), 7-17.
Cisilino, Calvo, Mazzetta and Blanco, 2009. FEA for understanding the cranial
mechanics of a dinosaur from Patagonia. iacm expressions. 26, 12-17.
Paulina-Carabajal, 2009. El neurocráneo de los dinosaurios Theropoda
de la Argentina: Osteología y sus implicancias filogenéticas.
PhD thesis, Universidad Nacional de La Plata. 554 pp.
Ruiz and Novas, 2009. New insights about the anatomy of the hand of Carnotaurus
sastrei (Theropoda: Abelisauridae). Journal of Vertebrate Paleontology.
29(3), 173A.
Persons, 2010. Anatomy of a speed demon: The caudal musculature of Carnotaurus
and the implications for abelisaurid locomotion and evolutionary trajectory.
Journal of Vertebrate Paleontology. Program and Abstracts 2010, 145A.
Carabajal, 2011. The braincase anatomy of Carnotaurus sastrei (Theropoda:
Abelisauridae) from the Upper Cretaceous of Patagonia. Journal of Vertebrate
Paleontology. 31(2), 378-386.
Persons and Currie, 2011. Dinosaur speed demon: The caudal musculature of Carnotaurus
sastrei and implications for the evolution of South American abelisaurids.
PLoS ONE. 6(10), e25763.
Ruiz, Torices, Serrano and Lopez, 2011. The hand structure of Carnotaurus
sastrei (Theropoda, Abelisauridae): Implications for hand diversity and
evolution in abelisaurids. Palaeontology. 54(6), 1271-1277.
Abelisaurinae Bonaparte and Novas, 1985
sensu Paul, 1988
Definition- (Abelisaurus comahuensis <- Carnotaurus sastrei)
(modified from Sereno, 1998)
Abelisaurus Bonaparte and
Novas, 1985
A. comahuensis Bonaparte and Novas, 1985
Early Campanian or Late Campanian-Early Maastrichtian, Late Cretaceous
Allen or Anacleto Formation, Rio Negro, Argentina
Holotype- (MC 11098) (~7.3 m) incomplete skull (856 mm)
Comments- Originally described
as being from the Allen Formation near Pellegrini Lake, Heredia and
Salgado (1999) reassigned the locality to the earlier Anacleto
Formation (contested by Delaloye and Garrido, 2017).
However, Gianechini et al. (2015) reported "it is locally known that
the specimen actually comes from the Allen Formation at the Sr.
Fernandez field, close to Salitral Moreno, Río Negro Province (F.
Fernandez, pers. comm.). Since no witnesses are alive and no field
notes were taken, more information is necessary to assess the
stratigraphic provenance of this taxon."
References- Bonaparte and Novas, 1985. Abelisaurus comahuensis,
n. g., n. sp., Carnosauria from the Late Cretaceous of Patagonia. Ameghiniana.
21, 259-265.
Bonaparte, 1991. The Gondwanian theropod families Abelisauridae and Noasauridae.
Historical Biology. 5, 1-25.
Heredia and Salgado, 1999. Posicionn estratigrafica de los estratos
supracretacicos portadores de dinosaurios en el Lago Pellegrini,
Patagonia septentrional, Argentina. Ameghiniana. 36, 229-234.
Paulina-Carabajal, 2009. El neurocráneo de los dinosaurios Theropoda
de la Argentina: Osteología y sus implicancias filogenéticas.
PhD thesis, Universidad Nacional de La Plata. 554 pp.
Carabajal, 2011. Braincases of abelisaurid theropods from the Upper Cretaceous
of North Patagonia. Palaeontology. 54(4), 793-806.
Gianechini, Apesteguia, Landini, Finotti, Valieri and Zandonai, 2015. New abelisaurid
remains from the Anacleto Formation (Upper Cretaceous), Patagonia, Argentina.
Cretaceous Research. 54, 1-16.
Delaloye and Garrido, 2017. Unidad de procedencia de Abelisaurus comahuensis
Bonaparte y Novas (Dinosauria, Theropoda, Abelisauridae) y su
paleoambiente, Lago Pellegrini, provincia de Río Negro, Argentina. I
Reunión de Paleovertebrados de la Cuenca Neuquina. Ameghiniana. 54(6),
R3.
Aucasaurus Coria, Chiappe and
Dingus, 2002
= "Aucasaurus" Dingus and Chiappe, 2001
A. garridoi Coria, Chiappe and Dingus, 2002
= "Aucasaurus garridoi" Dingus and Chiappe, 2001
= Abelisaurus garridoi (Coria, Chiappe and Dingus, 2002) Paul, 2010
Early Campanian, Late Cretaceous
Anacleto Formation of Rio Colorado Subgroup, Neuquen, Argentina
Holotype- (MCF-PVPH-236) (~6.10 m, adult) skull, mandible, cervical series, cervical
ribs, dorsal series, dorsal ribs, gastralia, sacrum, first-thirteenth caudal
vertebrae, distal caudal fragments, ten chevrons, scapulocoracoid, humerus,
radius, ulna, metacarpal I, metacarpal II, phalanx II-1, metacarpal III, phalanx
III-1, phalanx III-2, metacarpal IV, pelvis, femora, tibiotarsi, fibulae, distal
tarsal III, distal tarsal IV, metatarsal I, phalanx I-1, pedal ungual I, metatarsal
II, phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III, phalanx III-1,
phalanx III-2, phalanx III-3, pedal ungual III, metatarsal IV, phalanx IV-1,
phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, metatarsal V
Diagnosis- (from Coria et al., 2002) differs from Carnotaurus
in- longer and lower rostrum and external antorbital fenestra; horizontal ventral
margin of the antorbital fenestra; complete lateral exposure of the maxillary
fenestra; frontal swells instead of horns; sigmoidal outline of the dentigerous
margin of the maxilla; less developed coracoidal process; forelimb relatively
longer; humerus with a slender and craniocaudally compressed shaft and well-defined
condyles; proximal radius lacking a hooked ulnar process; chevrons with dorsally
open haemal canals.
References- Coria, Chiappe and Dingus, 2000. A new abelisaur theropod
from the Upper Cretaceous of Patagonia. Journal of Vertebrate Paleontology.
20(3), 36A-37A.
Dingus and Chiappe, 2001. Walking on Eggs: The Astonishing Discovery of Thousands
of Dinosaur Eggs in the Badlands of Patagonia. Scribner. 219 pp.
Coria, Chiappe and Dingus 2002. A new close relative of Carnotaurus sastrei
Bonaparte 1985 (Theropoda: Abelisauridae) from the Late Cretaceous of Patagonia.
Journal of Vertebrate Paleontology. 22(2), 460-465.
Paulina-Carabajal, 2009. El neurocráneo de los dinosaurios Theropoda
de la Argentina: Osteología y sus implicancias filogenéticas.
PhD thesis, Universidad Nacional de La Plata. 554 pp.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press.
320 pp.
Carabajal, 2011. Braincases of abelisaurid theropods from the Upper Cretaceous
of North Patagonia. Palaeontology. 54(4), 793-806.
Paulina-Carabajal and Succar, 2015. The endocranial morphology and inner ear
of the abelisaurid theropod Aucasaurus garridoi. Acta Palaeontologica
Polonica. 60(1), 141-144.