Averostra Paul, 2002
Definition- (Ceratosaurus nasicornis + Allosaurus fragilis)
(Ezcurra and Cuny, 2007)
Other definitions- (promaxillary fenestra homologous with Dromaeosaurus albertensis)
(modified from Paul, 2002)
(Ceratosaurus nasicornis, Vultur gryphus <- Coelophysis bauri) (Dal Sasso, Maganuco and Cau, 2018)
= Neotheropoda sensu Padian et al., 1999
Definition- (Ceratosaurus nasicornis + Passer domesticus)
(modified)
= Neotheropoda sensu Kischlat, 2000
Definition-
(Ceratosaurus nasicornis + Allosaurus fragilis) (modified)
Comments- Paul (2002) proposed Averostra for a clade of "avepods
that either possessed at least one accessory maxillary opening in the lateral
wall of the antorbital fossa that led into a bony mediorostral maxillary sinus,
or descended from such avepods, and are members of the clade that includes the
Dromaeosauridae." This included taxa generally recognized as ceratosaurs
and tetanurines, but excluded coelophysoids. The definition is here modified
to use Dromaeosaurus albertensis (as the eponymous species for Dromaeosauridae)
and to specify the promaxillary fenestra (as it is the first accessory maxillary
opening to evolve, and the only one present in taxa Paul considers basal averostrans
such as Ceratosaurus).
Ezcurra used Averostra for the ceratosaur+tetanurine
clade in several papers, and in 2007 with Cuny gave it a new node-based phylogenetic
definition with that extent- "Ceratosaurus nasicornis, Allosaurus
fragilis, and all the descendants of their common ancestor." However,
Paul's apomorphy-based definition is not limited to that clade as promaxillary
fenestrae have been identified in Dilophosaurus, Zupaysaurus,
Panguraptor, "Syntarsus" kayentakatae, Eodromaeus, herrerasaurines, Heterodontosaurus and Sacisaurus.
Regardless, Ezcurra's definition has become the consensus and has the
advantage of ignoring the ambiguity of when the promaxillary fenestra
homologous to dromaeosaurids' evolved.
References-
Padian, Hutchinson and Holtz, 1999. Phylogenetic definitions and nomenclature
of the major taxonomic categories of the carnivorous dinosaurs Dinosauria (Theropoda).
Journal of Vertebrate Paleontology. 19(1), 69-80.
Kischlat, 2000. Tecodoncios: A aurora dos Arcosaurios no Triassico. In Holz
and De Rose (eds.). Paleontologia do Rio Grande do Sul. 273-316.
Paul, 2002. Dinosaurs of the Air. The John Hopkins University
Press. 460 pp.
Ezcurra and Cuny, 2007. The coelophysoid Lophostropheus airelensis, gen.
nov.: A review of the systematics of "Liliensternus" airelensis
from the Triassic-Jurassic outcrops of Normandy (France). Journal of Vertebrate
Paleontology. 27(1), 73-86.
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.
Sinosaurus Young, 1940
= Shuangbaisaurus Wang, You, Pan and Wang, 2017
S. triassicus Young, 1940
pr= Sinosaurus "shawanensis" Young vide [author], 1979
= Dilophosaurus “sinensis” Dong, Hisa and Azuma, 1992
= Dilophosaurus sinensis Hu, 1993
= Shuangbaisaurus anlongbaoensis Wang, You, Pan and Wang, 2017
Sinemurian, Early Jurassic
Huangchiatien, Zhangjiawa Member (Dark Red Beds) of Lufeng Formation, Yunnan, China
Holotype- (IVPP V34) partial maxilla with teeth (21x14x9,
42x24x14, 20x12x8, 41x21x13, 19x17x11, 18x12x7 mm), maxillary or
dentary fragment, three teeth (25x24x12, 15x14x9, 21x19x10 mm)
Paratypes-
?(IVPP V36) eight teeth
?(IVPP V37) tooth
Referred- ?(IVPP V73) astragalocalcaneum (157 mm trans) (Young, 1951)
Sinemurian, Early Jurassic
Ehrchuanshan, Zhangjiawa Member (Dark Red Beds) of Lufeng Formation, Yunnan, China
Paratype- ?(IVPP V48) seven teeth
Sinemurian, Early Jurassic
Hei Koas Peng, Zhangjiawa Member (Dark Red Beds) of Lufeng Formation, Yunnan, China
Referred- ?(FMNH CUP 2001) twenty-four teeth (Simmons, 1965)
?(FMNH CUP 2002) two teeth (Simmons, 1965)
?(FMNH CUP 2003) ?third dorsal centrum (Simmons, 1965)
Sinemurian, Early Jurassic
Liuchiaching, Zhangjiawa Member (Dark Red Beds) of Lufeng Formation, Yunnan, China
Paratype- ?(IVPP V35) eleven teeth
Sinemurian, Early Jurassic
Ta Ti, Zhangjiawa Member (Dark Red Beds) of Lufeng Formation, Yunnan, China
Referred- ?(FMNH CUP 2004) four teeth (Simmons, 1965)
?(FMNH CUP 2005) three teeth (Simmons, 1965)
?(FMNH CUP 2097) partial dentaries, splenial, two teeth (Simmons, 1965)
?(FMNH CUP 2098) cervical centrum, two dorsal centra (Simmons, 1965)
Sinemurian, Early Jurassic
Zhangjiawa Member (Dark Red Beds) of Lufeng Formation, Yunnan, China
?(IVPP V33) prearticular (Young, 1951)
Hettangian, Early Jurassic
Heilongtan, Shawan Member (Dull Purplish Beds) of Lufeng Formation, Yunnan, China
(ZLJT0057; = LDM-LCA 10; 'ZLJT-037' of Stiegler, 2019) skull,
incomplete skeleton including femur (590 mm), possible osteoderms,
ossified tendons (Dong, 2003)
Hettangian, Early Jurassic
Hewenzi, Shawan Member (Dull Purplish Beds) of Lufeng Formation, Yunnan, China
(ZLJT01) (immature) premaxillae fragment, incomplete maxilla, maxilla fragment,
lacrimal, frontals, parietals, incomplete braincase, incomplete dentary, atlantal
intercentrum, two dorsal rib fragments, partial proximal caudal neural arch
(Xing, 2012)
Hettangian, Early Jurassic
Konglong Hill, Shawan Member (Dull Purplish Beds) of Lufeng Formation, Yunnan, China
(ZLJ0003) partial skull, incomplete skeleton (Xing, 2012)
Hettangian, Early Jurassic
Qinglongshan, Shawan Member (Dull Purplish Beds) of Lufeng Formation, Yunnan, China
(KMV 8701; Kunming-long; Kunming theropod; holotype of Dilophosaurus sinensis) (5.5 m) skull (525 mm), lower jaw (487 mm), nine
cervical vertebrae, cervical ribs, fifteen dorsal vertebrae, four sacral vertebrae,
thirty-six caudal vertebrae, chevrons, scapulae, coracoids, distal clavicle,
humerus, radius, ulna, metacarpals, ilium, pubis, ischium, femur (587 mm), tibia,
fibula, astragalus, calcaneum, distal tarsal IV, metatarsus, pes (Anonymous, 1989; described in Hu, 1993)
three teeth (25-30 mm) (Liston et al., 2014)
Hettangian, Early Jurassic
Shawan, Shawan Member (Dull Purplish Beds) of Lufeng Formation, Yunnan, China
?(IVPP V279) tooth (Young, 1951)
Hettangian, Early Jurassic
Tachung, Shawan Member (Dull Purplish Beds) of Lufeng Formation, Yunnan, China
Paratype- ?(IVPP V38) thirteen teeth
Referred-
(IVPP V504) partial maxilla (Young, 1951)
Hettangian-Sinemurian, Early Jurassic
Lufeng Formation, Yunnan, China
?(FMNH CUP 2095) ?third dorsal centrum (Simmons, 1965)
?(FMNH CUP 2096) three tooth fragments (Simmons, 1965)
Early Jurassic
Fengjiahe Formation, Yunnan, China
(CPM C2140ZA245; holotype of Shuangbaisaurus anlongbaoensis) partial skull (540 mm), partial mandible (Wang, You, Pan and Wang, 2017)
Diagnosis-
(after Hu, 1993) vertical groove in dorsal maxillary process; skull
ornamented with two fan-shaped crests that diverge obliquely and
laterally in dorsal perspective (also in Dilophosaurus); five premaxillary teeth.
(after Carrano et al., 2012) vertical groove on lateral premaxilla
adjacent to contact with maxilla.
(after Liston et al., 2014) crown angle unusually low for FABL of 15 mm at 50-64
degrees; high denticle density on distal carina, producing DSDI of 0.62-0.88.
Other diagnoses- (after Young, 1940) teeth irregular in size, laterally compressed with fine mesial and distal
serrations, and curved
distinctly backwards.
(after Young, 1948) gigantic size; maxilla high; teeth with long,
massive root; teeth compressed, sharply pointed and curving backwards;
mesial and distal carinae with fine serrations.
Hu (1993) largely included averostran symplesiomorphies in his diagnosis of Dilophosaurus sinensis
("high, large, and robust skull ...; nasal is oblique and long ...; 13
maxillary, and 13 dentary teeth which are laterally compressed and
possess small serrations anteriorly and posteriorly. Vertebrae are
typically carnosaurian, consisting of 9 cervical, 15 dorsal ... and 36
out of a possible 45 caudals presevered. The coracoid fenestra
penetrates dorsally, scapula is narrow and long, illium is low with an
extended posterior lobe, pubis is longer than ischium...; the humerus
is half as long as the femur, and the tibia is also shorter than the
femur. The femoral fourth trochanter lies medially at a point one third
down the shaft. Metatarsals are parallel, and pes phalanges are robust
with a formula of 2�3�4�5...") The presence of four incompletely
fused sacral vertebrae is probably ontogenetic, while pneumatized limbs
and a phalanx on pedal digit V are probably errors by Hu.
Comments-
Bien (1940) stated the holotype Huangchiatien site was excavated in December 1939, with Sinosaurus then listed as "Teratosaurus-like form". Young (1940) initailly described Sinosaurus
as "a much larger Carnosauria is represented by a fragmentary part of
skull, lower jaw and many isolated teeth and limbs." He said "the
general shape of the teeth resemble that of Teratosaurus and its closely related genera." Another preliminary description is found in Young (1946)- "Sinosaurus triassicus
material includes part of a left maxilla with teeth, many isolated
teeth, and a number of articulated skeletons. The maxilla and the teeth
seem to place it very close to Teratosaurus, both in osteological features and in size." Young later (1948) noted his manuscript describing and illustrating Sinosaurus
in detail "was ready for publication at the end of 1943" but delayed
due to WWII. Paratype teeth were found in 1937-1938, but may not be
correctly referred ("since there is only one big Carnosauria found from
the Lufeng area, all the teeth belong apparently to Sinosaurus triassicus"). He described his new taxon as a teratosaurid carnosaur, though Teratosaurus is now recognized as a pseudosuchian. The supposed accessory antorbital fenestrae in both Teratosaurus and Sinosaurus
are due to damage, so that Young's proposed difference in their
comparative height is invalid to distinguish the genera. Nor is the
maxilla of Sinosaurus taller than Teratosaurus, another supposed difference proposed by Young. His final character of Sinosaurus' teeth being "rather long stretched" is hard to interpret. Actual differences from Teratosaurus
are the unfused interdental plates, shallow external surface ventral to
the antorbital fossa and more recurved teeth, all shared with KMV 8701
and ZLJT01. Young described and referred a sacrum and pelves (IVPP
V21), but this specimen is a sauropodomorph (Walker, 1964). Referred
dorsal vertebrae IVPP V30 and V31 are also probably sauropodomorph,
comparable to IVPP V100.
Additional jaw and vertebral remians were referred by Young (1951) and
Simmons (1965), though none were done so based on diagnostic characters. IVPP
V504 is comparable in antorbital fossa shape, so is probably
correctly referred.
IVPP V33 was described by Young as an angular but resembles a prearticular more closely. Young described
a partial skeleton (IVPP V100), ilium (IVPP V88) and partial femur and proximal
tibia (IVPP V97) as Sinosaurusas
well. Heerden (1977) stated both plateosaurid and melanorosaurid
material was present in these specimens. Young also described
astragalocalcaneum IVPP V73 as a carnivorous dinosaur, and postulated
either it was a hitherto unknown taxon or (at least aprtially
correctly) it "actually represents the only skeletal of the Sinosaurus triassicus and the postcranial skeleton described and tentatively referred to S. triassius as V100, V88, V21 etc., belong to a large Plateosaurid still larger than Lufengosaurus magnus."
Welles and Long (1974) considered this an example of their
'ceratosauroid' tarsus type, consisting of non-tetanuran neotheropods
(note it and Dilophosaurus have their figures switched, so that IVPP V73 is figure 6). While it plausibly belongs to Sinosaurus, the Dilophosaurus sinensis holotype does differ in lacking astragalocalcanear fusion. FMNH CUP 2097 consists of
partial mandibles described by Simmons, who stated they differed from
the holotype in having teeth without mesial serrations and with greater
labiolingual compression. Indeed, ZLJT01 has mesial serrations even on
its first dentary tooth, so FMNH CUP 2097 may be incorrectly referred.
The teeth and centra referred by Simmons have not been described or
illustrated. Until the 2000s, Sinosaurus was generally viewed as Theropoda or Archosauria indet. with no further descriptive work being done on it.
Dilophosaurus sinensis- KMV 8701 was discovered in August 1987 and was first reported in an
anonymous 1989 article in the Japanese newspaper Asahi Shinbun as
Kunming-long, as it was found in the prefecture level Kunming City and
'long' means 'dragon' in Chinese. "The article included three
photographs: one showing the dinosaur in situ, one showing the mounted
skeleton, and a third showing a closeup of the skull" (Tanimoto,
1989). Tanimoto reported this in a short piece for Olshevsky's
magazine Archosaurian Articulations, which provides information from
the newspaper source that has since been shown to be wrong- 400 mm long
skull (actually 525 mm), 52 teeth (actually 62), and "probably a single
crest" (actually the paired crests converge
posteriorly). It was
later featured in Sattler's (1990) popular book misspelled as
"Kumming-long", with the same errors and the tooth number changed to
the still incorrect 32. Tanimoto noted it was similar to Dilophosaurus, and Hu submitted its description as Dilophosaurus sinensison
June 25th 1991. According to Olshevsky (2000), Dong et al.
(1992) used the name prior to its official publication in January 1993
however. Holtz (DML, 1995) incorrectly reported that the
unofficial
name "Kunmingsaurus" "was mentioned in Archosaurian Articulations some
years back" for this specimen, but Tanimoto's article only calls it
Kunming-long or the "Kunming theropod". KMV 8701 has been
confused
with the popular mounted specimen LDM-LCA 10 discovered in 1994, which
has a longer skull
among other differences. Carrano et al. (2012) believe the differences
are ontogenetic or taphonomic, as the skulls share the the diagnostic
premaxillary groove, a vertical anterior border of the maxilla, and a
similarly shaped and positioned promaxillary fenestra. Currie et al.
(in prep.) ascribe the differences to sexual dimorphism, but this has
yet to be published. In an abstract, Currie et al. (2019) state
"crest
development, anatomy of the femoral trochanters, and ossification of
the tibiotarsus show notable differences between the two specimens.
However, these same characters have been previously described in other
coelophysoids as possible sexual differences. Given that the two
skeletons were recovered from the same stratigraphic level (Zhangjiawa
Member of the Lower Lufeng Formation) and were separated by only 80 km,
it is most parsimonious to conclude that the specimens support the
suggestion that there was strong sexual dimorphism in all
coelophysoids." Wang et al. (2017) noted "complete referred
skeleton ... (LDM-L10) ... is currently housed at World Dinosaur Valley
theme park and catalogued as ZLJT-0057." Stiegler (2019) refers
to the same specimen, saying "'D. sinensis'
(ZLJT-037) has several postaxial neural spines which are "capped" by a
dorsal expansion, but what appears to be paint over the specimen
obscures whether these expansions have the same morphology as
osteoderms in other ceratosaurs" and that it has "ossified tendons
adjacent to the lateral aspect of the posterior dorsal, sacral, and/or
anterior caudal neural spines." Xing (2012 thesis and resulting
publications) has
described two additional partial skulls, ZLJ0003 and ZLJT01, collected
in 2006 and 2007 respectively.
Shuangbaisaurus- Wang et al. (2017) describe partial skull CPM C2140ZA245 from the equivalant Fengjiahe Formation as a new taxon of theropod Shuangbaisaurus anlongbaoensis. Discovered by February 2017, this shares parasagittal crests and a vertical premaxillary groove with "Dilophosaurus" sinensis.
Supposedly distinguishing characters are more posteriorly extensive
crests, posteroventrally angled jugal and reduced supratemporal
fenestrae. Cau (online 2017) correctly noted the former two can be
corrected by realigning the broken skull, while the squamosal and
postorbital could easily be crushed against the parietal to make the
supratemporal fenestra seem small. Currie et al. (2019) concluded all
specimens including Shuangbaisaurus "fall within the range of morphological variability represented by the Kunming and Lufeng skeletons."
Synonymous? Lamanna et al. (1998) doubted sinensis is referrable to Dilophosaurus
based on differences in premaxillary shape, tooth count, lack of antorbital
tooth row, shape of infratemporal fenestra and squamosal, and size and position
of external mandibular fenestra. Dong (2003) first stated it was similar
to, and perhaps synonymous with Sinosaurus triassicus, though without
citing supporting evidence. Xing (2012) also synonymizes the taxa without evidence,
and cites Currie et al. (in prep.) as supporting the synonymy. Liston et al.
(2014) noted two dental characters which may be behind this synonymy. Most recently, Currie et al. (2019) stated "the [Sinosaurus]
holotype (especially the maxilla) was studied in detail, and there can
be no doubt that the second skeleton [LDM-LCA 10] is referable to Sinosaurus triassicus."
This may be why Wang et al. (2017) say "only the isolated teeth
referred by Young could be located in IVPP collections during the
length of this study."
"shawanensis"- Olshevsky (DML,
2002) reported that within an incomplete set of pages he had, "On p. 9
and p. 17 the paper notes from the Lufeng Formation the species
Sinosaurus shawanensis (Young) among a number of well-known dinosaur
names." He listed the reference as "Stratigraphy of China,
Jurassic System, Summary, Chinese Academy of Geological Sciences, May
1979." and attributed the name to Anonymous, as there was no indication
of the author in the pages he possessed. While such a publication
has never surfaced, an identical situation is present in Cheng (1985),
a section of "The Jurassic System of China", volume 11 in the
Stratigraphy of China series. Perhaps Olshevsky's paper was a
summary of this series printed prior to the series itself, which began
in 1982. In any case, Cheng lists "Sinosaurus shawanensis
(Young)" alongside other taxa from his layer 5 of the Dark Red Beds of
the Lufeng Formation (equivalent to layer 6 of Luo and Wu, 1994).
Notably this is the only species with the author listed in parentheses,
which would normally indicate a species named shawanensis by Young was transferred to Sinosaurus by someone else, but while Young did name Sinosaurus triassicus neither he nor anyone else named a vertebrate species shawanensis (the only animals with that species name before 1985 are brachiopod Cryptospirifer shawanensis Jing et al., 1974 and small shelly fossil Phyllochites shawanensis Duan, 1983, neither from the Lufeng Formation). Young (1951) did refer one tooth from Shawan to Sinosaurus (IVPP V279), but this was to S. triassicus
and was from the Dull Purplish Beds, so doesn't match stratigraphically
with Cheng's taxon. As the holotype and most paratypes of S. triassicus
are from the Dark Red Beds, I think Olshevsky was correct when he noted
"Perhaps it is significant that Sinosaurus triassicus is not listed,
which might mean that Sinosaurus shawanensis is a synonym", as
indeed S. triassicus is not listed by Cheng either. Thus "shawanensis" is near certainly a typo for triassicus, but is still listed here as this can probably not be proven more than thirty years after the fact with the author dead.
Note Molina-Perez and Larramendi (2019) represent Sinosaurus "shawanensis" with isolated dorsal vertebra IVPP V31, which was referred to Sinosaurus triassicus
by Young (1948). Yet its size and morphology are similar to mid
dorsals of sauropodomorph skeleton IVPP V100, also referred to S. triassicus
by Young (1951) in an example of that era's habit of combining
sauropodomorph postcrania with jaws and teeth of carnivorous
archosaurs. Contra Molina-Perez and Larramendi, it was not "More
recent than Sinosaurus triassicus", being from the Dark Red Beds as well, and has no connection to the name "shawanensis."
Relationships- Carrano et al. (2012) recovered sinensis as a non-orionidan
tetanurine, but only three more step remove it from Averostra. Ten steps
were needed to place it in Coelophysoidea however, where they recovered Dilophosaurus.
As several other relevent taxa were not included (e.g. Zupaysaurus, Dracovenator,
Sarcosaurus),
this result is questionable. Larger taxon and character samples were
used by Wang et al. (2017) and Dal Sasso et al. (2018), who recovered
it as the most basal ceratosaur and as sister to Averostra
respectively.
References- Bien, 1940. Discovery of Triassic saurisehian and primitive mammalian
remains. Bulletin of the Geological Society of China. 20(3-4), 225-234.
Young, 1940. Preliminary notes on the Lufeng vertebrate fossils. Bulletin of the Geological Society of China. 20(3-4), 235-240.
Young, 1946. The Triassic vertebrate remains of China. American Museum Novitates. 1324, 14 pp.
Young, 1948. On two new saurischians from Lufeng, Yunnan.
Bulletin of the Geological Society of China. 28, 75-90.
Young, 1951. The Lufeng saurischian fauna in China. Palaeontologica Sinica.
C(13), 1-96.
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.
Simmons, 1965. The non-therapsid reptiles of the Lufeng Basin, Yunnan, China.
Fieldiana Geology. 15, 1-93.
Welles and Long, 1974. The tarsus of theropod dinosaurs. Annals of the South African Museum. 64, 191-218.
Heerden, 1977. The comparative anatomy of the postcranial skeleton and the relationships
of the South African Melanorosauridae (Saurischia: Prosauropoda). PhD Thesis,
University of the Orange Free State. 175 pp.
[author], 1979. Stratigraphy of China, Jurassic System, Summary.
Chinese Academy of Geological Sciences. [pp].
Cheng, 1985. The Lufeng-Dafang Subregion. In Wang, Cheng and Wang (eds.).
The Jurassic System of China. Stratigraphy of China. 11, 185-189.
Anonymous, 1989. [title]. Asahi Shinbun. May 29, 1989, [pp].
Tanimoto, 1989. The complete skeleton of an Early Jurassic theropod
from Yunnan, China. Archosaurian Articulations. March 1989, 69-70.
Sattler, 1990. The New Illustrated Dinosaur Dictionary. Lothrop, Lee & Shepard Books. 363 pp.
Olshevsky, 1991. A revision of the parainfraclass Archosauria Cope,
1869, excluding the advanced Crocodylia. Mesozoic Meanderings. 2, 196
pp.
Dong, Hisa and Azuma, 1992. The Hunt for the Asian Dinosaurs. Shogakukan. 142 pp.
Hu, 1993. A new Theropoda (Dilophosaurus sinensis
sp. nov.) from Yunnan, China. Vertebrata PalAsiatica. 31(1), 65-69.
Holtz, DML 1995. https://web.archive.org/web/20210524172844/http://dml.cmnh.org/1995Feb/msg00014.html
Lamanna, Smith, You, Holtz and Dodson, 1998. A reassessment of the Chinese theropod
dinosaur Dilophosaurus sinensis. Journal of Vertebrate Paleontology.
18(3), 57A.
Olshevsky, 2000. An annotated checklist of dinosaur species by continent. Mesozoic Meanderings. 3, 157 pp.
Olshevsky, DML 2002. https://web.archive.org/web/20181215172409/http://dml.cmnh.org/2002Apr/msg00630.html
Dong, 2003. Contributions of new dinosaur materials from China to dinosaurology.
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Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
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Xing, 2012. Sinosaurus from southwestern China. Masters Thesis. University
of Alberta. 267 pp.
Xing, Bell, Rothschild, Ran, Zhang, Dong, Zhang and Currie, 2013. Tooth loss
and alveolar remodeling in Sinosaurus triassicus (Dinosauria: Theropoda)
from the Lower Jurassic strata of the Lufeng Basin, China. Chinese Science Bulletin.
58(16), 1931-1935.
Liston, Naish, Hone, Tianyang and Jian-Rong, 2014. New data on Early Jurassic
theropod diversity and feeding behavior in the Lufeng Formation of Yunnan Province,
China. Journal of Vertebrate Paleontology. Program and Abstracts 2014, 169.
Xing, Paulina-Carabajal, Currie, Xu, Zhang, Wang, Burns and Dong, 2014. Braincase
anatomy of the basal theropod Sinosaurus from the Early Jurassic of China.
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Liston, Naish, Hone, Tianyang and Jian-Rong, 2015. New data on Early Jurassic
theropod diversity and feeding behaviour in the Lufeng Formation of Yunnan Province,
China. SVPCA 2015 abstracts, 92.
Xing, Wang, Snively, Zhang, Dong, Burns and Currie, 2015. Model-based identification
of mechanical characteristics of Sinosaurus (Theropoda) crests. Acta
Geologica Sinica. 89(1), 1-11.
Cau, online 2017. http://theropoda.blogspot.com/2017/03/shuangbaisaurus-un-sinosaurus.html
Wang, Stiegler, Amiot, Wang, Du, Clark and Xu, 2017 (online 2016). Extreme ontogenetic
changes in a ceratosaurian theropod. Current Biology. 27(1), 144-148.
Wang, You, Pan and Wang, 2017. A new crested theropod dinosaur from the
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Dal Sasso, Maganuco and Cau, 2018. The oldest ceratosaurian
(Dinosauria: Theropoda), from the Lower Jurassic of Italy, sheds light
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Currie, Xing, Wu and Dong, 2019. Anatomy and relationships of Sinosaurus triassicus
(Theropoda, Coelophysoidea) from the Lufeng Formation (Lower Jurassic)
of Yunnan, China. Canadian Society of Vertebrate Palaeontology 2019
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Molina-Perez and Larramendi, 2019. Dinosaur Facts and Figures: The
Theropods and Other Dinosauriformes. Princeton University Press. 288
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Stiegler, 2019. Anatomy, systematics, and paleobiology of noasaurid
ceratosaurs from the Late Jurassic of China. PhD thesis, The George
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Currie, Xing, Wu and Dong, in prep. Anatomy and relationships of Sinosaurus
triassicus ("Dilophosaurus sinensis") from the Lufeng Formation
(Lower Jurassic) of Yunnan, China.
"Allosaurus" medius
Marsh, 1888
= Antrodemus medius (Marsh, 1888) Hay, 1902
= Dryptosaurus medius (Marsh, 1888) Gilmore, 1920
= Labrosaurus medius (Marsh, 1888) Kuhn, 1939
Late Aptian-Early Albian, Early Cretaceous
Arundel Formation, Maryland, US
Holotype- (USNM 4972) tooth (75 mm)
Referred- ?(Chas Coffin coll.) tooth (Lull, 1911)
?(Goucher College 2521; lost) pedal phalanx III-1 (110 mm) (Lull, 1911)
?(USNM 3121) tooth (Lull, 1911)
?(USNM 3446) tooth (Lull, 1911)
?(USNM 5685; not Goucher College, contra Lull) tooth (76 mm) (Lull, 1911)
?(USNM 5693) tooth (Lull, 1911)
?(USNM 8447) tooth (Lull, 1911)
?(USNM 8502; = Goucher College 2534) anterior sacral centrum (90 mm) (Lull,
1911)
?(USNM 8503; = Goucher College 2614) proximal caudal centrum (107 mm) (Lull,
1911)
?(USNM 8504; = Goucher College 2536) proximal half of pedal phalanx II-1 (Lull,
1911)
Diagnosis- Provisionally indeterminate relative to Acrocanthosaurus
atokensis.
Comments- Lipka (pers. comm.) notes the holotype tooth is almost identical
to Acrocanthosaurus atokensis, so these taxa may be synonymous. None
of the referred material can be assigned to this taxon with certainty, as it
is not comparable (postcrania) or has not been compared in detail (teeth). Lull
(1911) originally believed USNM 8502 to be a posterior dorsal centrum.
References- Marsh, 1888. Notice of a new genus of Sauropoda and other
new dinosaurs from the Potomac Formation. American Journal of Science. 35, 89-94.
Hay, 1902. Bibliography and Catalogue of the Fossil Vertebrata of North America.
Bulletin of the United States Geological Survey. 179, 868 pp.
Lull, 1911. Systematic paleontology of the Lower Cretaceous deposits of Maryland:
Vertebrata. Maryland Geological Survey. Lower Cretaceous, 183-211.
Gilmore, 1920. Osteology of the carnivorous Dinosauria in the United States
National Museum with special reference to the genera Antrodemus (Allosaurus)
and Ceratosaurus. United States National Museum Bulletin. 110, l-154.
Kuhn, 1939. Beitrage zur Keuperfauna von Halberstadt. Palaeontologische Zeitschrift.
21, 258-286.
Allosaurus? trihedrodon (Cope,
1877) Glut, 1997
= Laelaps trihedrodon Cope, 1877
= Dryptosaurus trihedrodon (Cope, 1877) Hay, 1902
= Creosaurus trigonodon (misspelling of trihedrodon) (Cope, 1877)
Osborn, 1931
= Antrodemus trihedrodon (Cope, 1877) Kuhn, 1939
= Hypsirophus trihedrodon (Cope, 1877) Cope vide Chure, 2001
Kimmeridgian-Tithonian, Late Jurassic
Brushy Basin Memberr of the Morrison Formation, Colorado, US
Holotype- (lost) dentary, eight teeth
Referred- ?(AMNH coll., lost) femur, tibia (Chure, 2001)
?(lost) skull fragments, other bones (Chure, 2001)
Comments- The holotype is lost and was not described in enough detail
to support synonymy with Allosaurus or other large Morrison theropods.
AMNH 5780 was referred to this taxon as well, but is probably an Allosaurus
specimen.
References- Cope, 1877. On a carnivorous dinosaurian from the Dakota
beds of Colorado. Bulletin of the United States Geological Survey. 3(33), 805-806.
Hay, 1902. Bibliography and Catalogue of the Fossil Vertebrata of North America.
Bulletin of the United States Geological Survey. 179, 868 pp.
Osborn, 1931. Cope: Master Naturalist. Princeton: Princeton University Press,
New York. 740 pp.
Kuhn, 1939. Beitr�ge zur Keuperfauna von Halberstadt [Contributions to
the Keuper fauna of Halberstadt]. Palaeontologische Zeitschrift. 21, 258-286.
Glut, 1997. Dinosaurs - The Encyclopedia. McFarland Press, Jefferson, North
Carolina. 1076 pp.
Chure, 2001. On the type and referred material of Laelaps trihedrodon
Cope 1877 (Dinosauria: Theropoda). In Tanke and Carpenter (eds.). Mesozoic Vertebrate
Life: New Research Inspired by the Paleontology of Philip J. Currie. Indiana
University Press. 10-18.
Calamospondylus Fox vide Anonymous,
1866
C. oweni Fox vide Anonymous, 1866
Early Barremian, Early Cretaceous
Wessex Formation, England
Holotype- (lost) sacrum (152 mm), ilial fragments
Comments- This is not the same specimen as the Aristosuchus pusillus
holotype (Naish, 2002), nor is it definitively shown to be synonymous with Aristosuchus
or Calamosaurus.
References- Anonymous, 1866. Another new Wealden reptile. The Athenaeum.
2014, 740.
Naish, 2002. The historical taxonomy of the Lower Cretaceous theropods (Dinosauria)
Calamospondylus and Aristosuchus from the Isle of Wight. Proceedings
of the Geologists' Association. 113, 153-163.
"Capitalsaurus" Kranz, 1998
"C." potens (Lull, 1911) Molina-Perez and Larramendi, 2019
= Creosaurus potens Lull, 1911
= Dryptosaurus potens (Lull, 1911) Gilmore, 1920
Late Aptian-Early Albian, Early Cretaceous
Arundel Formation, Maryland, US
Holotype- (USNM 3049) (7-10 m) proximal caudal centrum (140 mm)
Referred- ?(USNM 8505) manual ungual I (Gilmore, 1920)
Comments- The holotype was collected by J. K. Murphy in a Washington
D.C. sewer. Comparison with Allosaurus (Madsen, 1976) indicates that
the holotype is probably from the fifth or sixth caudal vertebra. Based on this,
it is probably from an animal 7-10 meters long. Although Gilmore (1920) referred
the mid-portion of a large pedal ungual I (USNM 8505) to this taxon, this was
only based on size and provenence. No valid criterion suggest such an assignment
was warrented, so it will not be discussed further.
Kranz (1998) listed "Capitalsaurus" in his Table 3 as the name for a
large theropod vertebra, with large "teeth and various isolated
postcrania" listed as "(possibly "Capitalsaurus")". The genus is
a nomen nudum as the quotation marks suggest it was not "used as valid
for a taxon when proposed" (ICZN Article 11.5), and certainly was not
"accompanied by a description or definition that states in words
characters that are purported to differentiate the taxon" (Article
13.1.1) or "accompanied by the fixation of a type species" (Article
13.3). While the vertebra was not specified in the publication,
Kranz (pers. comm. 8-6-2005) informed me it was meant as a replacement
name for Creosaurus potens. Molina-Perez and Larramendi (2019) published the combination "Capitalsaurus" potens for USNM 3049, as a megaraptorian of "Doubtful identification." While this does officially tie "Capitalsaurus" to Creosaurus potens
in the literature, the former is still a nomen nudum as the authors
left it in quotation marks and did not explicitly indicate it as a new
genus (Article 16.1).
Relationships- Comparison to other theropods is difficult due to both
the fragmentary nature of the specimen and the few detailed descriptions of
caudal centra in the literature. As "Capitalsaurus" was discovered
in Cretaceous deposits, it is assumed that the centrum did not derive from a
basal theropod such as a coelophysoid or Dilophosaurus, which are only
known from the Triassic and Early Jurassic. This species has been referred to
Allosaurus and Dryptosaurus in the past, but is stratigraphically
closest to Acrocanthosaurus. It will be compared to these three genera
first, then to other genera that may be similar. The proximal caudals of Allosaurus
are amphiplatyan to slightly procoelous, the opposite of "Capitalsaurus".
Also, they are about as wide as tall, sometimes wider, and the ventral edge
is much more concave. The ventral surface has a slight groove instead of a keel.
Those of Dryptosaurus share the straighter ventral edge and are slightly
taller than wide (~1.05 times), but no further details can be discerned. Acrocanthosaurus
has caudal pleurocoels (like Carcharodontosaurus, but not Giganotosaurus),
a concave ventral margin and amphiplatyan or amphicoelous centra. The ventral
surface is grooved and the centra are 1-1.2 times taller than wide. The only
theropod described as having opisthocoelous caudals is the segnosaur Nothronychus.
This taxon differs from "Capitalsaurus" in having a median ventral
groove, pleurocoels, an autapmorphic posterolateral tubercle, larger chevron
facets and being slightly wider (1.16 times taller than wide). Among other segnosaurs,
at least Neimongosaurus and Segnosaurus lack opisthocoelous centra.
Several theropods are known to lack ventral grooves on the proximal caudals.
These include Elaphrosaurus, Carnotaurus, Eustreptospondylus, Suchomimus,
Sinraptor dongi, "Alashansaurus", Ornithomimus? sedens
and alvarezsaurids. Of these, only alvarezsaurids are known have ventral keels,
though the condition in most others is uncertain in this regard. Although most
other theropods (eg. Ceratosaurus, Torvosaurus, Monolophosaurus, Nedcolbertia,
Sinraptor hepingensis, Tyrannosaurus, Archaeornithomimus, Gallimimus, Microvenator,
Chirostenotes) are described as having a ventral groove, the condition in
Sinraptor dongi at least changes from convex in the proximal caudals
to grooved in the mid and posterior caudals. This suggests our knowledge of
which theropods have convex ventral surfaces on their proximal caudals is extremely
limited, and subject to change as specimens are described more fully. Although
alvarezsaurids do have ventral keels, they are otherwise quite dissimilar to
"Capitalsaurus" in having strongly procoelous centra. Several theropods
are similar to "Capitalsaurus" in having centra over 1.2 times taller
than they are wide, including Monolophosaurus, sinraptorids and Bagaraatan.
Theropods known to have more circular centra are Ceratosaurus, Carnotaurus,
Elaphrosaurus, Torvosaurus, Baryonyx, Piatnitzkyosaurus, Allosaurus, Acrocanthosaurus,
Carcharodontosaurus, Dryptosaurus, ornithomimids and oviraptorosaurs (which
are diagnosed in part by their wide caudal centra). Paravians have distinctively
subrectangular centra, so "Capitalsaurus" can be excluded from this
clade. The condition found in "Capitalsaurus", where the ventral edge
of the centrum is nearly straight, is extremely rare in theropods, being otherwise
noted in Dryptosaurus, tyrannosaurids and Bagaraatan. This can
vary greatly with position in some taxa such as Bagaraatan, so undue
emphasis shouldn't be placed on the character. While clearly not a derived oviraptorosaur
or paravian, the current phylogenetic utility of proximal caudal centra does
not allow placement more precise than assumed Averostra incertae sedis.
While currently unique compared to described theropod caudals, the amount of
variation between caudal centra in single specimens is just starting to be revealed
(Sinraptor dongi's ventral groove/keel; titanosaurid's articular surfaces
varying from opisthocoelous to procoelous; Bagaraatan's ventral edge
concavity). Because of this potentially high variation, I am extremely cautious
as to the taxonomic utility of this caudal centrum and only doubtfully retain
it as a valid taxon.
References- Lull, 1911. The Reptilia of the Arundel Formation.
Maryland Geological Survey. Lower Cretaceous, 173-211.
Gilmore, 1920. Osteology of the carnivorous Dinosauria in the United States
National Museum, with special reference to the genera Antrodemus (Allosaurus)
and Ceratosaurus. United States National Museum Bulletin. 110, l-154.
Madsen, 1976. Allosaurus fragilis: A revised osteology. Utah Geological
Survey Bulletin. 109, 1-163.
Kranz, 1998. Mostly dinosaurs: A review of the vertebrates of the Potomac Group
(Aptian Arundel Formation), USA. In Lucas, Kirkland and Estep (eds.). New Mexico
Museum of Natural History and Science Bulletin. 14, 235-238.
Molina-Perez and Larramendi, 2019. Dinosaur Facts and Figures: The
Theropods and Other Dinosauriformes. Princeton University Press. 288 pp.
"Coelurosaurus" Huene, 1929
Campanian-Maastrichtian, Late Cretaceous
Allen Formation, Rio Negro, Argentina
Material- (MLP CS 1478) partial ungual (~19 mm)
Comments- Coelurosaurus was listed by Huene (1929) in a faunal
list for MLP CS 1478, a partial ungual from the Allen Formation. As Olshevsky
(DML, 1999) noted, the name is probably a typographical error for Coelurosauria
made when translating the paper from German to Spanish. This is indicated by
the fact he never attaches a name to the specimen in the description or plates
(it's described under the heading "coelurosaur claw"). Since "Coelurosaurus"
was apparently not meant as a valid name when it was published (ICZN Article
11.5), it is a nomen nudum.
The ungual consists of the distal half, which exhibits an interesting combination
of features. It is highly curved and transversely compressed, suggesting it
is a tetanurine manual ungual or a paravian pedal ungual (though note both Mapusaurus
and alvarezsaurids differ in being straighter). The cross section at midlength
is triangular (expanded ventrally), unlike the roughtly oval shape of most theropod
manual unguals (e.g. Noasaurus, Fukuiraptor, Deinonychus)
or the blade-like shape of Megaraptor's manual unguals and paravian sickle
claws. Yet the shape is not similar to most theropod pedal unguals either, as
the ventral surface is concave and the ventral transverse expansion does not
flare past the sides of the ungual. Notably, the ungual is quite asymmetric,
with one wall of the ventral groove projecting further ventrally. This is more
prominent distally, where the groove faces more to the side than downward. The
asymmetry and ventral groove are characteristic of abelisaurid pedal unguals,
though these are more straight and broad. Noasaurid pedal unguals (as judged
by Masiakasaurus) are also straight, are only slightly asymmetrical and
are keeled ventrally. Noasaurus itself possesses a controversial ungual
most recently thought to be manual which is curved and compressed like "Coelurosaurus",
but is not very asymmetrical and has a ventral keel. Another possibility is
that "Coelurosaurus" belongs to a bird, as many birds are comparably
sized with highly curved pedal unguals. Unfortunately, comparable bird unguals
(e.g. Patagopteryx, Soroavisaurus, MACN PV RN 1105) are not described
in enough detail to be usefully compared. Huene considered it to be a manual
ungual based on its curvature and believed it was a distinct specimen. It is
here referred to Averostra incertae sedis due to its Cretaceous age.
References- Huene, 1929. Los saurisquios y ornitisquios del Cretac�o
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
Diplotomodon Leidy, 1868
= Tomodon Leidy, 1865 (preoccupied Dumeril, 1853)
D. horrificus (Leidy, 1865) Leidy, 1868
= Tomodon horrificus Leidy, 1865
Maastrichtian, Late Cretaceous
Navesink or Hornerstown Formation, New Jersey, US
Holotype- (ANSP 9680; holotype of Tomodon horrificus) tooth
Comments- This taxon is often associated with Dryptosaurus aquilunguis,
following Molnar (1990). However, the teeth of the latter taxon are not distinctive
in their shape, and more detailed comparisons have yet to be made.
References- Leidy, 1865. Memoir on the extinct reptiles of the Cretaceous
formations of the United States. Smithsonian Contributions to Knowledge. 14,
1-135.
Leidy, 1868. Remarks on Conosaurus of Gibbes. Proceedings of the Academy
of Natural Sciences of Philadelphia. 20, 200-202.
Molnar, 1990. Problematic Theropoda: "Carnosaurs". In Weishampel,
Dodson and Osmolska (eds.). The Dinosauria. University of California Press. 306-317
"Elaphrosaurus" iguidiensis
Lapparent, 1960b
= Elaphrosaurus "iguidiensis" Lapparent, 1957
Albain, Early Cretaceous
Continental Intercalaire, Algeria
Syntypes- (MNHN coll.; from Alrar) tooth
(MNHN coll.; from Timimoun) incomplete mid caudal vertebra (65 mm)
Aptian, Early Cretaceous
Elrhaz Formation, Niger
Syntype- (MNHN coll.; from El Rhaz) manual ungual (30 mm)
Referred- (MNN GDF coll.) (many individuals) dozen teeth, vertebrae,
limb elements, metatarsals, phalanges (Taquet, 1976)
Bathonian-Oxfordian, Middle-Late Jurassic
Tiouraren Formation of the Irhazer Group, Niger
Syntypes- (MNHN coll.; from Ebrechko) (many individuals) thirty-one
teeth
(MNHN coll.; from Ifayen Ign�re) distal caudal vertebra (55 mm)
Cenomanian, Late Cretaceous
Echkar Formation, Niger
Syntype- (MNHN coll.; from In Abangarit) distal caudal centrum (80 mm)
Early Cenomanian, Late Cretaceous
Continental Intercalaire, Tunisia
Syntypes- (MNHN coll.; from Guermessa) tooth
(MNHN coll.; from R�mada: Kanboute) tooth
(from Dahar) three teeth (Lapparent, 1951)
Early Cenomanian, Late Cretaceous
Continental Intercalaire, Libya (Giado)
(MNHN coll.) tooth,
Berriasian-Barremian, Early Cretaceous
Irhazer Shales Group, Niger (In Tedreft)
Albian-Early Cenomanian, Early Cretaceous-Late Cretaceous
Tegama Group, Niger (Tiguidi: Zinder piste)
Early Cenomanian, Late Cretaceous
Continental Intercalaire, Tunisia (Chenini)
Syntypes- (MNHN coll.; from Chenini, Giado, In Tedreft, and Tiguidi:
Zinder piste in addition to above localities) (many individuals) fourteen teeth,
five distal caudal vertebrae (40, 40 mm; from Abangarit, Ifayen Ign�re,
and/or Timimoun), distal femur, incomplete tibia (350 mm)
Comments- Lapparent (1957) originally used Elaphrosaurus iguidiensis
in three faunal lists- Southern Tunisia (Dahar cliff including Guermessa, Chenini,
Remada and Dehibat, and Giado), Northern Sahara (including Alrar, In Akhamil,
Djoua, Aoulef and Timimoun) and Southern Sahara (including Tamesna, Agades and
Tiguidi). As no other information about iguidiensis was provided, this
name fails ICZN Article 13.1.1 ("accompanied by a description or definition
that states in words characters that are purported to differentiate the taxon")
so was a nomen nudum. Among those localities, the only potential material of
iguidiensis published earlier are three teeth "of another species
of Megalosaurus" (besides saharicus) from Dahar cliff (either
Guermessa, Chenini or Remada) mentioned by Lapparent (1951). Lapparent (1960a)
later noted caudal vertebrae and fragments of two new Elaphrosaurus species
were found in Timimoun and In Tedreft, though the species were left unnamed.
Lapparent (1960b) officially named iguidiensis, basing it on remains
from fifteen localities which may not belong to the same taxon. The locality
of numerous remains was not stated in 1960, though there were four localities
listed as having iguidiensis
material (Chenini, Giado, In Tedreft, and Tiguidi: Zinder piste) when
no material was specified as being from them. The caudals were stated
to all be from Abangarit, Ifayen Ign�re and Timimoun, so the five
unspecified caudals must come from those areas. A tooth and bone are
from Giado, so the bone is either the femur or tibia. No holotype
was designated, so all 1960b material are syntypes. Which material
should be made the lectotype is uncertain, though the species name
references Tiguidi cliff where undetermined remains from Zinder piste
were found.
Lapparent (1960b) only differentiated iguidiensis from Elaphrosaurus
bambergi due to "constantly lesser size and some accentuated differences",
and from Spinostropheus (then Elaphrosaurus) gautieri due
to smaller size and "the form of the vertebrae". The figured teeth
are recurved and labiolingually compressed with small distal serrations extending
along the entire crown and similar-sized (DSDI ~.8) mesial serrations along
the apical half in at least the figured Alrar specimen. Elongation ranges greatly
between 1.15-~2.7 times FABL. Enamel is unornamented and the crowns are not
separated from the roots by a constriction. The two illustrated caudals are
elongated (2.23-2.86 times central height) and amphicoelous to amphiplatyan
with no transverse processes. The manual ungual from El Rhaz is said to have
vascular grooves that are "situated very high and have a very different
shape from those of large theropods" and the hindlimb material is basically
undescribed. No appendicular material is illustrated. None of the published
data allows assignment to Ceratosauria, Coelurosauria, or other averostran
groups.
References- Lapparent, 1951. D�couverte de Dinosauriens, associ�s
� une faune de Reptiles et de Poissons, dans le Cr�tac�
inf�rieur de l'Extr�me Sud tunisien [Discovery of dinosaurs associated
with a reptile and fish fauna in the Lower Cretaceous of extreme southern Tunisia].
Comptes Rendus de l'Acad�mie des Sciences � Paris. 232, 1430-1432.
Lapparent, 1957. The Cretaceous dinosaurs of Africa and India. Journal of the
Palaeontological Society of India. 2, 109-112.
Lapparent, 1960a. Les dinosaures du Sahara central [The dinosaurs of the central
Sahara]. Travaux de l'Institut de Recherches Saharienne.s 19(1-2), 7-24.
Lapparent, 1960b. Les dinosauriens du "Continental intercalaire" du
Sahara central. Memoirs of the Geological Society of France. 88A, 1-57.
Taquet, 1976. G�ologie et Pal�ontologie du Gisement de Gadoufaoua
(Aptien du Niger) [Geology and Paleontology of the Gadoufaoua Locality (Aptian
of Niger)]. Cahiers de Pal�ontologie, Centre National de la Recherche
Scientifique, Paris. 191 pp.
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.
"Futabasaurus" Lambert, 1990
Coniacian, Late Cretaceous
Ashizawa Formation of the Futaba Group, Japan
Material- (unknown collection; Futaba-ryu) (~1.5-2 m) partial tibia (~56
mm wide)
Comments- This specimen was originally mentioned by Hasegawa et al. (1987)
in an abstract as Futaba-ryu, as dinosaur remains in Japan are often given nicknames
ending in "ryu" (= dragon). Lambert (1990) inappropriately made it
into a genus name, listing it as "Futabasaurus" in a childrens' book.
It was mentioned as being a large carnosaur (sensu lato) from Japan that had
yet to be described. Dong et al. (1990) referred to it as Tyrannosauridae gen.
et sp. indet. and published a photograph. Olshevsky (1991) listed it as
an allosaurid without comment, and later (DML, 2001) as a probable junior synonym
of Tarbosaurus. Chure (2000) briefly discussed and illustrated the specimen,
excluding it from Allosauridae based on a few differences from Allosaurus
(lateral edge less elongated ventrally; medial edge not rounded and drawn out
medially). While these mean "Futabasaurus" is not Allosaurus
itself, they do little to pin down its relationships further. With only a low
quality photocopy to go by, I can't make any phylogenetic judgements.
A genus of elasmosaurid plesiosaur was later named Futabasaurus by Sato
et al. (2006), making the name unavailable for the theropod tibia.
References- Hasegawa, Watanabe, Oshida, Takizawa and Koda, 1987. Dinosaur
assemblage from the Futaba Group, Fukushima. Abstract of the Annual Meeting
of the Paleontological Society of Japan. 4.
Lambert, 1990. The Dinosaur Data Book. New York, Avon Books. 320 pp.
Dong, Hasegawa and Azuma, 1990. The Age of Dinosaurs in Japan and China. Fukui,
Japan. Fukui Prefectural Museum. 65 pp.
Matsukawa and Obata, 1994. Dinosaurs and sedimentary environments, in the Japanese
Cretaceous: A contribution to dinosaur facies in Asia based on molluscan paleontology
and stratigraphy. Cretaceous Research. 15, 101-125.
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.
Sato, Hasegawa and Manabe, 2006. A new elasmosaurid plesiosaur from the Upper
Cretaceous of Fukushima, Japan. Palaeontology. 49(3), 467-484.
Inosaurus Lapparent, 1960
I. tedreftensis Lapparent, 1960
Berriasian-Barremian, Early Cretaceous
Irhazer Group, Niger
Syntype- (MNNHN coll.; from In Tedreft) two anterior dorsal vertebrae
(30 mm), two posterior dorsal vertebrae (33 mm), two sacral vertebrae, five
mid-distal caudal vertebrae (50 mm), seven caudal vertebrae fragments, proximal
tibia
Albian-Early Cenomanian, Early Cretaceous-Late Cretaceous
Tegama Group, Niger
Syntypes- ?(MNNHN coll.; from In Abangarit) partial fourth sacral vertebra
fused to fifth sacral vertebra (44 mm)
?(MNNHN coll.; from In Abangarit) proximal caudal centrum (40 mm)
?(MNNHN coll.; from In Abangarit) mid caudal vertebra (30 mm)
?(MNNHN coll.; from In Abangarit) distal caudal vertebra (12 mm)
Early Cenomanian, Late Cretaceous
Baharija Formation, Egypt
Referred- ?(IPHG 1912 VIII 63c) caudal vertebra (Stromer, 1934)
?(IPHG 1912 VIII 63e) caudal vertebra (Stromer, 1934)
?(IPHG 1912 VIII 63g) caudal vertebra (Stromer, 1934)
Comments- The associated individual from In Tedreft is the specimen Lapparent
(1960) based Inosaurus' diagnosis and species name on, so should probably
be made the lectotype if it is redescribed. He also based the taxon on three
isolated caudals and a partial sacrum from In Abangarit. They were referred
to Inosaurus because of their shorteness relative to their height. In
addition, the mid caudal was said to "present some rather close characters"
to the proximal caudal, while the sacral centrum proportions were said to be
similar. Only the proximal caudal was illustrated. Finally, Lapparent found
three caudals described by Stromer (1934) from the Baharija Formation to be
similar, without further justification. The referral of the In Abangarit and
Baharija specimens to a single taxon, let alone Inosaurus, is dubious.
References- Stromer, 1934. Ergebnisse der Forschungsreisen Prof. E. Stromers
in den W�sten �gyptens. II. Wirbeltierreste der Baharije-Stufe (unterstes
Cenoman). 13. Dinosauria. Abhandlungen der Bayerischen Akademie der Wissenschaften
Mathematisch-naturwissenschaftliche Abteilung, Neue Folge. 22, 1-79.
Lapparent, 1960. Les dinosauriens du "Continental intercalaire" du
Sahara central. Memoirs of the Geological Society of France. 88A, 1-57.
"Katsuyamasaurus" Lambert, 1990
Middle-Late Aptian, Early Cretaceous
Kitadani Formation of the Akaiwa Subgroup of the Tetori Group, Japan
Material- (unknown collection; Katsuyama-ryu) (~4 m) (?) mid caudal vertebra
(68 mm), ulna (~200 mm)
Comments- This material was informally called "Katsuyama-ryu",
as found in Azuma (1991). Lambert (1990) inappropriately made it into a genus
name, listing it as "Katsuyamasaurus" in a childrens' book. Dong et
al. (1990) published photos of the remains, labeling them Allosauridae indet..
They were later described by Chure (2000), who suggested the caudal may derive
from an ornithopod. He noted it lacks lateral pleurofossae and was reminiscent
of iguanodonts, which are known from the same quarry (Fukuisaurus, described
by Kobayashi and Azuma, 2003). Olshevsky (DML, 2000) considered "Katsuyamasaurus"
a likely junior synonym of Fukuiraptor, which was discovered later in
the same quarry. However, the ulna differs from Fukuiraptor in being
straight proximally, with a larger olecranon process and a more prominent and
proximally projecting anteroproximal process. The large olecranon process excludes
it from Maniraptoriformes, but more precise affinities within Theropoda are
unknown at this time.
References- Dong, Hasegawa and Azuma, 1990. The Age of Dinosaurs in Japan
and China. Fukui, Japan. Fukui Prefectural Museum. 65 pp.
Lambert, 1990. The Dinosaur Data Book. New York, Avon Books. 320 pp.
Azuma, 1991. Early Cretaceous Dinosaur Fauna from the Tetori Group, central
Japan. Research on Dinosaurs from the Tetori Group (1). Professor S. Miura Memorial
Volume, 55-69.
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.
Olshevsky, DML 2000. https://web.archive.org/web/20191030133242/http://dml.cmnh.org/2000Dec/msg00399.html
Kobayashi and Azuma, 2003. A new iguanodontian (Dinosauria: Ornithopoda) from
the Lower Cretaceous Kitadani Formation of Fukui Perfecture, Japan. Journal
of Vertebrate Paleontology. 23(1), 166-175.
"Labrosaurus" "huene"
Huene vide Madsen and Welles, 2000
Late Jurassic
Szechuan, China
Material- tooth
Comments- Madsen and Welles (2000) state this was a nomen nudum based
on a tooth from the Jurassic of Szechuan mentioned by Huene (1956) on page 481,
and later listed by him (1958; p. 205) as a nomen nudum.
However, the only thing Huene says about Labrosaurus in his 1956 paper
is, "Labrosaurus Marsh Teeth, Grooves similar to Ceratosaurus
from the Upper Jurassic of Wyoming, of Tendaguru and from Szechuan." This
is on the very bottom of page 481, with "Huene, Palaeontologie" in
smaller type below, which repeats every eight pages in the volume presumably
to separate it into regular sections. Similarly, in his 1958 paper, Huene only
mentions Labrosaurus on page 205, where he lists "Labrosaurus
Marsh Upper Jurassic Szechuan, China" in his list of coelurosaurs. Thus
Huene did not name Labrosaurus "huene", but did refer to Labrosaurus
teeth from Szechuan. Chure (2000) states that Huene includes Szechuan in the
distribution of Labrosaurus, "an apparent reference to medially
ridged teeth described by Young (1942)." Young only mentions Labrosaurus
stechowi on page 299, to compare it to Chienkosaurus, noting the
latter lacks lingual fluting. Chienkosaurus and Szechuanosaurus
are both listed separately by Huene, so he did not intend to sink either into
Labrosaurus. Perhaps Huene was referring to fluted teeth from Szechuan
(which could be ceratosaurid) or perhaps it was a mistake. It is clear that
the species "huene" was a mistake by Madsen and Welles (2000) and
never intended as a valid species by anyone.
References- Young, 1942. Fossil vertebrates from Kuangyuan, N. Szechuan,
China. Bulletin of the Geological Society of China. 22(3-4), 293-309.
Huene, 1956. Palaontologie und Phylogenie der Niederen Tetrapoden. Jena, Gustav
Fischer, 716 pp.
Huene, 1958. Pre-Tertiary saurians of China. Vertebrata PalAsiatica. 2(4), 201-207.
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.
"Megalosaurus"
dunkeri Dames, 1884
= Prodeinodon dunkeri (Dames, 1884) Ruiz-Omenaca and Canudo, 2003
Berriasian, Early Cretaceous
Obernkirchen Sandstein, Germany
Holotype- (UM 84; lost) tooth (60x22x? mm)
Comments- Naish (2011) reports the holotype is lost.
This species is based on a single tooth described by Dames (1884) and illustrated
by Koken (1887), initially distinguished from Megalosaurus bucklandii
in being more transversely compressed and lacking mesial serrations.
Note abundant additional material was referred to Megalosaurus dunkeri
by Dollo (1909) and Lydekker (1888), then Altispinax dunkeri by Huene
(1926), but has not been described in detail so cannot be compared to the holotypes
of either. NHMUK 2559 and 2661 became the holotype and referred specimen of Megalosaurus
oweni (Lydekker, 1889 and 1890 respectively), eventually separated as Valdoraptor.
Most influential has been Huene's (1923) proposal of the genus Altispinax
for "the species described as M. dunkeri by Lydekker (Dames)"
... "distinguished from Megalosaurus by its enormously high neural
spines in the dorsal region" (referencing three high-spined dorsal vertebrae
catalogued as NHMUK R1828), which was misunderstood by future authors as proposing
a new genus for Dames' (1884) Megalosaurus dunkeri. After decades of
both tooth and vertebrae being called Altispinax dunkeri, Paul (1888)
formally separated the vertebrae as the species altispinax, resulting
in the recent consensus of Altispinax dunkeri for the tooth and Olshevsky's
1991 new combination Becklespinax altispinax for the vertebrae. The issue
was resolved by Maisch (2016), who found the ICZN supported Altispinax dunkeri
Huene, 1923 as a taxon based on vertebrae different from Megalosaurus dunkeri
Dames, 1884 based on the tooth (see Altispinax entry for details). Thus
the German tooth is the only specimen that can be referred to Megalosaurus
dunkeri. The Hastings Beds specimens contemporaneous with Altispinax
are listed under that entry, while the Weald Clay (Belgium, England) and Lower
Greensand (England) material is listed as Averostra here. Osi et al. (2010)
referred two additional teeth from the Weald Clay.
Osi et al. (2010) found dunkeri to clade with "Megalosaurus"
pannoniensis in a morphometric study, particularly when crown angle was
compared to FABL. However, this was based on three teeth from the Weald Clay
of England, not the holotype. Ruiz-Omenaca and Canudo (2003) proposed dunkeri
was a species of Prodeinodon based on the false conclusion the mesial
carina of the latter genus is unserrated, but it is actually serrated in both
holotype and paratype (pers. obs.). The lack of mesial serrations in dunkeri
may itself be due to wear.
References- Dames, 1884. Vorlegung eines Zahnes von Megalosaurus
aus dem Wealden des Deisters. Sitzungsberichte der Gesellschaft Naturforschender
Freunde zu Berlin. 1884, 186-188.
Koken, 1887. Die Dinosaurier, Crocodiliden und Sauropterygier des norddeutschen
Wealden. Palaeontologische Abhandlungen. 3(5), 311-419.
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.
Dollo, 1909. The fossil vertebrates of Belgium. Annals of the New York Academy
of Sciences. 19(4), 99-119.
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.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
464 pp.
Olshevsky, 1991. A revision of the parainfraclass Archosauria Cope, 1869, excluding
the advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.
Ruiz-Omenaca and Canudo, 2003. A new theropod dinosaur ("Prodeinodon"
sp.) from the Lower Cretaceous of La Cantalera (Teruel, Spain). Geogaceta. 34,
111-114.
Osi, Apesteguia and Kowalewski, 2010. Non-avian theropod dinosaurs from the
early Late Cretaceous of central Europe. Cretaceous Research. 31(3), 304-320.
Naish, 2011. Theropod dinosaurs. In Batten (ed.). English Wealden Fossils. The
Palaeontological Association. 526-559.
Maisch, 2016. The nomenclatural status of the carnivorous dinosaur genus Altispinax
v. Huene, 1923 (Saurischia, Theropoda) from the Lower Cretaceous of England.
Neues Jahrbuch f�r Geologie und Pal�ontologie - Abhandlungen. 280(2),
215-219.
"Megalosaurus" insignis
Eudes-Delongchamps and Lennier vide Lennier, 1870
= Streptospondylus insignis (Eudes-Deslongchamps and Lennier vide Lennier,
1870) Dep�ret and Savornin, 1928
= Erectopus insignis (Eudes-Deslongchamps and Lennier vide Lennier, 1870)
Stromer, 1931
Early Kimmeridgian, Late Jurassic
Saint-Adresse, Cap de La H�ve, Haute-Normandie, France
Holotype- (Museum du Havre coll.) partial tooth (~120 mm)
Comments- The holotype tooth was first reported by Valenciennes (1863)
as a megalosaur, then described as Megalosaurus insignis by Lennier (1870)
and illustrated in Lennier (1887).
Sauvage (1874) referred four teeth from Fort de la Cr�che (Mus�e
de Boulogne coll.), three teeth from Portel (Beaugrand coll.), a sacral fragment
and pedal ungual from Ch�tillon (Mus�e de Boulogne coll.) to Megalosaurus
insignis. Lennier (1887) referred a pedal phalanx and pedal ungual (both
Museum du Havre coll.), but these are sauropodan (Carrano et al., 2012). He
also referred another supposed pedal phalanx (Museum du Havre coll.) and osteoderm
(Poulain coll.), which are probably not theropod. Lydekker (1888) referred a
tooth (NHMUK 46388) from the Kimmeridge Clay and another (NHMUK 35553a) from Ningle.
Parent (1893) referred a pedal ungual (Lille Natural History Museum coll.) from
Wimereux. Sauvage (1894) referred a pedal phalanx from Ch�tillon (Mus�e
de Boulogne coll.), a tooth (Beaugrand coll.) from Moulin-Wibert, teeth from
Mont-Lambert, teeth from Wimille, and remains from Pembel. Sauvage (1898) referred
a tooth from the Oxfordian of Portugal. Lapparent (1943) referred seven teeth
(MNHN coll.) from the Solvay Company quarry. Lapparent and Zbyszewski (1957)
referred numerous specimens (mostly in the Geological Services Museum of Portugal
coll.) from the Callovian-Tithonian of Portugal to M. insignis (including
the holotype of Morosaurus marchei), much of the postcrania of which
may not be theropod, with at least one caudal series being teleosaurian. As
none of the theropod remains have been justified with shared derived characters,
and the postcrania cannot be compared to the holotype, all of the material is
here removed as Averostra indet. pending further study.
References- Valenciennes, 1863. D'une esp�ce de Ch�lonien
fossile d'un genre nouveau, trouv� dans la craie du cap la H�ve
par M. Lennier, du Havre, et d�crit par M. A. Valenciennes [On a species
of fossil chelonian of a new genus, found in the chalk of Cape H�ve by
Mr. Lennier, of Havre, and described by Mr. A. Valenciennes]. Compte Rendu des
S�ances de l'Acad�mie des Sciences. 46(8), 317-322.
Lennier, 1870. �tudes G�ologiques et Pal�ontologiques sur
l'Embouchure de la Seine et les Falaises de la Haute-Normandie [Geological and
Paleontological Studies on the Mouth of the Seine and the Cliffs of Haute-Normandie].
Imprimerie Eug�ne Costey, Havre. 1-245.
Sauvage, 1874. M�moire sur les dinosauriens et les crocodiliens des terrains
jurassiques de Boulogne-sur-Mer [Memoir on the dinosaurs and crocodilians of
the Jurassic deposits of Boulogne-sur-mer]. M�moires de la Soci�t�
G�ologique de France, s�rie 2. 10(2), 1-57.
Lennier, 1887. �tudes pal�ontologiques. Description des fossiles
du Cap de la H�ve. Bulletin de la Soci�t� G�ologique
de Normandie. 12, 17-98.
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.
Parent, 1893. Le Wealdien du Bas-Boulonnais. Annales de la Societe Geologique
du Nord. 21, 50-91.
Sauvage, 1894. Les dinosauriens du terrain jurassique sup�rieur du Boulonnais.
Bulletin de la Soci�t� G�ologique de France, 3e serie.
22, 465-470.
Sauvage, 1898. Les Reptiles et les Poissons des terrains M�sozo�ques
du Portugal [The reptiles and fishes from the Mesozoic terrains of Portugal].
Bulletin de la Soci�t� G�ologique de France, 3e s�rie.
26, 442-446.
Huene, 1926a. The carnivorous Saurischia in the Jura and Cretaceous formations,
principally in Europe. Revista Museo de La Plata. 29, 35-167.
Dep�ret and Savornin, 1928. La faune de Reptiles et de Poissons albiens
de Timimoun (Sahara alg�rien) [The Albian reptile and fish fauna of Timimoun
(Algerian Sahara)]. Bulletin de la Societ� G�ologique de France,
4e s�rie. 27, 257-265.
Stromer, 1931. Ergebnisse der Forschungsreisen Prof. E. Stromers in den W�sten
�gyptens. II. Wirbeltier-Reste der Bahar�jestufe (unterstes Cenoman).
10. Ein Skelett-Rest von Carcharodontosaurus nov. gen. Abhandlungen der
Bayerischen Akademie der Wissenschaften Mathematisch-naturwissenschaftliche
Abteilung, Neue Folge. 9, 1-23.
Lapparent, 1943. Les dinosauriens jurassiques de Damparis (Jura) [The Jurassic
dinosaurs of Damparis (Jura)]. M�moires de la Soci�t� G�ologique
de France (Nouvelle S�rie). M�moire 21(47), 1-21.
Lapparent and Zbyszewski, 1957. Les dinosauriens du Portugal [The dinosaurs
of Portugal]. M�moires des Services G�ologiques du Portugal, nouvelle
s�rie. 2, 1-63.
Mateus, 1999. Upper Jurassic dinosaurs from Lourinh� and Portuguese dinosaur
- with review of collecting in Laos. Geologisk Tidskrift. 1, 33-32.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
"Megalosaurus" lonzeensis
Dollo, 1903
= Ornithomimus lonzeensis (Dollo, 1903) Kuhn, 1965
= Struthiomimus lonzeensis (Dollo, 1903) Glut, 1997
Santonian, Late Cretaceous
Glauconite of Lonzee, Belgium
Holotype- (Mus�e royal d'histoire naturelle de Belgique coll.) ungual
Comments- Dollo (1883) originally distinguished this ungual from a supposed
Megalosaurus ungual from the Weald Clay because it had a less concave
proximal articulation with a reduced median ridge, and lacked ventral striations.
Although some authors such as Russell (1972) and Molnar (1990) have stated Dollo
named the species in his 1883 paper describing the holotype, he only referred
to it as the "dinosaurien carnivore de Lonzee". He later (1903) named
it as a new species of Megalosaurus, M. lonzeensis. Huene (1923)
considered the specimen an ornithomimid without specifying why, labeling it
Ornithomim.gen.Belgium in his phylogram. In 1926, Huene referred to the taxon
as Ornithomimidorum gen. A, which has been interpreted as a genus name by some
authors (e.g. Glut, 1997). Yet it is only the Latinized way of saying "ornithomimid
genus A", to indicate Huene thought a new ornithomimid genus was necessary
for lonzeensis, but did not wish to name it. Kuhn (1965) formally transferred
the species to Ornithomimus. Russell (1972) considered Megalosaurus
lonzeensis an indeterminate possible ornithomimid, but without published
reasons. Molnar (1989 pers. comm. to Glut, 1997) believed it was not an ornithomimid,
however. Glut (1997) incorrectly stated the species had been renamed ?Struthiomimus
lonzeensis, but that combination doesn't appear in any prior work. Carrano
et al. (2012) considered it a coelurosaur manual ungual based on "small
size, mediolaterally narrow dimensions and details of the vascular traces."
If the specimen is a pedal ungual, it is most probably from a basal coelurosaur
or paravian, as other theropods have pedal unguals which are less curved and
broader. Among manual unguals, it resembles Masiakasaurus most closely,
as other theropods' are generally deeper and more strongly curved. Of Carrano
et al.'s coelurosaur-like features, noasaurids are also small and have mediolaterally
compressed unguals (though the amount has not been reported), though the blood
groove is deeper proximally and more ventral distally in Masiakasaurus.
Traditional megalosaurids like Poekilopleuron and Dubreuillosaurus
are similar to other theropods in these aspects. Ornithomimosaur pedal unguals
are straight and broader, and further differ in having lateral and medial 'spurs'
instead of a flexor tubercle. Ornithomimosaur manual unguals differ in having
a distally placed flexor tubercle (except the deep, highly curved unguals of
Deinocheirus). They are also characteristic in having a transversely
expanded area ventral to the vascular grooves which ends far from the proximal
end of the ungual. This is not seen in "Megalosaurus" lonzeensis.
These comparisons indicate the taxon is neither a basal tetanurine nor an ornithomimosaur,
but may be a noasaurid manual ungual or a deinonychosaur pedal ungual III or
IV.
References- Dollo, 1883. Note sur les restes de dinosauriens rencontr�s
dans le Cr�tac� sup�rieur de la Belgique [Note on the dinosaur
remains found in the Upper Cretaceous of Belgium]. Bulletin du Mus�e
Royal d'Histoire Naturelle de Belgique. 2, 205-221.
Dollo, 1903. Les dinosauriens de la Belgique. Comptes Rendus de l' Acad�mie
des Sciences de Paris. 136, 565-567.
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.
Kuhn, 1965. Saurischia (Supplementum 1). In Fossilium Catalogus 1. Animalia.
109, 1-94.
Russell, 1972. Ostrich dinosaurs from the Late Cretaceous of western Canada. Canadian
Journal of Earth Sciences. 9(4), 375-402.
Molnar, 1990. Problematic Theropoda: "Carnosaurs". In Weishampel,
Dodson and Osmolska (eds.). The Dinosauria. University of California Press,
Berkeley, Los Angeles, Oxford. 306-317.
Glut, 1997. Dinosaurs - The Encyclopedia. McFarland Press. 1076
pp.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Megalosaurus? monasterii
(Muenster, 1836) Windolf, 1997
= Saurocephalus monasterii Muenster, 1836
Oxfordian, Late Jurassic
Korallenkalk Formation, Hanover, Germany
Holotype- tooth
Comments- Originally referred to the saurodontid fish genus Saurocephalus
by Muenster (1846), Windolf (1997, 1998) recognized it as theropod and used
the new combination Megalosaurus monasterii. Carrano et al. (2012) stated
the tooth could not be identified past Theropoda indet..
References- Muenster, 1836. Ueber die im Korallenkalk das Lindner Berges
bei Hannover vorkommenden Ueberreste von Fischen, mit Beschreibung und Abbildung
einiger neuen Arten [On the remains of fishes occurring in the Coral Chalk of
Lindner Mountain in Hannover, with description and images of some new forms].
in Muenster and Wissman (eds.). Beitrage zur Petrefacten-Kunde. 7, 36-50.
Windolf, 1997. Theropoden-Zahne aus dem Oberen Jura Niedersachsens [Theropod
teeth from the Upper Jurassic of Niedersachsen]. In Sachs, Rauhut and Weigert
(eds.). Terra Nostra. 1. Treffen der deutschsprachigen Pal�oherpetologen.
Extended Abstracts. Duesseldorf, Germany. 33-34.
Windolf, 1998. Dinosaurierfunde in Niedersachsen [Dinosaur finds in Lower Saxony].
Arbeitskreis Pal�ontologie Hannover. 26, 1-7.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
"Megalosaurus" pombali
Lapparent and Zbyszewski, 1957
Callovian-Oxfordian, Middle-Late Jurassic
Pombal, Leiria, Portugal
Syntype- (Faculty of Sciences of Lisbon coll.?; suggested lectotype) incomplete
tooth
Middle-Late Jurassic
Ribamar, Portugal
Syntype- ?(Faculty of Sciences of Lisbon coll.?) tooth
Late Kimmeridgian-Early Tithonian, Late Jurassic
Lourinha Formation, Portugal
Syntype- ?(Faculty of Sciences of Lisbon coll.?) incomplete tooth (~110
mm)
Kimmeridgian, Late Jurassic
Torrinha (Batalha), Portugal
Syntype- ?(Faculty of Sciences of Lisbon coll.?) anterior dorsal vertebra
(90 mm)
Late Kimmeridgian, Late Jurassic
Porto de Barcas, Alcobaca Formation, Portugal
Syntypes- ?(Faculty of Sciences of Lisbon coll.) partial anterior dorsal
vertebra (85 mm), partial proximal caudal vertebra (75 mm)
?(MG coll.) two mid caudal vertebrae (160
mm), two mid caudal vertebrae (120, 130 mm)
Comments- Megalosaurus pombali was founded on several different
specimens from numerous localities in Middle-Late Jurassic Portugal. Lapparent
and Zbyszewski (1957) distinguished it from M. insignisby
its larger size (untrue), greater labiolingual thickness, and mesial
carina restricted to the apical third. The vertebrae are supposedly
united by their strong transverse and ventral constriction. As these
characters have yet to be compared to the wide variety of generic
averostran teeth now known, they are unlikely to be diagnostic, and
the remains could be from multiple taxa. No lectotype has been
selected, but the tooth from Pombal is the obvious choice given the
tooth-based diagnosis and the species' etymology. Mocho et al.
(2016) referred several of the syntype vertebrae to Sauropoda- MG 4811
from Albergaria dos Doze to Sauropoda indet.; and MG 4819, 4821 and
4826 ("A vertebra broken in two but entirely of the same type, although
slightly smaller") from Port de Barros to Diplodocinae indet..
Mocho et al. (2017) found the syntype supposed "very powerful posterior
dorsal (Pl. XIII, fig. 31, 32, 33) whose face has a somewhat less
triangular shape (Torres Vedras Museum)" (MMLT 602528) is a proximal
caudal vertebra of Sauropoda indet..
References- Lapparent and Zbyszewski, 1957. Les dinosauriens du Portugal. M�moires des Services G�ologiques
du Portugal, nouvelle s�rie. 2, 1-63.
, , and ,
2016.
Systematic review of Late Jurassic sauropods from the Museu Geol�gico collections (Lisboa, Portugal). Journal of Iberian Geology. 42, 227-250.
Mocho, Royo-Torres,
Escaso, Malafaia, de Miguel Chaves, Narv�ez, P�rez-Garc�a, Pimentel, Silva and
Ortega, 2017. Upper Jurassic sauropod record in the Lusitanian Basin
(Portugal): Geographical and lithostratigraphical distribution.
Palaeontologia Electronica. 20.2.27A: 1-50.
Megalosaurus? "tibetensis"
Zhao, 1985
Early Jurassic
Middle Daye Group, Tibet, China
Material- (IVPP coll?)
Comments- This specimen was discovered in 1976 (An et al., 2021) and first reported by Zhao (1983) who while discussing
the evolution of dinosaurs in China noted "primitive carnosaurs (Megalosaurus
Buckland)" in the Early Jurassic. It might be surmised Zhao was referring
to an undescribed Chinese specimen of Megalosaurus, which is strengthened
by the later mention of a new Megalosaurus species from the same deposits
as other Early Jurassic taxa Zhao mentions (Lufengosaurus? "changduensis",
"Damalasaurus", ?Scelidosaurus). 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) and Zhao and Cheng (1985) list the new species Megalosaurus tibetensis
from the Early Jurassic Middle Daye Group of Qamdo, Tibet. It is listed as undescribed
?megalosaurid from the Early Jurassic Daye Group of Xizang Zizhiqu by Weishampel
(1990). Zhang and Li (1997) list this theropod as being from the Middle Daye
Formation of Daye, Qamdo County, Xizang. The Daye Formation itself seems to
be Early Triassic, so a referral to a Daye Group seems more likely. Weishampel
et al. (2004) list it as undescribed theropod from the Daye Group of Xizang.
It is listed as the megalosaurid Megalosaurus tibetensis Zhao sp. nov.
(MS) in Fang et al. (2006), suggesting Zhao's monograph was indeed never published
and is still a manuscript. It is probably not referrable to Megalosaurus
based on the older age, but has not been described or figured so remains a nomen
nudum.
References- Zhao, "1983" [unpublished]. The Mesozoic vertebrate
remains of Xizang (Tibet), China. The Series of the Scientific Expeditions to
the Qinghai-Xizang Plateau. Palaeontology of Xizang. 2, 1-200.
Zhao, 1983. Phylogeny and evolutionary stages of Dinosauria. Acta Palaeontologica
Polonica. 28(1-2), 295-306.
Zhao, 1985. The Jurassic Reptilia. In Wang, Cheng and Wang (eds.). The Jurassic
System of China. Stratigraphy of China. 11, 286-289, 347, plates 10 and 11.
Zhao and Cheng, 1985. The Qamdo-Simao Subregion. In Wang, Cheng and Wang (eds.).
The Jurassic System of China. Stratigraphy of China. 11, 174-179.
Weishampel, 1990. Dinosaurian distribution. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria. University of California Press. 63-139.
Zhang and Li, 1997. Mesozoic Dinosaur Localities in China and Their Stratigraphy.
In Wolberg, Sump and Rosenberg (eds.). Dinofest International, Proceedings of
a Symposium sponsered by Arizona State University. A Publication of The Academy
of Natural Sciences. 265-273.
Olshevsky, DML 1999. https://web.archive.org/web/20200720012936/http://dml.cmnh.org/1999Nov/msg00507.html
Weishampel, Barrett, Coria, Le Loeuff, Xu, Zhao, Sahni, Gomani and Noto, 2004.
Dinosaur Distribution. In Weishampel, Dodson and Osmolska (eds.). The Dinosauria
Second Edition. University of California Press. 517-606.
Fang, Zhang, Lu, Han, Zhao and Li, 2006. Collision between the Indian Plate
and the paleo-Asian late and the appearance of Asian dinosaurs. Geological Bulletin
of China. 25(7), 862-873.
An, Wang, Li, Wang and Wang, 2021. New discovery of Jurassic dinosaur
fossils in Chaya area, Qamdu district, Tibet. Geological Bulletin of
China. 40(1), 189-193.
"Ornithocheirus" hilsensis
Koken, 1883
Berriasian, Early Cretaceous
Obernkirchen Sandstein, Germany
Holotype- distal pedal phalanx II-2 (~105-145 mm)
Diagnosis- Indeterminate within Averostra.
Comments- Koken (1883) originally identified this as a distal metacarpal
IV belonging to the pterosaur genus Ornithocheirus. Meyer and Dames (1884)
disagreed, noting that in pterosaurs the distal condyles are much larger than
the shaft and that the sides do not have ligament pits. Meyer further believed
"O." hilsensis' bone was pneumatic, which he viewed as similar
to theropods. Koken and Kayser (1885) replied, stating the morphology could
still be congruent with a pterosaurian identity. Williston (1885, 1886) firmly
supported a theropod identity, based on his observations of material at the
YPM.
Based on Koken's (1883) figure, the specimen is the distal half of a theropod
left pedal phalanx II-2. It is 59 mm long as preserved, probably ~105-145 mm
when complete based on proportions in other taxa. It is 32.3 mm broad with condyles
37 mm tall. The articular surface is deeply ginglymoid, with both extensor and
flexor grooves, the latter more extensive. There is a slight extensor pit. The
lateral condyle is broader than the medial condyle and slightly higher in distal
view, though both are equal in distal and ventral extent. The condyles are more
pointed dorsodistally than ventrodistally and possess large ligament pits on
both sides.
It can be identified as pedal phalanx II-1 because other phalanges are either
more transversely flared distally (making their distal profiles wide) and/or
more stout. Mayer and Dames were correct to note it strongly differs from pterosaurs
such as Pteranodon in having smaller condyles which have prominant ligament
pits. These condyles are also slightly dorsally displaced, unlike the strongly
ventrally displaced condyles in Pteranodon. It is extremely similar to
such taxa as Majungasaurus and Neovenator.
The size is much larger than nearly all coelurosaurs except
tyrannosauroids, meaning it is unlikely to belong to this clade, and it
does not show the dorsally displaced ligament pits of dromaeosaurids.
It may be from any variety of large theropod (neoceratosaur,
megalosauroid, carnosaur, tyrannosauroid) and shows no distinctive
characteristics, leaving it a nomen dubium.
References- Koken, 1883. Die Reptilien der norddeutschen unteren Kreide.
Zeitschrift der deutschen Geologischen Gesellshaft. 35, 735-827.
Meyer and Dames, 1884. Ueber Ornithocheirus hilsensis Koken und �ber
Zirkonzwillinge. Zeitschrift der deutschen Geologischen Gesellshaft. 36, 664-665.
Koken and Kayser, 1885. �ber Ornithocheirus hilsensis, Koken. Zeitschrift
der deutschen Geologischen Gesellshaft. 37, 214-215.
Williston, 1885. Uber Ornithocheirus hilsensis, Koken. Zoologischer Anzeiger.
8, 628-629.
Koken, 1886. Ueber Ornithocheirus hilsensis Koken. Zoologischer Anzeiger.
9, 21-23.
Williston, 1886. �ber Ornithocheirus hilsensis Koken. Zoologischer
Anzeiger. 9, 282-283.
Dames, 1886. [Comments on Meyer and Dames 1884, Koken and Kayser 1885, Williston
1885, Koken 1886 and Williston 1886]. Neues Jahrbuch fur Mineralogie, Geologie
und Palaeontologie. 1886(3), 113-114.
Orthogoniosaurus Das-Gupta,
1931
O. matleyi Das-Gupta, 1931
Late Maastrichtian, Late Cretaceous
Lameta Formation, India
Holotype- (GI coll.) posterior tooth
Comments- The type tooth of Orthogoniosaurus matleyi was discovered
in 1924.
References- Das-Gupta, 1931. On a new theropod dinosaur (Orthogoniosaurus
matleyi, n. gen. et n. sp.) from the Lameta beds of Jubbulpore. Journal
and Proceedings of the Asiatic Society of Bengal. 26, 367-369.
Huene and Matley, 1933. The Cretaceous Saurischia and Ornithischia of the Central
Provinces of India. Memoirs of the Geological Survey of India. Palaeontologica
Indica. 21, 1-72.
Romer, 1956. Osteology of the Reptiles. University of Chicago Press. 772 pp.
Walker, 1964. Triassic reptiles from the Elgin area: Ornithosuchus and
the origin of carnosaurs. Philosophical Transactions of the Royal Society of
London, Series B, Biological Sciences. 248, 53-134.
Romer, 1966. Vertebrate Paleontology, 3rd edition. University of Chicago Press,
Chicago. 468 pp.
Molnar, 1990. Problematic Theropoda: "Carnosaurs". In Weishampel,
Dodson and Osmolska (eds.). The Dinosauria. University of California Press. 306-317.
Glut, 1997. Dinosaurs - The Encyclopedia. McFarland Press. 1076
pp.
Tykoski and Rowe, 2004. Ceratosauria. In Weishampel, Dodson and Osmolska (eds.).
The Dinosauria Second Edition. University of California Press. 47-70.
Orthogoniosaurus? rawesi
(Lydekker, 1890) Olshevsky, 1991
= Massospondylus rawesi Lydekker, 1890
= Megalosaurus rawesi (Lydekker, 1890) Vianey-Liaud, Jain and Sahni,
1987
Late Maastrichtian, Late Cretaceous
Takli Formation, India
Holotype- (NHMUK R4190) tooth
Comments- Considered non-dinosaurian by Galton (pers. comm. to Glut,
1989, in Glut 1997). However, Carrano et al. (2012) considered it theropod and
noted the fine serrations and stout proportions resembled abelisaurids.
References- Lydekker, 1890. Note on certain vertebrate remains from the
Nagpur District. Records of the Geological Survey of India. 23, 20-24.
Vianey-Liaud, Jain and Sahni, 1987. Dinosaur eggshells (Saurischia) from the
Late Cretaceous intertrappeans and Lameta Formation (Deccan, India). Journal
of Vertebrate Paleontology. 7(4), 408-424.
Olshevsky, 1991. A revision of the parainfraclass Archosauria Cope, 1869, excluding
the advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.
Glut, 1997. Dinosaurs - The Encyclopedia. McFarland Press. 1076
pp.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Ozraptor Long and Molnar, 1998
= "Austroraptor" Pigdon, DML 1997
O. subotaii Long and Molnar, 1998
Bajocian, Middle Jurassic
Colalura Sandstone, Western Australia, Australia
Holotype- (UWA 82469) (~1.6-2 m) distal tibia (~170-200 mm, 40 mm wide)
Other diagnoses- The diagnostic characters listed by Long and Molnar
(1998) (high rectangular ascending process with straight dorsal margin; centrally
placed vertical ridge in astragalar facet; weakly developed medial condyle)
are also found in at least Velocisaurus and Austrocheirus. However,
differences in the extent and width of the central ridge and other features
may provide a valid diagnosis with further study.
Comments- This was originally found in 1967 and identified as chelonian
by the NHMUK. It was later repared and reidentified as theropod by Long in 1990s.
Rauhut (2005) noted Ozraptor shared a centrally placed vertical ridge
in the astragalar facet with unnamed Tendaguru abelisauroid MB R 1750, so referred
it to that clade. More recently, Rauhut (2012) found that non-ceratosaurs such
as Chuandongocoelurus, Aerosteon and Juratyrant also have
well-developed ridges, with fainter ridges present in several coelurosaurs.
Furthermore, Rajasaurus, Majungasaurus, Pycnonemosaurus,
Quilmesaurus and Masiakasaurus lack the ridge, leaving it only
known in Velocisaurus and Austrocheirus among named ceratosaurs.
Rauhut (2012) placed Ozraptor in Theropoda indet., but here it is in Averostra
incertae sedis pending further study, as no non-averostrans survived until
the Middle Jurassic or have tall astragalar ascending processes.
References- Pigdon, DML 1997. https://web.archive.org/web/20191030133231/http://dml.cmnh.org/1997Sep/msg00942.html
Long and Molnar, 1998. A new Jurassic theropod dinosaur from Western Australia.
Records of the Western Australian Museum. 19(1), 221-229.
Rauhut, 2005. Post-cranial remains of 'coelurosaurs' (Dinosauria, Theropoda)
from the Late Jurassic of Tanzania. Geological Magazine. 142(1), 97-107.
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.
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.
Paraxenisaurus Serrano-Bra�as, Espinosa-Ch�vez, Maccracken, Guti�rrez-Blando, de Le�n-D�vila and Ventura, 2020
P. normalensis Serrano-Bra�as, Espinosa-Ch�vez, Maccracken, Guti�rrez-Blando, de Le�n-D�vila and Ventura, 2020
Late Campanian, Late Cretaceous
Cerro del Pueblo Formation, Mexico
Holotype- (BENC 2/2-001) partial astragalocalcaneum, proximal phalanx I-1(?), partial metatarsal II, phalanx II-1
(115 mm), proximal phalanx II-2(?), proximal metatarsal III(?), distal metatarsal III, proximal
phalanx III-3(?), distal metatarsal IV(?), phalanx IV-1 (104 mm), phalanx
IV-3 (67 mm), phalanx IV-4 (45 mm), partial pedal ungual IV
Diagnosis- (after
Serrano-Bra�as et al., 2020) non-arctometatarsalian pes, where
proximal end of metatarsal III is expanded and has a proximally ovoid
outline (also in abelisaurids); medial condyle of metatarsal II flares
strongly medially; distinctively broad pedal unguals (also in
abelisaurids).
(proposed) posterior surface of distal quarter of metatarsal II with deep groove for m. flexor digitorum longus II tendon.
Other diagnoses- Many proposed
apomorphies of Paraxenisaurus are only scorable in paratype
specimens, which cannot be referred to the taxon as they lack
diagnostic comparable material.
The following are only scorable in BENC 1/2-0054- strongly curved and
transversely compressed manual ungual I that has
distally placed flexor tubercle divided by deep sulcus; deeply concave
proximal articular surface of manual ungual I, which is twice taller
than wide, giving it an elliptical outline; metacarpal III that has an
expanded proximal articular end, similar in width with that of
metacarpal II. The following are only scorable in BENC 1/2-0092-
distal caudal vertebrae in which dorsoventrally low prezygapophyses
with nearly vertical articular surfaces are found on the more proximal
caudals, and prezygapophyses that face ventromedially are found on the
most distal caudals.
Of the other proposed characters, the supposed attachment site for metatarsal I on metatarsal II is actually a groove for the m. flexor digitorum longus II tendon, which Serrano-Branas et al. confused for the raised scar for the m. gastrocnemius pars medialis in e.g. Garudimimus, which is also present Gallimimus
so was not related to metatarsal I. Metatarsal I actually lacks
an obvious attachment site on metatarsal II in non-bird
theropods. "Distal end of
metatarsal III is wider transversely than anteroposteriorly and has a
semi-ginglymoid articular surface" is true of most theropods.
Of supposed diagnostic pedal ungual characters, ventral
curvature is plesiomorphic and ventral angling with the
proximal end held vertically is common in theropods and present in e.g. Garudimimus and Beishanlong.
The proximodorsal process "changing its position" is using a difference
between paratype 30/2-001's mostly horizontal processes and the
intended holotype's more vertical process as a character, which
presupposes they are the same taxon. The ventral fossa
surrounding a ridge-like flexor tubercle is also present in Harpymimus, Garudimimus, Beishanlong and
large Dinosaur Park ornithomimid unguals (NMC 1349, RTMP 1967.19.145) and is not
shown in the intended holotype but is claimed to be "partially broken."
This leaves the medial foramen, which might be a valid character in
paratype ungual III and holotype ungual IV (paratype ungual II is damaged in that area), but might also be
taphonomic, as there are many other small circular areas of damage (e.g.
center of proximal surface of ungual IV). While
the two unguals in 30/2-001 are similar to each other, that of the
intended holotype is more strongly curved, has that smaller more
dorsally angled proximodorsal process, is wider in proximal view, and
lacks the expanded ventral half characteristic of ornithomimosaurs that
is present in the other specimen.
Comments- The material was
discovered in the 1990s and described in a pre-print released online on
April 24 2020. As this was electronic
and had no mention of ZooBank, it was a nomen nudum (ICZN Article
8.5.3. states names published electronically 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") until August 2020 when that issue was
published physically.
Serrano-Branas et al. (2020) referred four paratype specimens to Paraxenisaurus from different localities in the same formation- BENC
1/2-0054 consists of several manual fragments, BENC 1/2-0092 is several
distal caudal vertebrae, BENC 30/2-001 is two pedal unguals, and BENC
1/2-0091 consists of three forelimb fragments, a distal femur and
distal metatarsal. The latter element is supposed to be a
metatarsal IV, which shares no apparent characters with the distal
metatarsal in the holotype. The pedal unguals of 30/2-001 differ
from the type ungual in several characters besides a supposed medial
foramen in one, which may be taphonomic (see Other diagnoses
above). These specimens are all placed in Ornithomimidae on this
website.
The authors recovered Paraxenisaurus in Deinocheiridae with Garudimimus
using the coelurosaur matrix of Choiniere, but this dataset is plagued
by numerous misscorings. Worse yet, most of the authors' reported
deinocheirid synapomorphies don't actually match scorings in their
matrix, suggesting systemic error. For instance, of the supposed
deinocheirid synapomorphies present in the holotype- "an astragalus
with indistinct or poorly separated condyles" and "a dorsoventrally
thicker shaft of metatarsal IV in cross-section" are present in all
scored ornithomimosaurs and indeed most Mesozoic tetanurines; "a less
broad and well-rounded distal end of femur" is an averostran character
again present in most taxa and all ornithomimosaurs; "the presence of
anteroposteriorly short pedal phalanges of digit IV, with proximal and
distal articular surfaces very close together" is scored as true in all
ornithomimosaurs (and abelisaurs and tyrannosaurines among other
taxa). Only "the presence of a rectangular cross-section of metatarsal III" and "a
ventrally concave shape in lateral view of the ventral surface of pedal
unguals" would resolve as joining Paraxenisaurus with Deinocheirus
in their matrix, but are also both plesiomorphies found in e.g.
abelisaurs. The final character, "a distally-placed metatarsal I
attached to the distal quarter of metatarsal II", is misinterpreted in Paraxenisaurus as detailed above and is also scored as true for
almost all tetanurines in their matrix, so wouldn't resolve as a
deinocheirid character in any case.
Among material in the intended holotype, the supposed proximal end of
manual phalanx II-2 or III-3 is very poorly preserved but matches the
size and morphology of a pedal phalanx I-1, which would make more sense
preservationally as the rest of the material is from the distal
hindlimb. If the specimen is an abelisaurid, this would also
eliminate the objection that their manual elements are far more robust
than this. Metatarsal fragments are all very poorly preserved,
with only the distal end of III being obviously identifiable, though
two of the other fragments are distal metatarsals. The supposed
proximal end of metatarsal III is particularly broken, and interpreted
by Serrano-Branas et al. as being oval in a strictly extensor-flexor
axis unlike any other theropod. Most Late Cretaceous theropods
had transversely compressed metatarsal III proximal outlines, even
carcharodontosaurids and Deinocheirus.
In addition, these and the wider therizinosauroids have a rectangular
shape in proximal view. The only contemporaneous theropods with
similar outlines are abelisaurids like Majungasaurus,
if the long axis is tilted posteromedially and a medial tip is
added. The pedal ungual is broader than carcharodontosaurids and
coelurosaurs, but comparable to abelisaurids. If added to Hartman et al.'s maniraptoromorph matrix, it emerges as a ceratosaur closest to Aucasaurus as
far as taxa with well preserved feet are concerned, but that matrix
also doesn't include characters particular to ceratosaurs and isn't
great with pedal characters in general. Ceratosaurs are also
unknown from Cretaceous North America, though the idea of an
abelisaurid making its way from South America to Mexico isn't
completely beyond plausibility. Still, I would place the specimen
as Averostra incertae sedis pending a better description of the
tarsus and of the real bone surfaces on supposed proximal metatarsal
III.
References- Serrano-Bra�as,
Espinosa-Ch�vez, Maccracken, Guti�rrez-Blando, de Le�n-D�vila and
Ventura, 2020. Paraxenisaurus normalensis, a large deinocheirid
ornithomimosaur from the Cerro del Pueblo Formation (Upper Cretaceous),
Coahuila, Mexico. Journal of South American Earth Sciences. 101, 102610.
"Prodeinodon" kwangshiensis
Hou, Yeh and Zhao, 1975
Aptian, Early Cretaceous
Xinlong Formation, Guangxi, China
Syntypes- (IVPP V4795) tooth (73x28x13 mm), three teeth
Referred- ?(NP01) tooth (Amoit et al., 2010)
?(NP02) tooth (Amoit et al., 2010)
?(NP06) tooth (Amoit et al., 2010)
? teeth (Mo et al., 2015)
Comments- This taxon was discovered in 1973 and described as a new species
of Prodeinodon by Hou et al. (1975), though the paper has yet to be translated
from Chinese. Thus the rationale for referring kwangshiensis to Prodeinodon,
the purported diagnostic characters, and the descriptive details remain unknown
to Western authors. Note the Xinlong Formation used to be called the Napai Formation
(Mo et al., 2015). One of the four teeth described by Hou et al. is ~25-45%
larger than the others, so may belong to a different individual and make the
hypodigm more correctly termed syntypes than a holotype. Okazaki (1992) proposed
kwangshiensis might be referrable to Wakinosaurus based on the
high labiolingual compression and rather straight distal edge. Mo et al. (2014)
noted kwangshiensis differs from the Xinlong carcharodontosaurid tooth
NHMG 10858 in being smaller, more elongate (height/FABL ratio 2.61 vs. 1.92),
having apically angled distal serrations, and lacking longitudinal ridges along
the distal edge. Yet the size and elongation are more variable in e.g. Tyrannosaurus,
so these may not be important factors. Mo et al. (2016) stated "teeth generally
similar to those described by Hou, Yeh & Zhao (1975) are fairly common in
the Xinlong Formation", illustrating an example as a carcharodontosaurid.
kwangshiensis may end up being a carcharodontosaurid, and/or could be
synonymous with Datanglong from the same formation.
References- Hou, Yeh and Zhao, 1975. Fossil reptiles from Fusui, Kwangshi.
Vertebrata PalAsiatica. 13(1), 24-33.
Okazaki, 1992. A new genus and species of carnivorous dinosaur from the Lower
Cretaceous Kwanmon Group, northern Kyusyu. Bulletin of the Kitakyushu Museum
of Natural History 11, 87-90.
Amiot, Buffetaut, Lecuyer, Wang, Boudad, Ding, Fourel, Hutt, Martineau, Medeiros,
Mo, Simon, Suteethorn, Sweetman, Tong, Zhang and Zhou, 2010. Oxygen isotope
evidence for semiaquatic habits among spinosaurid theropods. Geology. 38, 139-142.
Mo, Huang, Xie and Buffetaut, 2014. A megatheropod tooth from the Early Cretaceous
of Fusui, Guangxi, southern China. Acta Geologica Sinica (English Edition).
88(1), 6-12.
Mo, Buffetaut, Tong, Amiot, Cavin, Cuny, Suteethorn, Suteethorn and Jiang, 2016 (online 2015).
Early Cretaceous vertebrates from the Xinlong Formation of Guangxi (southern
China): A review. Geological Magazine. 153(1), 143-159.
Prodeinodon Osborn, 1924
P. mongoliensis Osborn, 1924
Early Cretaceous
Huhteeg Svita (=Oshih Formation), Mongolia
Holotype- (AMNH 6265) partial tooth
Paratype- (AMNH 6531) tooth (47 mm)
Referred- ? maxillary tooth, fragmentary tibia (~1 m; lost), fragmentary
fibula (~1 m; lost) (Bohlin, 1953)
References- Osborn, 1924. Sauropoda and Theropoda of the Lower Cretaceous
of Mongolia. American Museum Novitates. 128, 1-7.
Bohlin, 1953. Fossil reptiles from Mongolia and Kansu. Reports from the Scientific
Expedition to the North-Western Provinces of China Under Leadership of Dr Sven
Hedin. The Sino-Swedish Expedition. Publication 37(6), 113 pp.
Szechuanosaurus Young, 1942
Not Szechuanosaurus-
Camp (1935) described a fragmentary specimen UCMP 32102 from the middle
Chongqing Group of Rongxian (= Jung-Hsien), Sichuan as Megalosauridae
gen. et sp. indet., but Young (1942) later wrote "The general structure
of [Szechuanosaurus campi
syntype] V236 with the way of serrations fits so well with the
Junghsien tooth, we feel that there is practically no doubt in
regarding them as identical" "and prefer to consider the Junghsien
tooth as belonging also to the new form" Szechuanosaurus. This despite previously stating UCMP 32102 "is bigger and straighter than all" Szechuanosaurus syntype teeth. Compared to Szechuanosaurus,
UCMP 32102 is indeed larger (~69 vs. ~32 and ~47 mm), with a much
greater crown height/base ratio (~314% vs. ~224% and
~267%), making it less tapered. UCMP 32102 is different from the Szechuanosaurus syntypes and is considered
Averostra incertae sedis here.
Dong et al. (1978) list Szechuanosaurus
"yandonensis" as a new species in a faunal list of taxa from the Wujiaba
quarry of the Shangshaximiao Formation. There is no description or illustration,
making this a nomen nudum. In 1983, Dong et al. note there was only a single
large theropod skeleton in the Wujiaba quarry, CV 00214, described by them as a neotype
of Szechuanosaurus campi. It can be implied that Dong et al. originally
believed CV 00214 to be a new species of Szechuanosaurus, but later decided
to include it in S. campi. Note that Dong et al.'s attempt at a neotype
designation is invalid, as the ICZN requires the original type(s) to be lost
or destroyed (Article 75.3.4) and that the new specimen "came as nearly
as practicable from the original type locality" (Article 75.3.6) whereas
CV 00214 is from a different stratigraphic group than Szechuanosaurus'
types and at least one Szechuanosaurus campi syntype still exists (Wu et al., 2020). To make CV 00214 a neotype merely due to the suggested undiagnosability
of S. campi's syntypes would require an ICZN petition (Article 75.5).
Carrano et al. (2012) found CV 00214 to be sister to Yangchuanosaurus shangyouensis
in their analysis and referred it to that species.
He (1984) briefly described a series of remains ("many carnosaur
specimens, including many teeth, cervical vertebrae, dorsal vertebrae,
more than forty caudal vertebrae, complete ischium, femur, tibia and
fibula, as well as relatively complete humerus, coracoid and claws"
[translated]) from the Shangshaximiao Formation of Hexi Commune,
Sichuan. He referred these to Szechuanosaurus campi
because the syntypes were also found in the suburbs of Guangyuan and
believed to be from the Shangshaximiao Formation based on faunal
similarities and fossil abundance, "there is no significant difference
in shape and size" between S. campi
and the Hexi teeth, and "there is no evidence of the existence of two
or more carnosaurs" from that horizon. However, the teeth of S. campi
have not been shown to be diagnostic within e.g. Metriacanthosauridae,
multiple taxa with megalosaur-grade teeth are now known from the
Shangshaximiao (Leshansaurus, Yangchuanosaurus shangyouensis, Sinraptor hepingensis), and S. campi itself may be from the Penglaizhen Formation or slightly lower Shuining
Formation instead.
Gao (1993) described ZDM 9011 (and three incomparable referred specimens) from Dashanpu in the Xiashaximiao Formation as Szechuanosaurus zigongensis based on a number of
characters shared with CV 00214 and He's Hexi Commune tetanurine
material that are largely symplesiomorphies. Notably, the only preserved teeth in S. zigongensis are in the referred specimens ZDM 9012 and 9013, so the type isn't even comparable to S. campi. Carrano et al. (2012) found zigongensis to be sister to Yangchuanosaurus
shangyouensis instead and renamed it Yangchuanosaurus zigongensis.
References-
Camp, 1935. Dinosaur remains from the province of Szechuan.
University of California Publications, Bulletin of the Department of Geological
Sciences. 23(14), 467-471.
Young, 1942. Fossil vertebrates from Kuangyuan, N. Szechuan, China. Bulletin
of the Geological Society of China. 22(3-4), 293-309.
Dong, Zhang, Li and Zhou, 1978. [A new carnosaur discovered in
Yongchuan, Sichuan]. Chinese Science Bulletin. 23(5), 302-304.
Dong, Zhou and Zhang, 1983. Dinosaurs from the Jurassic of Sichuan. Palaeontologica
Sinica. Whole Number 162, New Series C, 23, 136 pp.
He, 1984. The Vertebrate Fossils of Sichuan. Sichuan Scientific and Technical
Publishing House, Chengdu, Sichuan. 168 pp.
Gao, 1993. A new species of Szechuanosaurus from the
Middle Jurassic of Dashanpu, Zigong, Sichuan. Vertebrata PalAsiatica. 31(4),
308-314.
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.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Wu, Shi, Dong, Carr, Yi and Xu, 2020 (online 2019). A new tyrannosauroid from the
Upper Cretaceous of Shanxi, China. Cretaceous Research. 108, 104357.
S. campi Young, 1942
Tithonian?, Late Jurassic
IVPP locality 47, upper Guangyuan Group, Sichuan, China
Syntypes- (IVPP V235) partial anterior lateral tooth (?x~22x13 mm)
(IVPP V236; lost) partial anterior lateral tooth (~47x21x14 mm)
(IVPP V238A; lost) partial lateral tooth (?x17x12 mm)
(IVPP V238B; lost) partial anterior tooth
(IVPP V238C; lost) fragmentary lateral tooth (?x14x7 mm)
(IVPP V239; lost) incomplete tooth (~32x12.5x7 mm)
Bathonian-Callovian?, Middle Jurassic
IVPP locality 49, middle Guangyuan Group, Sichuan, China
Referred- ?(IVPP coll.) few teeth (Young, Bien and Mi, 1943)
Diagnosis- Provisionally indeterminate relative to at least Metriacanthosauridae.
Other diagnoses- Young's original diagnosis was - "Teeth of Megalosaurus-type,
distinctly compressed with anterior and posterior denticulations. They
are pointed and moderately curved." This describes most theropod
teeth including Megalosaurus itself, so perhaps Young intended Szechuanosaurus to be distinct purely based on geography.
Comments-
The syntype teeth were collected in late Spring 1941. Young
stated they "resemble, in many respects, those teeth described by
Janensch from Tendaguru of Africa", citing Janensch's Theropod Type A
and Type B teeth, which are megalosaur-grade teeth that have not been
analyzed recently. Note Dong et al.'s (1983) Figure 39 is
mislabeled, with 1 being the Chienkosaurus lectotype V237A and supposed Hsisosuchus
tooth V237B (not V235), 2 being V239 (not V236) and 3 being V236 (not
V238). Also note Wu et al. (2009) were incorrect to state
Szechuanosaurus "was established by Young in 1942 on the basis of four
isolated teeth", not realizing IVPP V238 is composed of pieces of three
different teeth. Wu et al.
(2020) report "we only found one tooth (IVPP V 235) in the IVPP
collections and the others are reported missing." This makes IVPP
V235 an obvious candidate to become the lectotype, although this would
need to be proposed officially and is unfortunate as V236 and 239 were
better preserved. They photograph IVPP V235 in their Figure 5F.
Young states "The denticulations of V238 and V239 are comparatively
finer and the size of them are smaller as compared with those of V235
and V236", but never gives measurements of these densities.
However, the plate would agree with the assessment as IVPP V236 has 12
serrations per 5 mm at midheight distally, IVPP V238A has 17 and IVPP
V239 has ~19 although its figure isn't clear. Wu et al. (2022)
show IVPP V235 has 8.5 serrations per 5 mm mesially at the middle or
base of the crown.
Young 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 (layer 8b in Young et al., 1943), Szechuanosaurus
may be from the Penglaizhen Formation or slightly lower Shuining
Formation. 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.
Young et al. (1943) list "Szechuanosaurus campi
Young" from layer 5b, which is presumably "a few isolated teeth"
mentioned by Young as deriving from locality 49, though they remain
undescribed. IVPP locality 49 is described as being "a few
kilometers N. of the city [Guangyuan] in the hills along the
Chengtu-Sian highway", and the highway connecting Chengdu and Xi'an (as
the cities are now called) is China National Highway 108, or G108.
References- Young, 1942. Fossil vertebrates from Kuangyuan, N. Szechuan,
China. Bulletin of the Geological Society of China. 22(3-4), 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.
Dong, Zhou and Zhang, 1983. Dinosaurs from the Jurassic of Sichuan. Palaeontologica
Sinica. Whole Number 162, New Series C, 23, 136 pp.
He, 1984. The Vertebrate Fossils of Sichuan. Sichuan Scientific and Technical
Publishing House, Chengdu, Sichuan. 168 pp.
Chure, 2000. A new species of Allosaurus from the Morrison Formation
of Dinosaur National Monument (Utah-Colorado) and a revision of the theropod
family Allosauridae. PhD thesis. Columbia University. 964 pp.
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.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Wu, Shi, Dong, Carr, Yi and Xu, 2020 (online 2019). A new tyrannosauroid from the
Upper Cretaceous of Shanxi, China. Cretaceous Research. 108, 104357.
S? sp. indet. (Dong, 1977)
Kimmeridgian-Tithonian, Late Jurassic
Nanhuxiang, Kalaza Formation, Xinjiang, China
Material- (IVPP coll.) teeth (~45x~17x? mm)
Comments- Collected between 1964 and 1966 at "Nanhu,
southeast of Qiketai" (translated), this was reported to be from the
Houyanshan Formation of the Hongshan Series (Dong, 1977).
Weishampel (1990) lists this as a synonym of the Keilozo Formation
(citing Dong, pers. comm.), which is now known as the Kalaza
Formation. Qiketai is more commonly known as Qiketaizhen (Qiketai
Town), while Nanhu is Nanhuxiang (Nanhu Township). Dong only says
"Tooth fossils collected by us can be included in this species",
referring to Szechuanosaurus campi. The
partial "Theropod tooth" in Plate II Figure 9 is presumably one of
these, and if at X1 scale is identical in FABL to S. campi
syntype IVPP
V238A and very similar in outline from what is preserved. Based
on this and the similar age, the taxonomic assignment is retained
pending detailed study.
References- Dong, 1977. On the dinosaurian remains from Turpan, Xinjiang.
Vertebrata PalAsiatica. 15(1), 59-66.
Weishampel, 1990. Dinosaurian distribution. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria. University of California Press. 63-139.
Teinurosaurus Nopcsa, 1928
= Saurornithoides Nopcsa, 1928 (preoccupied Osborn, 1924)
= Caudocoelus Huene, 1932
T. sauvagei (Huene, 1932) Olshevsky, 1978
= Caudocoelus sauvagei Huene, 1932
Tithonian, Late Jurassic
Mont-Lambert Formation, Hauts-de-France, France
Holotype- (BHN2R 240; = Boulogne Museum 500) incomplete distal caudal vertebra (75 mm)
Diagnosis- Provisionally indeterminate relative to Kaijiangosaurus, Tanycolagreus and Ornitholestes.
Other diagnoses- (after Huene, 1932; compared to Elaphrosaurus) centrum wider; narrower ventral surface; ventral median groove wider; transversely narrower prezygapophyses.
While Huene attmpted to distinguish Teinurosaurus from Elaphrosaurus,
only the wider median ventral groove is apparent in existing photos of
the former. This is compared to the one distal caudal of the
latter figured in ventral view, but as Kobayashi reports grooves become
distally narrower in Harpymimus while Ostrom reports they become distally wider in Deinonychus,
groove width is not considered taxonomically distinctive at our current
level of understanding. Indeed, this lack of data is most
relevent to both diagnosing and identifying Teinurosaurus.
Very few taxa have detailed descriptions of distal caudal vertebrae or
more than lateral views figured, let alone indications of variation
within the distal caudal series. So the facts that Fukuiraptor and Deinonychus share ventrally concave central articulations with Teinurosaurus in their single anteriorly/posteriorly figured distal caudal vertebra, or that Afromimus, "Grusimimus" and Falcarius
also have have wide ventral grooves in their few ventrally figured
distal caudals, are not considered taxonomically important.
Comments- Sauvage (1897-1898;
in a section written in January 1898) first mentioned a distal caudal
vertebra he referred to the ornithischian Iguanodon prestwichii (now recognized as the basal styracosternan Cumnoria prestwichii) - "We
are disposed to regard as belonging to the same species the caudal
vertebra of a remote region, the part which we figure under n � 7, 8"
[translated]. Note Galton (1982) was incorrect in
claiming Sauvage reported on this specimen in his 1897 paper (written
December 6), which includes a section on prestwichii
nearly identical to the 1897-1898 one but which lacks the paragraph
describing this vertebra. This could provide a specific date of
December 1897 to January 1898 for the discovery and/or recognition of
the specimen. Huene (1932) correctly noted Sauvage mislabeled
plate VII figure 8 as dorsal view, when it is in ventral view as
understood by the text. Compared to Cumnoria,
the caudal is more elongate (length 3.93 times posterior height
compared to 2.54 times at most), has a ventral median groove instead of
a keel, and the prezygapophyseal base in 71% of the anterior central
height compared to ~30-40%, all typical of averostrans. Nopcsa (1928)
recognized its theropod nature and in his list of reptile genera meant
to use a footnote to propose Teinurosaurus
as a "new name for the piece described and figured by Sauvage (Direct.
Traveaux Geol. Portugal Lisbonne 1897-1898, plate VII, Fig. 7-10) as
late caudal of Iguanodon Prestwichi." Teinurosaurus
is listed as an aublysodontine megalosaurid (not as an ornithomimine,
contra Galton), roughly equivalent to modern Eutyrannosauria.
However due to a typographical error, the footnote's superscript 1 was
placed after Saurornithoides instead of Teinurosaurus. Sauvage (1929) corrected this in an addendum- "footnote 1 does not refer to Saurornithoides (line 19 from below) but to Teinurosaurus
(last line of text)." Unfortunately, Huene missed the addendum,
and thus wrote "Nopcsa recognized in 1927 (43, p. 183) that this was a
coelurosaur and intended to give it a name, but used one already used
by Osborn, namely "Saurornithoides" (91, 1924, p. 3- 7). For this reason, a new name had to be given here" [translated]. Huene's proposed new name was Caudocoelus sauvagei, placed in Coeluridae and "somewhat reminiscent of Elaphrosaurus."
Huene is also perhaps the first of several authors to place the
specimen in the Kimmeridgian, when it is actually from the Tithonian
(Buffetaut and Martin, 1993; as Portlandian). Galton wrote
"Lapparent and Lavocat (1955: 801) gave a line drawing of the vertebra
after Sauavage (1898) and included it in the section on Elaphrosaurus" and that the specimen "was referred to Elaphrosaurus
by Lapparent and Lavocat (1955)." This was perhaps done because
Huene explicitly compared the two, ironically making it the only taxon
distinguished from Teinurosaurus at the time. Most of Huene's characters cannot be checked in the few published photos of Teinurosaurus, but the ventral median sulcus is indeed much wider than Elaphrosaurus. Ostrom (1969) was the first author to detail Nopcsa's (1929) addendum, stating "Nopcsa's name Teinurosaurus has clear piority over Huene's Caudocoelus, but since Nopcsa failed to provbide a specific name, Teinurosaurus is not valid." Olshevsky (1978) solved this by writing "Teinurosaurus has clear priority over Caudocoelus, as noted in Ostrom 1969, and it is certainly a valid generic name. The species Caudocoelus sauvagei is proposed here as the type species of the genus Teinurosaurus, resulting in the new combination Teinurosaurus sauvagei
(von Huene 1932) as the proper name of the type specimen." He
also claimed "the specimen itself, unfortunately, was destroyed during
World War II and thus must remain a nomen dubium." This was
repeated by Galton, but as Buffetaut et al. (1991) wrote- "Contrary to
a widespread opinion (expressed, for instance, by Lapparent, 1967), the
vertebra in question has survived two world wars and years of neglect,
like a large part of the other fossil reptile remains in the
collections of the Boulogne Natural History Museum (see Vadet and Rose,
1986)." Olshevsky noted Steel misunderstood Nopsca in a different
way, believing Teinurosaurus instead of Aublysodon was a "name, proposed by Cope in 1869 ... used instead of Deinodon", as stated under superscript 2. Galton did have the first modern opinion on Teinurosaurus' affinities, stating "In addition to Elaphrosaurus, elongate prezygapophyses occur in the allosaurid Allosaurus and the dromaeosaurid Deinonychus,
so this caudal vertebra can only be identified as theropod, family
incertae sedis." Buffetaut and Martin (1993) agreed, saying "no
really distinctive characters that would allow a familial assignment
can be observed." Ford (2005 online) gave the type repository as
"Dortigen Museum", but this is a misunderstanding based on Huene's
"Boulogne-sur-mer (Nr. 500 im dortigen Museum)", which
translated is "Boulogne-sur-mer (No. 500 in the museum there)",
referring to the Boulogne Museum where it has always been held.
It was originally number 500, but was recatalogued at some point.
Sauvage lists the vertebra's length as 75 mm and his plate at natural
size would have it be 79 mm, Huene lists it as 11 cm (110 mm) and his
figure at 1:2 size would have it be 152 mm. Galton's drawing with
supposed 5 cm scale would have it be 235 mm, while Buffetaut and
Martin's plate with scale would leave it at 74 mm. As Huene's and
Galton's figures are taken from Sauvage's original plate and the newest
and unique photo matches Sauvage's reported length almost exactly, 75
mm is taken as the correct length.
Relationships- While prior authors haven't specified Teinurosaurus' relationships past Theropoda (besides Lapparent and Lavocat's apparent synonymy with Elaphrosaurus),
there are several ways to narrow down its identity. Only averostrans are known from the Late Jurassic onward, so
coelophysoid-grade taxa are excluded. Some theropod clades were
too small to have a 75 mm caudal, including most non-tyrannosauroid
coelurosaurs besides ornithomimosaurs, therizinosaurs and
eudromaeosaurs. The former two are unknown from the Jurassic, and
additionally paravians like eudromaeosaurs lack any neural spine by the
time the centrum gets as elongate as Teinurosaurus (e.g. by caudal 12 in Deinonychus at elongation index of 2.4). Teinurosaurus has an elongation index (centrum length/height) of 3.9, which also excludes Ceratosauridae, Beipiaosaurus
+ therizinosauroids and oviraptorosaurs. Prezygapophyses basal
depth is significantly less in ceratosaurids, megalosaurids, carnosaurs
except Neovenator, compsognathids, Fukuivenator and Falcarius. Remaining taxa are elaphrosaur-grade ceratosaurs, piatnitzkysaurids, Neovenator and basal tyrannosauroids.
References- Sauvage, 1897. Notes sur les Reptiles Fossiles (1). Bulletin de la Soci�t� g�ologique de France. 3(25), 864-875.
Sauvage, 1897-1898. Vertebres Fossiles du Portugual, Contributions
a l'etude des poissions et des reptiles du Jurassique et du Cretaceous. Direction
des Travaux Geologiques Portugal. 1-46.
Osborn, 1924. Three new Theropoda, Protoceratops zone, central Mongolia.
American Museum Novitates. 144, 1-12.
Nopcsa, 1928. The genera of reptiles. Palaeobiologica. 1,
163-188.
Nopcsa, 1929. Addendum "The genera of reptiles". Palaeobiologica. 2, 201.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte.
Monographien zur Geologie und Palaeontologie. 4(1), 361 pp.
Lapparent and Lavocat, 1955. Dinosauriens. In Piveteau (ed.). Traite de Paleontologie. Masson et Cie. 5, 785-962.
Lapparent, 1967. Les dinosaures de France. Sciences. 51, 4-19.
Ostrom, 1969. Osteology of Deinonychus antirrhopus, an unusual theropod
from the Lower Cretaceous of Montana. Peabody Museum of Natural History Bulletin.
30, 1-165.
Steel, 1970. Part 14. Saurischia. Handbuch der Pal�oherpetologie/Encyclopedia
of Paleoherpetology. Gustav Fischer Verlag. 87 pp.
Olshevsky, 1978. The archosaurian taxa (excluding the Crocodylia). Mesozoic
Meanderings. 1, 50 pp.
Galton, 1982. Elaphrosaurus, an ornithomimid dinosaur from the Upper
Jurassic of North America and Africa. Pal�ontologische Zeitschrift. 56,
265-275.
Vadet and Rose, 1986. Catalogue commente des types et figures de dinosauriens,
ichthyosauriens, sauropterygiens, pterosauriens et cheloninens du Mus�e
d'Histoire Naturelle de Boulogne-sur-Mer.
In E. Buffetaut, Rose and Vadet (eds.). V�rt�br�s Fossiles du Boulonnais. M�moires de la Soci�t� Acad�mique du Boulonnais.
1(2), 85-97.
Rose, 1987. Redecouverte d'une vertebre caudale reptilienne
(Archosauriens) de status controverse et provenant des terrains
jurassiques superieurs du Boulonnais. Bulletin de la Soci�t� acad�mique
du Boulonnais. 1(5), 150-153.
Buffetaut, Cuny and le Loeuff, 1991. French Dinosaurs: The best record in Europe? Modern Geology. 16(1-2), 17-42.
Buffetaut and Martin, 1993. Late Jurassic dinosaurs from the Boulonnais
(northern France): A review. Revue de Pal�obiologie. 7(vol. sp�c.),
17-28.
Ford, 2005 online. http://www.paleofile.com/Dinosaurs/Theropods/Teinurosaurus.asp
Wakinosaurus Okazaki, 1992
W. satoi Okazaki, 1992
Late Hauterivian, Early Cretaceous
Sengoku Formation of the Wakino Subgroup of the Kwanmon Group, Japan
Holotype- (KMNH VP 000,016) partial tooth (~70 mm)
Comments- Okazaki (1990) originally identified the holotype as Megalosauridae
indet..
References- Okazaki, 1990. Discovery of dinosaur remain from the Kwanmon
Group. Abstract of the Annual Meeting of the Paleontological Society of Japan.
37
Okazaki, 1992. A new genus and species of carnivorous dinosaur from the Lower
Cretaceous Kwanmon Group, Northern Kyushu. Bulletin of the Kitakyushu Museum
of Natural History. 11, 87-90.
Walgettosuchus Huene, 1932
W. woodwardi Huene, 1932
= Megalosaurus woodwardi (Huene, 1932) Olshevsky, 1991
Albian, Early Cretaceous
Sandstone Member of the Griman Creek Formation, New South Wales, Australia
Holotype- (NHMUK R3717) incomplete distal caudal vertebra (63 mm)
Comments- Huene (1932) compared the apoparently long prezygapophyses (based on the
depth of their bases?) to Elaphrosaurus and Ornithomimus, both of which he viewed
as coelurosaurs. He felt it "quite possible, if not probable" that
Walgettosuchus was an ornithomimid. It has more recently been placed
in Theropoda indet., as in Agnolin et al. (2010). As Elaphrosaurus is
now considered a ceratosaur and only averostrans are known from the Cretaceous,
Walgettosuchus is here placed in Averostra pending further study.
While Olshevsky (1991) listed Megalosaurus woodwardi as a junior synonym,
he included no responsible author, nor has an earlier work using that combination
for the Walgettosuchus material (as opposed to European finds) been located.
References- Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre
Entwicklung und Geschichte. Monographien zur Geologie und Palaeontologie. 4(1),
viii + 361 pp.
Olshevsky, 1991. A revision of the parainfraclass Archosauria Cope, 1869, excluding
the advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.
Agnolin, Ezcurra, Pais and Salisbury, 2010. A reappraisal of the Cretaceous
non-avian dinosaur faunas from Australia and New Zealand: Evidence for their
Gondwanan affinities. Journal of Systematic Palaeontology. 8(2), 257-300.
Yangchuanosaurus? "longqiaoensis"
Li, Zhang and Cai, 1999
Late Jurassic
Penglaizhen Formation, Sichuan, China
Comments- This is only published as a faunal listing, so it is unknown
if the specimen is really referrable to Yangchuanosaurus.
Reference- Li, Zhang and Cai, 1999. The Characteristics of the Composition
of the Trace Elements in Jurassic Dinosaur Bones and Red Beds in Sichuan Basin.
Geological Publishing House, Beijing. 155 pp.
undescribed Averostra (Buckley, McCrea and Currie, 2005)
Middle Turonian, Late Cretaceous
Kaskapau Formation, British Columbia, Canada
Material- two teeth
References-
Buckley, McCrea and Currie, 2005. Theropod teeth from the Upper
Cretaceous Kaskapau (Middle Turonian) and the Wapiti (Upper Campanian -
Lower Maastrichtian) formations of north-eastern British Columbia,
Canada. Journal of Vertebrate Paleontology. 25(3), 40A-41A.
Rylaarsdam, Varban, Plint, Buckley and McCrea, 2006. Middle Turonian dinosaur
paleoenvironments in the Upper Cretaceous Kaskapau Formation, northeast British
Columbia. Canadian Journal of Earth Sciences. 43(6), 631-652.
unnamed averostran (Marsh, 1881)
Late Tithonian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Wyoming, US (Quarry 9)
Material- (YPM 1996) ?fifth cervical vertebra (46.8 mm)
?...(YPM 1997) incomplete ?fourth cervical vertebra (40.4 mm)
Comments- Originally assigned to Coelurus
by Marsh (1881), Ostrom (1980) noted they were from a different quarry
than the type specimens. Makovicky (1997) described them in
detail as a new taxon of coelurosaur more derived than Coelurus.
Considering similarities to small ceratosaurs like elaphrosaurines,
they are provisionally placed as Averostra incertae sedis here pending
further study.
References- Marsh, 1881. A new order of extinct Jurassic reptiles (Coeluria). American Journal
Science. 21, 339-341.
Ostrom, 1980. Coelurus and Ornitholestes: Are they the same? In
Jacobs (ed.). Aspects of Vertebrate History. Museum of Northern
Arizona Press. 245-256.
Makovicky, 1997. A new small theropod from the Morrison Formation of Como Bluff,
Wyoming. Journal of Vertebrate Paleontology. 17(4), 755-757.
undescribed Averostra (Madsen, 1976)
Tithonian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Colorado, US (Mygatt-Moore
quarry)
(MWC coll.) sixteen teeth (Kane, 2020)
Tithonian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Utah, US (Cleveland-Lloyd
quarry)
Material- (UUVP 30-727) tibia (374 mm) (Madsen, 1976)
(UUVP 40-264) femur (385 mm) (Madsen, 1976)
(UUVP 40-299) tibia (379 mm) (Madsen, 1976)
(UUVP 40-584) maxilla (Madsen, 1976)
(UUVP 509) maxilla (Madsen, 1976)
(UUVP 1599) tibia (550 mm) (Madsen, 1976)
(UUVP 1974) dentary (Madsen, 1976)
(UUVP 1987) metatarsal III (265 mm) (Madsen, 1976)
(UUVP 2308) metatarsal III (251 mm) (Madsen, 1976)
(UUVP 2584) tibia (358 mm) (Madsen, 1976)
(UUVP 2775) metatarsal III (145 mm) (Madsen, 1976)
(UUVP 2909) astragalus (Madsen, 1976)
(UUVP 2977) tibia (433 mm) (Madsen, 1976)
(UUVP 2998) humerus (280 mm) (Madsen, 1976)
(UUVP 3233) tibia (444 mm) (Madsen, 1976)
(UUVP 3872) femur (350 mm) (Madsen, 1976)
(UUVP 4909) humerus (277 mm) (Madsen, 1976)
Late Tithonian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Wyoming, US (Reed's Quarry
12)
(YPM 58268) distal ?tibia (Dalman, 2014)
(YPM coll.) proximal ?ulna (Dalman, 2014)
Kimmeridgian-Tithonian, Late Jurassic
Morrison Formation, Colorado, US
(DMNS coll.) twenty-two teeth (Kane, 2020)
Comments- The Cleveland-Lloyd specimens were listed as Unidentified by
Madsen (1976), and may belong to lesser known Morrison theropods like Stokesosaurus
and Marshosaurus. Dalman (2014) referred YPM 58268 to Allosauroidea as
a distal tibia, but the broad mediolateral expansion distally and apprent intercondylar
notch suggests it may be a distal humerus instead. Similarly, Dalman has YPM
coll. as a coelurosaurid [sic] proximal humerus, but Nair (pers. comm., 2015)
correctly notes it resembles a proximal ulna more.
References- Madsen, 1976. Allosaurus fragilis: A revised osteology.
Utah Geological and Mineral Survey Bulletin. 109, 1-163.
Dalman, 2014. New data on small theropod dinosaurs from the Upper Jurassic Morrison
Formation of Como Bluff, Wyoming, USA. Volumina Jurassica. 12(2), 181-196.
Kane, 2020. Identifying Jurassic theropod genera using GIS maps of tooth serrations. The Society
of Vertebrate Paleontology 80th
Annual Meeting, Conference Program. 197.
undescribed averostran (Hilton, 2003)
Campanian, Late Cretaceous
Chico Formation, California, US
Material- (SC VRD57) (juvenile) limb bone fragment
Reference- Hilton, 2003. Dinosaurs and Other Mesozoic Reptiles of California.
University of California Press. 312 pp.
undescribed Averostra (Chapman, Deck, Varricchio and Jackson, 2004)
Late Albian-Cenomanian, Early-Late Cretaceous
Wayan Formation, Idaho, US
Material- (IMNH 2251/49806) (juvenile) mid-posterior dorsal centrum (49.9
mm) (Krumenacker, Simon, Scofield and Varricchio, 2017)
(IMNH 2251/50850) incomplete mid caudal vertebra (Krumenacker, Simon, Scofield
and Varricchio, 2017)
Comments- Chapman et al. (2004) reported "limited, non-diagnostic
remains from a larger theropod" and "unprepared theropod fossils",
but these were presumably described by Krumenacker et al. (2017).
References- Chapman, Deck, Varricchio and Jackson, 2004. Cretaceous Wayan
Formation of Idaho: A preliminary report. 24(3), 151-152.
Krumenacker, Simon, Scofield and Varricchio, 2017 (online 2016). Theropod dinosaurs from the
Albian-Cenomanian Wayan Formation of eastern Idaho. Historical Biology. 29(2), 170-186.
undescribed Averostra (Bonde, 2008)
Albian, Early Cretaceous
Willow Tank Formation, Nevada, US
Material- teeth
Reference- Bonde, 2008. Paleoecology and taphonomy of the Willow Tank
Formation (Albian), southern Nevada. Masters thesis, Montana State University.
96 pp.
undescribed averostran (D'Emic, 2013)
Early Albian, Early Cretaceous
Paluxy Formation of the Trinity Group, Texas, US
Material- (SMU 61741) squamosal
References- D'Emic, 2013 (online 2012). Revision of the sauropod dinosaurs of the Lower
Cretaceous Trinity Group, southern USA, with the description of a new genus.
Journal of Systematic Palaeontology. 11(6), 707-726.
unnamed Averostra (DeCourten, 1991)
Early Albian, Early Cretaceous
Ruby Ranch Member of Cedar Mountain Formation, Utah, US
Material- (UUVP 904) tooth (DeCourten, 1991)
ilum (Kirkland et al., 1997)
Comments- DeCourten (1991) assigned this tooth to Acrocanthosaurus,
but Kirkland and Parrish (1995), Kirkland et al. (1997) and Harris (1998) disagreed,
citing coarser serrations (1/mm as opposed to Acrocanthosaurus' 2/mm).
Harris suggested the ilium may belong to the same individual or taxon, but noted
it cannot be compared with Acrocanthosaurus.
References- Decourten, 1990. The Long Walk Quarry: A new horizon in dinosaur
research. Canyon Legacy. 6, 15-22.
DeCourten, 1991. The Long Walk Quarry and tracksite: Unveiling the mysterious
Early Cretaceous of the Dinosaur Triangle region. In Averett (ed.). Guidebook
for dinosaur quarries and tracksites tour, Western Colorado and Eastern Utah.
Grand Junction: Grand Junction Geological Society. 19-25.
Kirkland and Parrish, 1995. Theropod teeth from the Lower and Middle Cretaceous
of Utah. Journal of Vertebrate Paleontology. 15(3), 39A.
Kirkland, Britt, Burge, Carpenter, Cifelli, DeCourten, Eaton, Hasiotis and Lawton,
1997. Lower to Middle Cretaceous dinosaur faunas of the Central Colorado Plateau:
A key to understanding 35 million years of tectonics, sedimentology, evolution,
and biogeography. Brigham Young University Geology Studies. 42, 69-103.
Harris, 1998. Large, Early Cretaceous theropods in North America. In Lucas,
Kirkland and Estep (eds.). New Mexico Museum of Natural History and Science
Bulletin. 14, 225-228.
unnamed averostran (Fiorillo and Currie, 1994)
Late Albian, Early Cretaceous
Mussentuchit Member of the Cedar Mountain Formation, Utah, US
Material- (CM 72651) tooth fragment
Comments- Fiorillo (1999) described a tooth fragment with labiolingually
elongate serrations and no blood grooves, which he felt was very similar to
specimens described as Theropod A by Fiorillo and Currie (1994).
Reference- Fiorillo and Currie, 1994. Theropod teeth from the Judith
River Formation (Upper Cretaceous) of south-central Montana. Journal of Vertebrate
Paleontology. 14, 74-80.
Fiorillo, 1999. Non-mammalian microvertebrate remains from the Robison Eggshell
site, Cedar Mountain Formation (Lower Cretaceous), Emery County, Utah. In Gillette
(ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous
Publication. 99-1, 259-268.
undescribed averostran (Templeman and Williamson, 2013)
Santonian-Early Campanian, Late Cretaceous
Menefee Formation, New Mexico, US
Material- fragmentary postcrania
Reference- Templeman and Williamson, 2013. A vertebrate fauna from the
Santonian-Lower Campanian Menefee Formation, San Juan Basin, New Mexico. Journal
of Vertebrate Paleontology. Program and Abstracts 2013, 224.
Unnamed Averostra (Jasinski, Sullivan and Lucas, 2011)
Early Maastrichtian, Late Cretaceous
Naashoibito Member of Ojo Alamo Formation, New Mexico, US
Material- (NMMNH P-28367) incomplete tooth
(NMMNH P-28369) incomplete tooth
(SMP VP-1318) incomplete caudal vertebra
(SMP VP-2069) partial ?premaxillary tooth (?x7x5 mm)
(SMP VP-2176) incomplete caudal centrum
(SMP VP-2434) ?cranial fragment
(SMP VP-2435) ?cranial fragment
(SMP VP-2500) two ?parietal fragments, braincase fragment
(SMP VP-2521) ?cranial fragment
(SMP VP-2626) (medium to large) incomplete vertebra, fragments
(SMP VP-2709) incomplete metatarsal
(SMP VP-2781) incomplete pedal ungual
(SMP VP-2788) pedal ungual
(SMP VP-3357) pedal phalanx ?III-1
Reference- Jasinski, Sullivan and Lucas, 2011. Taxonomic composition
of the Alamo Wash local fauna from the Upper Cretaceous Ojo Alamo Formation
(Naashoibito Member), San Juan Basin, New Mexico. In Sullivan, Lucas and Spielmann
(eds.). Fossil Record 3. New Mexico Museum of Natural History and Science Bulletin.
53, 216-271.
undescribed averostran (Horner, 1979)
Late Campanian, Late Cretaceous
Englishtown or Marshalltown Formation, New Jersey, US
Material- (AMNH 7624) incomplete tooth (FABL ~14 mm)
Comments- Listed as Dryptosaurus aquilunguis from "MAESTRICHTIAN: Mt.
Laurel, Navesink and New Egypt Formations" by Horner (1979), Gallagher
(1993) states the locality listed on the museum label ("About 6 mi. NW
of Freehold, N. J.") "is mapped as Englishtown-Marshalltown Fm. area"
"on the New Jersey Geological Map (Lewis and Kummel, 1910-1912)."
The AMNH online catalogue includes a photo which indicates this is a
recurved tooth ~2.6 times taller than its FABL with small serrations,
suggesting it is tyrannosauroid or dromaeosaurid considering its
stratigraphy.
References- Horner, 1979. Upper Cretaceous dinosaurs from the Bearpaw Shale (marine) of
south-central Montana with a checklist of Upper Cretaceous dinosaur
remains from marine sediments in North America. Journal of Paleontology.
53(3), 566-577.
Gallagher, 1993. The Cretaceous/Tertiary mass extinction event in the northern Atlantic coastal plain. The Mosasaur. 5, 75-154.
undescribed Averostra (Horner, 1979)
Late Campanian, Late Cretaceous
Wenonah Formation, New Jersey, US
Material- (Johnson coll.; cast MAPS A1226a) caudal vertebra
(MAPS A1210b) phalanx
Comments- Listed as Dryptosaurus aquilunguis
from "MAESTRICHTIAN: Mt.
Laurel, Navesink and New Egypt Formations" by Horner (1979), Baird
(1986) gave a "caudal (MAPS A1226a, Johnson coll.) from the Wenonah" as
a specimen that had been assigned to Ornithomimus antiquus. Without more information, whether this is tyrannosauroid, ornithomimosaurian or neither is unknown.
Horner (1979) listed MAPS A1210b from the same localities as Coelosaurus antiquus,
which seems to be the same specimen called MAPS A12106 and MAPS 12106
by Gallagher (1993). Given the notation of MAPS A1226a, it seems
likely A1210b is the correct number, and that the b was transcribed
incorrectly as a 6 by Gallagher. Gallagher states it is "listed
as Dryptosaurus? sp.
(phalanx) from Wenonah Formation." Again the similarity between
tyrannosauroid and ornithomimosaur pedal phalanges means its identity
is uncertain until it is described, especially as Gallagher referred
all reported "Coelosaurus" specimens to Dryptosaurus.
References- Horner, 1979. Upper Cretaceous dinosaurs from the Bearpaw Shale (marine) of
south-central Montana with a checklist of Upper Cretaceous dinosaur
remains from marine sediments in North America. Journal of Paleontology.
53(3), 566-577.
Baird, 1986. Upper Cretaceous reptiles from the Severn Formation of Maryland.
The Mosasaur. 3, 63-85.
Gallagher, 1993. The Cretaceous/Tertiary mass extinction event in the northern Atlantic coastal plain. The Mosasaur. 5, 75-154.
unnamed Averostra (Schwimmer, Sanders, Erickson and Weems, 2015)
Middle Campanian, Late Cretaceous
Coachman Formation, South Carolina, US
Material- (ChM PV7366) distal ?metatarsal
(ChM PV8833) limb shaft fragment
(ChM PV9149) distal ?humerus
(ChM PV9150) distal ?tibia
Comments- ChM PV7366 was listed twice in the materials list, once as
a manual phalanx. Schwimmer et al. (2015) assign the supposed humeral and tibial
ends to ?Maniraptora indet., though one condyle of the second specimen is far
more ventrally projected unlike most tibiae.
Reference- Schwimmer, Sanders, Erickson and Weems, 2015. A Late Cretaceous
dinosaur and reptile assemblage from South Carolina, USA. Transactions of the
American Philosophical Society. 105(2), 157 pp.
undescribed Averostra (Ramirez-Valesco and Hernandez-Rivera, 2015)
Late Cretaceous
unknown formation, Arenales, Mexico
Material- (uncollected) vertebrae
(uncollected) caudal vertebrae
Late Cretaceous
unknown formation, Palau, Mexico
(IGM coll.) tooth
Reference- Ramirez-Valesco and Hernandez-Rivera, 2015. Diversity of Late
Cretaceous dinosaurs from Mexico. Bolet�n Geol�gico y Minero.
126(1), 63-108.
undescribed averostran (Ramirez-Valesco and Hernandez-Rivera, 2015)
Campanian, Late Cretaceous
San Carlos Formation, Mexico
Material- (IGM coll.) phalanx
Reference- Ramirez-Valesco and Hernandez-Rivera, 2015. Diversity of Late
Cretaceous dinosaurs from Mexico. Bolet�n Geol�gico y Minero.
126(1), 63-108.
unnamed Averostra (Torres-Rodriguez, 2006)
Late Campanian, Late Cretaceous
Aguja Formation, Mexico
Material- (IGM 6213) tooth (Torres-Rodriguez, 2006)
(IGM coll.) tooth fragment (Monroy-Mujica, 2009)
(IGM coll.) tooth (Monroy-Mujica, 2009)
(IGM coll.) tooth (Monroy-Mujica, 2009)
(IGM coll.) tooth (Ramirez-Valesco and Hernandez-Rivera, 2015)
(IGM coll.) phalanx (Ramirez-Valesco and Hernandez-Rivera, 2015)
References- Torres-Rodr�guez, 2006. Ter�podos del Cret�cico
Superior del Estado de Coahuila, M�xico. MS thesis, Universidad Nacional
Aut�noma de M�xico. 91 pp.
Monroy-M�jica, 2009. Microvertebrados f�siles Cret�cicos
Tard�os (Campaniano Tard�o) de la Formaci�n Aguja en el
Noroeste de Coahuila, M�xico. Tesis Licenciatura. Universidad Nacional
Aut�noma de M�xico. 111 pp.
Ramirez-Valesco and Hernandez-Rivera, 2015. Diversity of Late Cretaceous dinosaurs
from Mexico. Bolet�n Geol�gico y Minero. 126(1), 63-108.
unnamed Averostra (Rodriguez de la Rosa and Cevallos-Ferriz, 1998)
Late Campanian, Late Cretaceous
Cerro del Pueblo Formation, Mexico
Material- (IGM-7714) partial caudal centrum
(IGM-7716) distal pedal phalanx
Comments- Rodriguez de la Rosa and Cevallos-Ferriz (1998) assigned IGM-7714
to Theropoda indet. and IGM-7716 to probable Dromaeosauridae, but the somewhat
centrally placed collateral ligement pit and uncertain phalangeal identity make
assignment more precise than Averostra uncertain.
Reference- Rodriguez de la Rosa and Cevallos-Ferriz, 1998. Vertebrates
of the El Pelillal locality (Campanian, Cerro del Pueblo Formation), southeastern
Coahuila, Mexico. Journal of Vertebrate Paleontology. 18(4), 751-764.
unnamed Averostra (Rodriguez-de la Rosa and Aranda-Manteca, 1999)
Late Campanian, Late Cretaceous
El Gallo Formation, Mexico
Material- (FCM 06/053) tooth (~20x9.5x5.6 mm) (Rodr�guez-de la Rosa
and Aranda-Manteca, 1999)
(LACM 17696) tooth (Hilton, 2003)
(LACM 17697) tooth (Hilton, 2003)
(LACM 17701) tooth (Hilton, 2003)
(LACM 17704) tooth (Hilton, 2003)
(LACM 17714) teeth (Hilton, 2003)
(LACM 20879) tooth (Hilton, 2003)
(LACM 20889) phalanx, distal phalanx (Hilton, 2003)
(LACM 28993) teeth (Hilton, 2003)
(LACM 28997) teeth (Hilton, 2003)
(LACM 42563) tooth (Hilton, 2003)
(LACM 42564) tooth (Hilton, 2003)
(LACM 42565) manual ungual (Hilton, 2003)
(LACM 42571) distal manual phalanx (Hilton, 2003)
(LACM 42574) tooth (Hilton, 2003)
(LACM 42631) tooth (Hilton, 2003)
(LACM 42638) manual phalanx (Hilton, 2003)
(LACM 42669) tooth (Hilton, 2003)
(LACM 42685) tooth (Hilton, 2003)
(LACM 42687) tooth (Hilton, 2003)
(LACM 42703) ungual (Hilton, 2003)
(LACM 42704) tooth (Hilton, 2003)
(LACM 42705) tooth (Hilton, 2003)
(LACM 52458) teeth (Hilton, 2003)
(LACM 57871) tooth fragment (Hilton, 2003)
(LACM 101163) teeth (Hilton, 2003)
(LACM 101164) manual phalanx (Hilton, 2003)
(LACM 101173) tooth (Hilton, 2003)
(LACM 101182) caudal vertebra (Hilton, 2003)
(LACM 101183) tooth (Hilton, 2003)
(LACM 101184) teeth (Hilton, 2003)
(UCMP coll.) vertebrae, ribs (Hilton, 2003)
(UCMP coll.) tooth (Hilton, 2003)
(IGM coll.) tooth (Romo de Vivar, 2011)
(IGM coll.) tooth (Romo de Vivar, 2011)
(IGM coll.) tooth (Romo de Vivar, 2011)
Comments- FCM 06/053 (Rodr�guez-de la Rosa and Aranda-Manteca,
1999, 2000) is a tooth notable for its high DSDI and distal serrations being
located inside a groove. Note while one might be tempted to compare this to
Sinornithosaurus, which also has a high DSDI and supposedly venom-related
grooves, the grooves of the latter are located labially, not distally.
References- Rodr�guez-de la Rosa and Aranda-Manteca, 1999. Theropod
teeth from the Late Cretaceous El Gallo Formation, Baja California, Mexico.
VII International Symposium on Mesozoic Terrestrial Ecosystems. 56.
Rodr�guez-de la Rosa and Aranda-Manteca, 2000.Where there venomous theropods?
Journal of Vertebrate Paleontology. 20(3), 64A.
Hilton, 2003. Dinosaurs and other Mesozoic Reptiles of California. University
of California Press. 356 pp.
Romo de Vivar, 2011. Microvertebrados Cret�cicos Tard�os del �rea
de El Rosario, Baja California, M�xico. MS thesis, Universidad Nacional
Aut�noma de M�xico. 146 pp.
undescribed Averostra (Ramirez-Valesco and Hernandez-Rivera, 2015)
Late Campanian-Maastrichtian, Late Cretaceous
Cabullona Group, Mexico
Material- (MPF coll.) metatarsal fragment
(MPF coll.) phalanx fragment
Reference- Ramirez-Valesco and Hernandez-Rivera, 2015. Diversity of Late
Cretaceous dinosaurs from Mexico. Bolet�n Geol�gico y Minero.
126(1), 63-108.
undescribed averostran (Ramirez-Valesco and Hernandez-Rivera, 2015)
Late Campanian-Early Maastrichtian, Late Cretaceous
Olmos Formation, Mexico
Material- (IGM coll.) tooth fragments
Reference- Ramirez-Valesco and Hernandez-Rivera, 2015. Diversity of Late
Cretaceous dinosaurs from Mexico. Bolet�n Geol�gico y Minero.
126(1), 63-108.
unnamed possible Averostra (Lindgren, Currie, Rees, Siverson, Lindstr�m
and Alwmark, 2008)
Early Berriasian, Early Cretaceous
Skyttegaard Member of the Rabekke Formation, Denmark
Material- (MGUH 28410a-d) fragmented premaxillary tooth
(MGUH coll.) tooth fragment
Comments- Lindgren et al. (2008) referred a-d to Maniraptora incertae
sedis, noting the varying serration size is found in taxa with normally serrationless
teeth.
References- Lindgren, Currie, Rees, Siverson, Lindstr�m and Alwmark,
2008. Theropod dinosaur teeth from the lowermost Cretaceous Rabekke Formation
on Bornholm, Denmark. Geobios. 41, 253-263.
Bonde, 2012. Danish dinosaurs: A review. In Godefroit (ed.). Bernissart Dinosaurs
and Early Cretaceous Terrestrial Ecosystems. Indiana University Press. 434-451.
unnamed averostran (Poropat, Einarsson, Lindgren, Bazzi, Lagerstam
and Kear, 2016)
Early Campanian, Late Cretaceous
Ugnsmunnarna, Sweden
Material- (LO 12095t) incomplete tibia
Reference- Poropat, Einarsson, Lindgren, Bazzi, Lagerstam and Kear, 2016 (online 2015).
Late Cretaceous dinosaurian remains from the Kristianstad Basin of southern
Sweden. In Kear, Lindgren, Hurum, Mil�n and Vajda (eds.). Geological
Society, London, Special Publications. 434, 231-239.
undescribed averostran (Metcalf and Walker, 1994)
Early Bathonian, Middle Jurassic
Chipping Norton Formation, England
Material- (GLRCM coll.; C) premaxillary tooth (3.4 mm)
Comments- This was labeled as "dromaeosaur-like" by Metcalf
and Walker (1994).
It is a premaxillary tooth which is serrated on both carinae, though the mesial
serrations do not extend as basally. Serrations are very flat and low with no
blood pits. Serration density is 5-6/mm mesially and distally.
These are all fairly basal characters, suggesting the tooth may come from a
basal tetanurine or carnosaur, or perhaps a very basal coelurosaur.
Reference- Metcalf and Walker, 1994. A new Bathonian microvertebrate
locality in the English Midlands. In Fraser and Sues (eds.). In the Shadow of
the Dinosaurs- Mesozoic Small Tetrapods. Cambridge University Press.
322-332.
unnamed Averostra (Benson, 2009)
Middle Bathonian, Middle Jurassic
Taynton Limestone Formation (=Stonesfield Slate), England
Material- (MB R 2351) distal tibia (32.1 mm wide) (Galton and Molnar, 2005)
(OUM J12003) proximal tibia (Benson, 2009)
(OUM J29776) tooth (Benson, 2009)
(OUM J29778) tooth (Benson, 2009)
Comments-
Though Ezcurra and Agnolin (2012) referred MB R 2351 to Abelisauroidea,
Rauhut (2012) noted their listed characters have broader distributions
and are not consistently found in abelisauroids. He regarded it as
Averostra indet., which seems valid as some ceratosaurs (e.g. Quilmesaurus,
Majungasaurus) and most tetanurines have similar amounts of distal anteroposterior
compression.
OUM J12003 is a tibia which differs from Megalosaurus
in being smaller, with a thin fibular crest. Benson (2009) suggested it may be the
same taxon as the tetanurine distal tibia MB R 2351. OUM J29776 is a tooth which
differs from Megalosaurus in having a much higher serration density-
18 per 5 mm mesially and 18.5 per 5 mm mesially. Faint enamel wrinkles are present
and the mesial serrations extend 40% of the crown length. OUM J29778 also has
fine serrations- 14.5 per 5 mm on both carinae, and has mesial serrations over
half the crown length. Both are smaller than Megalosaurus and have interdenticular
sulci which are are short and perpendicular to the carinae.
References- 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.
Benson, 2009. An assessment of variability in theropod dinosaur
remains from the Bathonian (Middle Jurassic) of Stonesfield and New Park Quarry,
UK and taxonomic implications for Megalosaurus bucklandii and Iliosuchus
incognitus. Palaeontology. 52(4), 857-877.
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.
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.
undescribed Averostra (Lydekker, 1888)
Kimmeridgian, Late Jurassic
Kimmeridge Clay, England
Material- (NHMUK 46388) partial tooth (Lydekker, 1888)
(private coll.) two pedal phalanges (Brokenshire and Clarke, 1993)
Comments- NHMUK 46388 was referred to Megalosaurus insignis by Lydekker (1888), but is neither
described nor illustrated.
The pedal phalanges were described as ornithomimid by Brokenshire and Clarke
(1993), but Martill et al. (2006) could not place it more exactly than Theropoda
indet..
References- 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.
Delair, 1973. The Dinosaurs of Wiltshire. Whiltshire Archaeology and Natural
History Magazine. 68, 1-7.
Brokenshire and Clarke, 1993. Important recently collected
dinosaurian remains from the Lower Kimmeridge Clay at Weymouth. Proceedings
of the Dorset Natural History and Archaeological Society. 115, 177-178.
Martill, Naish and Earland, 2006. Dinosaurs in marine strata: Evidence from
the British Jurassic, including a review of the allochthonous vertebrate assemblage
from the marine Kimmeridge Clay Formation (Upper Jurassic) of Great Britain.
In Colectivo Arqueol�gico-Paleontol�gico Salense (ed.). Actas
de las III Jornadas sobre Dinosaurios y su Entorno. Salas de los Infantes (Burgos,
Espa�a). 35-72.
unnamed Averostra (Milner, 2002)
Berriasian, Early Cretaceous
Purbeck Limestone Group, England
Material- ?(NHMUK 48251 in part) nine teeth (Fowler, 2007 online)
(NHMUK 44806) tooth (64 mm) (Lydekker, 1888)
(NHMUK R 2566a) tooth (28.5 mm) (Woodward, 1895)
(NHMUK R 2566b) tooth (31.5 mm) (Woodward, 1895)
(NHMUK R 2566c) tooth (26.5 mm) (Woodward, 1895)
(NHMUK R 2567a) tooth (Milner, 2002)
(NHMUK R 2567b) tooth (Milner, 2002)
(NHMUK R 2821) tooth (56.8 mm) (Milner, 2002)
(NHMUK R 6908; = DORCM G 80) partial metatarsal III (~280 mm) (Milner, 2002)
Early Berriasian, Early Cretaceous
Marly Freshwater Member of Lulworth Formation of Purbeck Limestone Group, England
(CAMSM J 13956) pedal phalanx III-1 (24 mm) (Milner, 2002)
Comments- Lydekker (1888) assigned NHMUK 44806 to Megalosaurus dunkeri,
while it was later assigned to Altispinax dunkeri (Huene, 1926) and Megalosaurus
sp. (Delair, 1959). NHMUK R 2566 was assigned to Megalosaurus sp.
by Woodward (1895). Milner (2002) felt these and three others (NHMUK R 2567 and
2821) were closer to allosaurids than megalosaurids based on their higher DSDI.
However, Dubreuillosaurus has a comparably high DSDI, so this is not
a valid character for distinguishing carnosaurs from basal tetanurines. The
teeth are retained in Averostra indet. until they are described in more detail.
Neither these nor the pedal phalanx are illustrated. Fowler (2007 online) searched
the crocodilian NHMUK remains and found nine teeth that are probably theropod
instead.
The distal metatarsal III was found before 1954 but only described in 2002 by
Milner. She tentatively assigned it to the Eumaniraptora based on slenderness,
non-arctometatarsal condition and spatiotemporal occurance (earlier than known
oviraptorosaurs, which are incorrectly said to be only known from Mongolia and
Canada). However, most eumaniraptorans have ginglymoid third metatarsi (Bambiraptor,
Velociraptor, Deinonychus, Dromaeosaurus, Achillobator,
Rahonavis, Shenzhouraptor, Hulsanpes, basal Avebrevicauda),
a subarctometatarsus (Pedopenna, Sinornithosaurus, Graciliraptor,
Archaeopteryx), or both (Neuquenraptor, Microraptor, Troodontidae,
Buitreraptor). Possible exceptions are scansoriopterygids, Jixiangornis
and Yandangornis, though the former are only known from juvenile material
and the latter two are illustrated and described poorly. So contra Milner, I
find this metatarsal to most likely not be eumaniraptoran. Many non-paravian
theropods can be excluded due to arctometatarsaly or subarctometatarsaly (tyrannosaurids,
ornithomimosaurs, mononykines, basal oviraptorosaurs) or ginglymoidy (Compsognathus,
Dilong, allosaurids, Acrocanthosaurus, Sinraptor, Torvosaurus).
However, there are still several equally plausible alternatives left for the
Purbeck metatarsal, which resembles not only oviraptorids, but also such varied
taxa as Fukuiraptor, Falcarius, Tanycolagreus, Ornitholestes,
Coelurus, Nqwebasaurus, Elaphrosaurus and Masiakasaurus.
I recommend classifying it as Averostra indet..
References- 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.
Woodward, 1895. Note on megalosaurian teeth discovered by Mr. J. Alstone in
the Portlandian of Aylesbury. Proceedings of the Geologists' Association. 14,
31-32.
Huene, 1926. The carnivorous Saurischia in the Jura and Cretaceous Formations,
principally in Europe. Revista del Museo de La Plata. 29, 1-167.
Delair, 1959. The Mesozoic reptiles of Dorset. Part 2. Proceedings of the Dorset
Natural History and Archaeological Society. 80, 52-90.
Milner, 2002. Theropod dinosaurs of the Purbeck Limestone Group, southern England.
In Milner and Batten (eds.). Life and Environments in Purbeck Times. Special
Papers in Palaeontology. 68, 191-201.
Fowler, 2007 online. http://www.denverfowler.com/publications/Fowler_2007_SVP.htm
unnamed averostran (Seeley, 1899)
Valanginian, Early Cretaceous
Hastings Group, England
Material- (lost) distal femur
Comments- Seeley (1899) described this as a bird most similar to grebes,
though Galton and Martin (2002) thought it wasn't a bird. Naish (2000) thought
it was very similar to Wessex Formation femur MIWG 6214 except for possessing
an extensor/patellar groove. MIWG 6214 has most recently been referred to Ornithomimosauria
by Allain et al. (2014).
References- Seeley, 1899. On evidence of a bird from the Wealden beds
of Ansty Lane, near Cuckfield. Quarterly Journal of the Geological Society of
London. 55, 416-418.
Naish, 2000. A small, unusual theropod (Dinosauria) femur from the Wealden Group
(Lower Cretaceous) of the Isle of Wight, England. Neues Jahrbuch f�r Geologie
und Pal�ontologie Monatshefte. 2000, 217-234.
Galton and Martin, 2002. Postcranial anatomy and systematics of Enaliornis
Seeley, 1876, a footpropelled diving bird (Aves: Ornithurae: Hesperornithiformes)
from the Early Cretaceous of England. Revue de Paleobiologie. 21(2), 489-538.
Allain, Vullo, Le Loeuff and Tournepiche, 2014. European ornithomimosaurs (Dinosauria,
Theropoda): An undetected record. Geologica Acta. 12(2), 127-135.
undescribed Averostra (Lydekker, 1888)
Hauterivian-Barremian, Early Cretaceous
Weald Clay, England
Material- (NHMUK 28958) partial posterior dorsal centrum (Lydekker, 1888)
(NHMUK 37691) dorsal centrum (Lydekker, 1888)
(NHMUK 44806) tooth (Lydekker, 1888)
(NHMUK R139) two sacral fragments (Lydekker, 1888)
(NHMUK R141) dorsal centrum (Lydekker, 1888)
(NHMUK R210) tooth (Lydekker, 1888)
(NHMUK R1997) tooth (Osi et al., 2010)
(NHMUK R15909) tooth (Osi et al., 2010)
Comments- Most of this material was referred to Megalosaurus dunkeri
by Lydekker (1888), then Altispinax dunkeri by Huene (1926), but has
not been described in detail so cannot be compared to the holotypes of either.
They may belong to Baryonyx, Altispinax, Valdoraptor, Neovenator, Calamosaurus,
Eotyrannus or another Wealden theropod. Osi et al. (2010) described two
additional supposed Megalosaurus dunkeri teeth, finding them and NHMUK
R210 to clade with "Megalosaurus" pannoniensis in a morphometric
study, but these are all from the Weald Clay of England, so may not belong to
dunkeri in the first place.
References- 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.
Huene, 1926. The carnivorous Saurischia in the Jura and Cretaceous formations,
principally in Europe. Revista del Museo de La Plata. 29, 1-167.
Osi, Apesteguia and Kowalewski, 2010. Non-avian theropod dinosaurs from the
early Late Cretaceous of central Europe. Cretaceous Research. 31(3), 304-320.
undescribed Averostra (Mantell, 1827)
Barremian, Early Cretaceous
Upper Weald Clay, England
Material- (NHMUK 2141) posterior dorsal neural arch (Mantell, 1827)
(NHMUK 2294) partial caudal vertebra (Lydekker, 1888)
....(NHMUK 2295) partial caudal vertebra (Lydekker, 1888)
(NHMUK 2315) tooth (Lydekker, 1888)
(NHMUK 2332) tooth (Lydekker, 1888)
(NHMUK 2482) pedal ungual (Mantell, 1827)
(NHMUK 2501) pedal phalanx ?-1 (Lydekker, 1888)
(NHMUK 2503) pedal phalanx III-2 (Mantell, 1827)
(NHMUK 2513) tooth (Lydekker, 1888)
(NHMUK 2553; cast) metacarpal (Lydekker, 1888)
(NHMUK 2574) distal metatarsal III (Lydekker, 1888)
(NHMUK 2680) metatarsal III (Lydekker, 1888)
(NHMUK 2828) tooth (Lydekker, 1888)
(NHMUK 3221) tooth (Lydekker, 1888)
(NHMUK 3222) tooth (Lydekker, 1888)
(NHMUK 3223-3224) four teeth (Lydekker, 1888)
(NHMUK 3225) tooth (Lydekker, 1888)
(NHMUK 3333) tooth (Lydekker, 1888)
(NHMUK 3640) proximal pedal phalanx ?-1 (Lydekker, 1888)
(NHMUK 26012) tooth (Lydekker, 1888)
(NHMUK 28422) tooth (Lydekker, 1888)
(NHMUK 36495) partial metacarpal (Lydekker, 1888)
(NHMUK 36496) partial metacarpal (Lydekker, 1888)
(NHMUK 36551) proximal pedal phalanx ?-1 (Mantell, 1827)
(NHMUK 36522) partial tooth (Lydekker, 1888)
(NHMUK 36522a) three teeth (Lydekker, 1888)
(NHMUK 36523) tooth (Lydekker, 1888)
(NHMUK 39197) tooth (Lydekker, 1888)
(NHMUK R235) tooth (Lydekker, 1888)
(NHMUK R1105) incomplete manual ungual ?III (Owen, 1857)
Comments- Most of this material was referred to Megalosaurus dunkeri
by Lydekker (1888), then Altispinax dunkeri by Huene (1926), but has
not been described in detail so cannot be compared to the holotypes of either.
They may belong to Baryonyx, Altispinax, Valdoraptor, Neovenator, Calamosaurus,
Eotyrannus or another Wealden theropod.NHMUK 2141 and NHMUK 2482 were initially
illustrated as Iguanodon by Mantell (1827). NHMUK R1105 and NHMUK 2482
were described and illustrated by Owen (1857) as Megalosaurus bucklandii
(plate X, fig. 1-4 and 5 respectively). NHMUK R2559 was referred to M. dunkeri
by Lydekker (1888), but was made the holotype of Megalosaurus oweni by
the author the next year. NHMUK 2574 and 2680 were referred to oweni by
Lydekker (1890), though this has not been confirmed by recent studies. Note
Ford (www.paleofile.com) mistyped NHMUK 39197 as NHMUK 39497, and NHMUK 2513 as NHMUK 35523.
References- Mantell, 1827. Illustrations of the Geology of Sussex: The
Fossils of Tilgate Forest. Lupton Relfe, [pp].
Owen, 1857. Monograph on the Fossil Reptilia of the Wealden and Purbeck Formations.
Part III. Dinosauria (Megalosaurus) (Wealden). Palaeontographical Society
Monographs. 9, 1-26.
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, 1926. The carnivorous Saurischia in the Jura and Cretaceous formations,
principally in Europe. Revista del Museo de La Plata. 29, 1-167.
undescribed Averostra (Hutt, 2001)
Barremian, Early Cretaceous
Wessex Formation, England
Material- (IWCMS 1995.234) tooth
(IWCMS 1995.235) tooth
(IWCMS 1996.154) sacrum, fragments
(IWCMS 1997.152) premaxilla
(IWCMS 2000.1108) metatarsal
(MIWG 46) tooth
(MIWG 1473) humeral shaft
?(MIWG 1498) fragment
?(MIWG 1587) fragment
?(MIWG 3006) fragment
?(MIWG 3240) fragment
(MIWG 3644) fragment
(MIWG 5120) tooth
(MIWG 5121) tooth
(MIWG 5126) tooth
(MIWG 5129) teeth
(MIWG 5134) teeth
(MIWG 5181) jaw fragment
(MIWG 5182) jaw fragment
(MIWG 5185) tooth fragment
(MIWG 5229) rib
(MIWG 5139) tooth
(MIWG 5245) fragment
(MIWG 5251) rib
(MIWG 5309) rib
(MWIG 5358) teeth
(MWIG 5424) caudal vertebra
(MWIG 5439) tooth
(MIWG 5456) sacrum
(MIWG 5820) cervical vertebra
(MIWG 5823) dorsal vertebra
(MIWG 5924) centrum
(MIWG 5832) pectoral or pelvic elements
(MIWG 5962) teeth
(MIWG 6350) pubis, femur
(MIWG 6351) tooth
(MIWG 6515) cervical vertebra
(MIWG 6516) cervical vertebra
(MIWG 6479) caudal vertebra
(MIWG 6869) tooth
(MIWG 6913) ?dorsal vertebra
(MIWG 7049) tibia
References-
Hutt, 2001. Catalogue of Wealden Group Dinosauria in the Museum
of Isle of Wight Geology. In Martill and Naish (eds.). Dinosaurs of the Isle
of Wight. The Palaeontological Association. 411-422.
undescribed Averostra (Lydekker, 1888)
Aptian, Early Cretaceous
Lower Greensand, England
Material- (NHMUK 40455) partial distal caudal vertebra
(NHMUK 42032) dorsal centrum
Comments- This material was referred to Megalosaurus dunkeri by
Lydekker (1888), then Altispinax dunkeri by Huene (1926), but has not
been described and is probably too late to be either species.
References- 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.
Huene, 1926. The carnivorous Saurischia in the Jura and Cretaceous formations,
principally in Europe. Revista del Museo de La Plata. 29, 1-167.
unnamed possible theropod (Seeley, 1876)
Late Albian, Early Cretaceous
Cambridge Greensand, England
Material- (SMC B55281) anterior sacrum
Comments- SMC B55281 was identified by Seeley (1876) as a partial sacrum
of Enaliornis barretti, reidentified as a pterosaur notarium by Galton
and Martin (2002a), then identified again as a possible theropod sacrum by Galton
and Martin (2002b).
References- Seeley, 1876. On the British fossil Cretaceous birds. Quarterly
Journal of the Geological Society of London. 32, 496-515.
Galton and Martin, 2002a. Enaliornis, an Early Cretaceous hesperornithiform
bird from England, with comments on other Hesperornithiformes. In Chiappe and
Witmer (eds). Mesozoic birds: Above the heads of dinosaurs. Berkeley: University
of California Press. 317-338.
Galton and Martin, 2002b. Postcranial anatomy and systematics of Enaliornis
Seeley, 1876, a foot-propelled diving bird (Aves: Ornithurae: Hesperornithiformes)
from the Early Cretaceous of England. Revue de Paleobiologie. 21(2), 489-538.
undescribed Averostra (Lapparent and Zbyszewski, 1957)
Callovian-Oxfordian, Middle-Late Jurassic
Colmeias, Leiria, Portugal
Material- (Geological Services Museum of Portugal coll.?) tooth
Comments- This was referred to Megalosaurus insignis, but is neither
described nor illustrated.
Reference- Lapparent and Zbyszewski, 1957. Les dinosauriens du Portugal
[The dinosaurs of Portugal]. M�moires des Services G�ologiques
du Portugal, nouvelle s�rie. 2, 1-63.
unnamed possible Averostra (Lapparent and Zbyszewski, 1957)
Callovian-Oxfordian, Middle-Late Jurassic
Ferven�a, Leiria, Portugal
Material- (Geological Services Museum of Portugal coll.?) distal caudal
vertebra (54 mm), distal caudal vertebra (51 mm)
Comments- These were referred to Megalosaurus insignis, but cannot
be compared to the holotype. They were not reported to be associated and may
not be theropod.
Reference- Lapparent and Zbyszewski, 1957. Les dinosauriens du Portugal
[The dinosaurs of Portugal]. M�moires des Services G�ologiques
du Portugal, nouvelle s�rie. 2, 1-63.
undescribed Averostra (Lapparent and Zbyszewski, 1957)
Callovian-Oxfordian, Middle-Late Jurassic
Pombal, Leiria, Portugal
Material- (Geological Services Museum of Portugal coll.?) four teeth
Comments- These were referred to Megalosaurus insignis, but are
neither described nor illustrated. They were not reported to be associated.
Reference- Lapparent and Zbyszewski, 1957. Les dinosauriens du Portugal
[The dinosaurs of Portugal]. M�moires des Services G�ologiques
du Portugal, nouvelle s�rie. 2, 1-63.
unnamed possible Averostra (Lapparent and Zbyszewski, 1957)
Callovian-Oxfordian, Middle-Late Jurassic
Salir do Porto, Leiria, Portugal
Material- (Geological Services Museum of Portugal coll.?) three caudal vertebrae
Comments- These were referred to Megalosaurus insignis, but cannot
be compared to the holotype. They were not reported to be associated and may
not be theropod.
Reference- Lapparent and Zbyszewski, 1957. Les dinosauriens du Portugal
[The dinosaurs of Portugal]. M�moires des Services G�ologiques
du Portugal, nouvelle s�rie. 2, 1-63.
undescribed averostran (Sauvage, 1898)
Oxfordian, Late Jurassic
Portugal
Material- tooth
Comments- This was referred to Megalosaurus insignis, but is neither
described nor illustrated.
Reference- Sauvage, 1898. Les Reptiles et les Poissons des terrains M�sozo�ques
du Portugal [The reptiles and fishes from the Mesozoic terrains of Portugal].
Bulletin de la Soci�t� G�ologique de France, 3e s�rie.
26, 442-446.
undescribed averostran (Rauhut, 2000)
Kimmeridgian, Late Jurassic
Guimarota, Alcoba�a Formation, Leiria, Portugal
Material- (IPFUB Gui Th 5) manual ungual
Reference- Rauhut, 2000. The dinosaur fauna from the Guimarota mine.
In Martin and Krebs (eds.). Guimarota - A Jurassic Ecosystem. Verlag Dr. Friedrich
Pfeil. 75-82.
undescribed Averostra (Sauvage, 1894)
Kimmeridgian, Late Jurassic
Alcoba�a Formation, Crasto and Pembel or Pombal, Portugal
Comments- This was referred to Megalosaurus insignis, but is neither
described nor illustrated.
References- Sauvage, 1894. Les dinosauriens du terrain jurassique sup�rieur
du Boulonnais. Bulletin de la Soci�t� G�ologique de France,
3e serie. 22, 465-470.
Sauvage, 1898. Vert�br�s fossiles du Portugal: Contributions �
l'�tude des poissons et des reptiles du jurassique et du cr�tacique.
Direction des Travaux Geologiques du Portugal. Memoires. Comiss�o do
Servi�o Geol�gico de Portugal. 1-46.
unnamed possible averostran (Lapparent and Zbyszewski, 1957)
Kimmeridgian, Late Jurassic
Praia Areia Branca, Leiria, Portugal
Material- (Geological Services Museum of Portugal coll.?) posterior dorsal
vertebra (50 mm)
Comments- This was referred to Megalosaurus insignis, but cannot
be compared to the holotype. The triangular centrum suggests it may not be theropod.
References- Lapparent and Zbyszewski, 1957. Les dinosauriens du Portugal
[The dinosaurs of Portugal]. M�moires des Services G�ologiques
du Portugal, nouvelle s�rie. 2, 1-63.
unnamed possible averostran (Lapparent and Zbyszewski, 1957)
Kimmeridgian, Late Jurassic
Cesareda, Leiria, Portugal
Material- (Geological Services Museum of Portugal coll.?) mid caudal
centrum (76 mm)
Comments- This was referred to Megalosaurus insignis, but cannot
be compared to the holotype and may not be theropod.
Reference- Lapparent and Zbyszewski, 1957. Les dinosauriens du Portugal
[The dinosaurs of Portugal]. M�moires des Services G�ologiques
du Portugal, nouvelle s�rie. 2, 1-63.
unnamed Averostra (Lapparent and Zbyszewski, 1957)
Kimmeridgian, Late Jurassic
Porto de Barcas, Lisboa, Portugal
Material- (Geological Services Museum of Portugal coll.?) tooth
(Geology Laboratory of the Faculty of Sciences of Lisbon coll.) distal femur
Comments- These were referred to Megalosaurus insignis. The tooth
is neither described nor illustrated, and the femoral fragment cannot be compared
to the holotype and may not be theropod. The remains were not reported to be
associated.
Reference- Lapparent and Zbyszewski, 1957. Les dinosauriens du Portugal
[The dinosaurs of Portugal]. M�moires des Services G�ologiques
du Portugal, nouvelle s�rie. 2, 1-63.
unnamed Averostra (Lapparent and Zbyszewski, 1957)
Kimmeridgian, Late Jurassic
Ourem, Santarem, Portugal
Material- (Geological Services Museum of Portugal coll.?) tooth, sacral
centrum (90 mm), sacral centrum (80 mm), two partial distal caudal vertebrae,
ulnar fragment
Comments- These were referred to Megalosaurus insignis. The tooth
is neither described nor illustrated, and the postcrania cannot be compared
to the holotype and may not be theropod. The remains were not reported to be
associated, and indeed the sacrals are different sizes.
Reference- Lapparent and Zbyszewski, 1957. Les dinosauriens du Portugal
[The dinosaurs of Portugal]. M�moires des Services G�ologiques
du Portugal, nouvelle s�rie. 2, 1-63.
undescribed Averostra (Lapparent and Zbyszewski, 1957)
Late Kimmeridgian, Late Jurassic
Foz do Arelho, Leiria, Portugal
Material- (Geological Services Museum of Portugal coll.?) six teeth
Comments- These were referred to Megalosaurus insignis, but are
neither described nor illustrated. They were not reported to be associated.
Reference- Lapparent and Zbyszewski, 1957. Les dinosauriens du Portugal
[The dinosaurs of Portugal]. M�moires des Services G�ologiques
du Portugal, nouvelle s�rie. 2, 1-63.
unnamed Averostra (Lapparent and Zbyszewski, 1957)
Late Kimmeridgian-Early Tithonian, Late Jurassic
Lourinha Formation, Portugal
Material- (Geological Services Museum of Portugal coll.?) five teeth
(Geology Laboratory of the Faculty of Sciences of Lisbon coll.) two manual unguals
(15, ~17 mm)
Comments- These were referred to Megalosaurus insignis, though
the teeth are neither described nor illustrated. The unguals cannot be compared
to the holotype. None of the material was reported to be associated.
Reference- Lapparent and Zbyszewski, 1957. Les dinosauriens du Portugal
[The dinosaurs of Portugal]. M�moires des Services G�ologiques
du Portugal, nouvelle s�rie. 2, 1-63.
unnamed Averostra (Lapparent and Zbyszewski, 1957)
Late Kimmeridgian-Early Tithonian, Late Jurassic
Atalaia, Sobral Formation, Lisbon, Portugal
Material- (Geological Services Museum of Portugal coll.?) nine teeth (60,
38, 25 mm)
Comments- These were referred to Megalosaurus insignis and were
not reported to be associated.
Reference- Lapparent and Zbyszewski, 1957. Les dinosauriens du Portugal
[The dinosaurs of Portugal]. M�moires des Services G�ologiques
du Portugal, nouvelle s�rie. 2, 1-63.
undescribed averostran (Malafaia, Mocho, Escaso, Dantas and Ortega, 2019)
Late Tithonian, Late Jurassic
Praia de Cambelas, Freixial Formation, Portugal
Material- (SHN.019/1) (small) tooth
Comments- Malafaia et al. (2019) listed this as "a tooth of an indeterminate small theropod" associated with the Lusovenator paratype.
Reference-
Malafaia, Mocho, Escaso, Dantas and Ortega, 2019 (online 2018).
Carcharodontosaurian remains (Dinosauria, Theropoda) from the Upper
Jurassic of Portugal. Journal of Paleontology. 93, 157-172.
unnamed Averostra (Sauvage, 1898)
Aptian, Early Cretaceous
Boca do Chapim, Portugal
Material- teeth (Sauvage, 1898)
(Geological Services Museum of Portugal coll.?) two tooth fragments (FABL 20
mm)
Comments- Sauvage (1898) referred teeth to Megalosaurus superbus.
Lapparent and Zbyszewski (1957) referred two tooth fragments from the Aptian
of Portugal to the same species. but they could belong to any ceratosaur, basal
tetanurine, carnosaur or basal coelurosaur.
References- Sauvage, 1898. Vert�br�s fossiles du Portugal:
Contributions � l'�tude des poissons et des reptiles du jurassique
et du cr�tacique. Direction des Travaux Geologiques du Portugal. Memoires.
Comiss�o do Servi�o Geol�gico de Portugal. 1-46.
Lapparent and Zbyszewski, 1957. Les dinosauriens du Portugal [The dinosaurs
of Portugal]. M�moires des Services G�ologiques du Portugal, nouvelle
s�rie. 2, 1-63.
unnamed Averostra (Sauvage, 1898)
Late Campanian-Maastrichtian, Late Cretaceous
Viso, Coimbra, Portugal
Referred- ?(Geological Services Museum of Portugal coll.?) partial tooth
(~30x12x8 mm), tooth (26 mm), tooth fragment (Sauvage, 1898)
?(Geological Services Museum of Portugal coll.?) incomplete manual ungual (~22
mm), incomplete manual ungual (~17 mm), incomplete manual ungual (~13 mm) (Lapparent
and Zbyszewski, 1957)
Comments- The teeth were referred to Megalosaurus sp. by Sauvage
(1898), then they and the unguals were referred to Megalosaurus pannoniensis
by Lapparent and Zbyszewski (1957). The unguals are not comparable to the holotype,
though the measured tooth matches it in labiolingual compression and mesiodistal
elongation. Whether any specimens were associated is unknown.
References- Sauvage, 1898. Vert�br�s fossiles du Portugal:
Contributions � l'�tude des poissons et des reptiles du jurassique
et du cr�tacique. Direction des Travaux Geologiques du Portugal. Memoires.
Comiss�o do Servi�o Geol�gico de Portugal. 1-46.
Lapparent and Zbyszewski, 1957. Les dinosauriens du Portugal [The dinosaurs
of Portugal]. M�moires des Services G�ologiques du Portugal, nouvelle
s�rie. 2, 1-63.
unnamed Aversotra (Casanovas-Cladellas et al., 1993)
Tithonian-Berriasian, Late Jurassic-Early Cretaceous
Villar del Arzobispo Formation, Spain
Material- (BZ-3) partial tooth (Casanovas-Cladellas et al., 1993)
(FS-1) incomplete tooth (?x37.1x20.6 mm) (Su�er, Santisteban and Galobart,
2005)
(GAOO/CL/14) tooth (Barco and Ruiz-Ome�aca 2001a)
(LP-1) partial tooth (?x19.4x15 mm) (Su�er, Santisteban and Galobart,
2005)
(LV-1) tooth (67.5x24.7x11.1 mm) (Su�er, Santisteban and Galobart, 2005)
(MPZ01/97) proximal caudal centrum (Barco and Ruiz-Ome�aca 2001b)
(PM-1) partial pedal ungual (Su�er, Santisteban and Galobart, 2005)
tooth (Abella and Su�er, 2004)
three teeth, one bone (Su�er and Mart�n, 2009)
fifteen partial teeth (Gasco et al., 2012)
Comments- BZ-3 was assigned to Carnosauria by Casanovas-Cladellas et
al. (1993), then to Theropoda indet. by Ruiz-Ome�aca and Canudo (2003).
References- Casanovas-Cladellas, Santaf�-Llopis and Santisteban-Bov�,
1993. First dinosaur teeth from the Lower Cretaceous of Benicatazara (Aras de
Alpuente, Valencia). Revue de Pal�obiologie. Special volume 7, 37-44.
Barco and Ruiz-Ome�aca, 2001a. Primeros dientes de ter�podos (Dinosauria,
Saurischia) en la Formaci�n Villar del Arzobispo (Tit�nico-Berriasiense):
Yacimientos Cuesta Lonsal y Las Cerradicas 2 (Galve, Teruel). Publicaciones
del Seminario de Paleontolog�a de Zaragoza. 5, 239-246.
Barco and Ruiz-Ome�aca, 2001b. Primeros restos postcraneales de ter�podos
(Dinosauria, Saurischia) en la Formaci�n Villar del Arzobispo (Tit�nico-Berriasiense):
Un centro vertebral caudal del yacimiento Carretera (Galve, Teruel). Publicaciones
del Seminario de Paleontolog�a de Zaragoza. 5, 247-254.
Ruiz-Ome�aca and Canudo, 2003. Dinosaurios (Saurischia, Ornithischia)
en el Barremiense (Cret�cico inferior) de la Pen�nsula Ib�rica.
In P�rez-Lorente (Ed.). Dinosaurios y otros reptiles Mesozoicos en Espa�a.
Ciencias de la Tierra. 26, 269-312.
Canudo and Ruiz-Omenaca, 2003. Los restos directos de dinosaurios teropodos
(excluyendo Aves) en Espana. In P�rez-Lorente (Ed.). Dinosaurios y otros
reptiles Mesozoicos en Espa�a. Ciencias de la Tierra. 26, 347-373.
Abella and Su�er, 2004. Un nuevo diente aislado de ter�podo del
yacimiento de El Chopo (Alpuente, Los Serranos, Valencia). Libro de Res�menes
del II Encuentro de J�venes Investigadores en Paleontolog�a. 87-88.
Su�er, Santisteban and Galobart, 2005. Nuevos restos de Theropoda del
Jur�sico Superior-Cret�cico Inferior de la Comarca de Los Serranos
(Valencia). Revista Espa�ola de Paleontologia. N. Extra X, 93-99.
Su�er and Mart�n, 2009. Un nuevo yacimiento del tr�nsito
Jur�sico-Cret�cico de Alpuente (Los Serranos, Valencia, Espa�a):
Resultados preliminares. Paleolusitana. 1, 441-447.
Gasc�, Cobos, Royo-Torres, Mampel and Alcal�, 2012. Theropod teeth
diversity from the Villar del Arzobispo Formation (Tithonian-Berriasian) at
Riodeva (Teruel, Spain). Palaeobiodiversity and Palaeoenvironments. 92(2), 273-285.
undescribed Aversotra (Santos-Cubedo, Poza, Suner and de Santisteban,
2010)
Aptian, Early Cretaceous
Arcillas de Morella Formation, Spain
Material- teeth (Santos-Cubedo, Poza, Suner and de Santisteban, 2010)
proximal chevron (268 mm) (Selles, Santos-Cubedo and Poza, 2011)
References- Santos-Cubedo, Poza, Suner and de Santisteban, 2010. New
remains of a titanosaur (Dinosauria: Sauropoda) from the Early Cretaceous of
Spain. Journal of Vertebrate Paleontology. Program and Abstracts 2010, 157A.
Selles, Santos-Cubedo and Poza, 2011. Injury in a theropod dinosaur from the
Early Cretaceous of Spain. Journal of Vertebrate Paleontology. Program and Abstracts
2011, 192.
unnamed possible averostran (Blanco, Mendez and Marmi, 2015)
Late Maastrichtian, Late Cretaceous
Cingles de Cal Ros, Tremp Formation, Spain
Material- (IPS-81878) incomplete limb element
Reference- Blanco, Mendez and Marmi, 2015. The fossil record of the uppermost Maastrichtian
Reptile Sandstone (Tremp Formation, northeastern Iberian Peninsula). Spanish
Journal of Palaeontology. 30(1), 147-160.
undescribed averostran (Ortega, Escaso, Perez Garcia, Torices and Sanz,
2009)
Late Campanian-Early Maastrichtian, Late Cretaceous
Villalba de la Sierra Formation, Spain
Material- cranial and postcranial elements
Comments- Ortega et al. (2009) refer to this as a 'basal large theropod',
which given the provenence is most likely an abelisaurid.
Reference- Ortega, Escaso, Perez Garcia, Torices and Sanz, 2009. The
vertebrate diversity of the Upper Campanian-Lower Maastrichtian "Lo Hueco"
fossil-site (Cuenca, Spain). Journal of Vertebrate Paleontology. 29(3), 159A-160A.
unnamed Aversotra (Lapparent, 1943)
Late Oxfordian, Late Jurassic
Solvay Company quarry, Damparis, Jura, France
Material- (MNHN coll.) seven teeth (110, 22, 11, 10 mm)
Comments- These were referred to Megalosaurus insignis.
Reference- Lapparent, 1943. Les dinosauriens jurassiques de Damparis
(Jura) [The Jurassic dinosaurs of Damparis (Jura)]. M�moires de la Soci�t�
G�ologique de France (Nouvelle S�rie). M�moire 21(47),
1-21.
undescribed averostran (Sauvage, 1894)
Middle Kimmeridgian, Late Jurassic
Moulin-Wibert, Boulogne, France
Material- (Beaugrand coll.) tooth
Comments- This was referred to Megalosaurus insignis, but neither
described nor illustrated.
Reference- Sauvage, 1894. Les dinosauriens du terrain jurassique sup�rieur
du Boulonnais. Bulletin de la Soci�t� G�ologique de France,
3e serie. 22, 465-470.
unnamed Averostra (Sauvage, 1874)
Late Kimmeridgian, Late Jurassic
Ch�tillon, Pas-de-Calais, France
Material- (Mus�e de Boulogne coll.) two fused sacral centra
(Mus�e de Boulogne coll.) pedal ungual
(Mus�e de Boulogne coll.) pedal phalanx IV-?
Comments- The sacrals and ungual were referred to Megalosaurus insignis
by Sauvage (1874). He referred a pedal phalanx to ?Iguanodon, but later
(1894) reassigned it to M. insignis.
Reference- Sauvage, 1874. M�moire sur les dinosauriens et les
crocodiliens des terrains jurassiques de Boulogne-sur-Mer. M�moires
de la Soci�t� G�ologique de France, s�rie 2. 10(2),
1-57.
Sauvage, 1894. Les dinosauriens du terrain jurassique sup�rieur du Boulonnais.
Bulletin de la Soci�t� G�ologique de France, 3e serie.
22, 465-470.
undescribed Averostra (Sauvage, 1894)
Late Kimmeridgian, Late Jurassic
Wimille, Bouogne, France
Material- teeth
Comments- These were referred to Megalosaurus insignis, but neither
described nor illustrated. Note while Huene (1926) lists Megalosaurus
teeth from Wimille near Montagne Rouge, Sauvage (1894) does not list any from
there, only Cumnoria teeth and eroded fragments.
References- Sauvage, 1894. Les dinosauriens du terrain jurassique sup�rieur
du Boulonnais. Bulletin de la Soci�t� G�ologique de France,
3e serie. 22, 465-470.
Sauvage, 1900. Catalogue des Reptiles trouves dans le terrain jurassique-superieur
du Boulonnais. Compte rendu de l’Association francaise pour l’avancement
des sciences. 1899, 416-419.
Huene, 1926a. The carnivorous Saurischia in the Jura and Cretaceous formations,
principally in Europe. Revista Museo de La Plata. 29, 35-167.
undescribed Averostra (Sauvage, 1894)
Middle Tithonian, Late Jurassic
Mont-Lambert, Boulogne, France
Material- teeth
Comments- These were referred to Megalosaurus insignis.
References- Sauvage, 1894. Les dinosauriens du terrain jurassique sup�rieur
du Boulonnais. Bulletin de la Soci�t� G�ologique de France,
3e serie. 22, 465-470.
Sauvage, 1914. Catalogue des reptiles jurassiques du Boulonnais. Bulletin de
la Soci�t� acad�mique de Boulogne-sur-Mer. 10, 1-12.
Huene, 1926a. The carnivorous Saurischia in the Jura and Cretaceous formations,
principally in Europe. Revista Museo de La Plata. 29, 35-167.
unnamed Averostra (Sauvage, 1874)
Middle Tithonian, Late Jurassic
Fort de la Cr�che, Pas-de-Calais, France
Material- (Mus�e de Boulogne coll.) tooth (110 mm)
(Mus�e de Boulogne coll.) incomplete tooth (~70 mm)
(Mus�e de Boulogne coll.) incomplete tooth
(Mus�e de Boulogne coll.) tooth (>17 mm)
Comments- These were referred to Megalosaurus insignis.
Reference- Sauvage, 1874. M�moire sur les dinosauriens et les
crocodiliens des terrains jurassiques de Boulogne-sur-Mer. M�moires
de la Soci�t� G�ologique de France, s�rie 2. 10(2),
1-57.
undescribed averostran (Lydekker, 1888)
Late Tithonian, Late Jurassic
Ningle, Pas-de-Calais, France
Material- (NHMUK 35553a) tooth
Comments- This was referred to Megalosaurus insignis, but is neither
described nor illustrated.
Reference- 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.
unnamed Averostra (Sauvage, 1882b)
Early Cretaceous
Bar-le-Duc, Meuse, France
Material- (Pierson coll.) pedal phalanx II-1 (100 mm)
? (Pierson coll.) pedal phalanx (65 mm)
Comments- Sauvage (1882b) described two pedal phalanges as belonging
to Erectopus superbus, specifying one came from Bar-le-Duc. Huene (1926a,
b) retained that referral and listed both as deriving from that area. Chure
(2000) states Huene (1926a) described this element as part of the type material,
but the pedal phalanges Huene mentions in the type description are in figures
3.3 and 4.4 of Sauvage's paper (metacarpal I and pedal phalanx IV-?, both from
the type), whereas this phalanx is in figure 4.3. The phalanges may belong to
any large theropod, and are not comparable to Erectopus' type.
References- Sauvage, 1882b. Recherches sur les reptiles trouv�s
dans le Gault de l'est du bassin de Paris [Research on the reptiles found in
the Gault of the eastern Paris Basin]. M�moires de la Soci�t�
G�ologique de France, s�rie 3. 2(4), 1-42.
Huene, 1926a. The carnivorous Saurischia in the Jura and Cretaceous formations,
principally in Europe. Revista Museo de La Plata. 29, 35-167.
Huene. 1926b. On several known and unknown reptiles of the order Saurischia
from England and France. Annals and Magazine of Natural History.17, 473-489.
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.
unnamed averostran (Parent, 1893)
Early Cretaceous
Wealden Group(?), Wimereux, Pas-de-Calais, France
Material- (Lille Natural History Museum coll.) pedal ungual (~100 mm),
elements
Comments- Parent (1893) and Sauvage (1894) both referred this ungual
to Megalosaurus insignis.
References- Parent, 1893. Le Wealdien du Bas-Boulonnais. Annales de la
Societe Geologique du Nord. 21, 50-91.
Sauvage, 1894. Les dinosauriens du terrain jurassique sup�rieur du Boulonnais.
Bulletin de la Soci�t� G�ologique de France, 3e serie.
22, 465-470.
Buffetaut, Cuny and le Loeuff, 1991. French Dinosaurs: The best record in Europe?
Modern Geology. 16(1-2), 17-42.
unnamed Averostra (Barrois, 1875)
Early Albian, Early Cretaceous
Phosphate bearing beds of La Penthieve near Louppy-de-Chateau, Meuse, France
Material- (Mus�e de la Facult� des Sciences de Lille coll.)
tooth (Barrois, 1875)
(Pierson coll.) distal femur (Sauvage, 1882a)
Comments- Barrois (1875) and Sauvage (1876) described a tooth from Louppy-de-Chateau
as Megalosaurus. Sauvage (1882a) briefly described a partial skeleton
from there as a new species Megalosaurus superbus, referring a large
distal femur found in the same area to the same taxon. He later (1882b) described
both in more detail and indicated the tooth was also thought to be from that
species. Huene (1926a, b) believed the distal femur to come from a different
taxon of carnosaur from M. superbus, which he renamed Erectopus
in 1923. Huene (1932) later kept the tooth referred to superbus. The
tooth is typical of large theropods while the femur has never been illustrated,
though both may indeed belong to Erectopus.
References- Barrois, 1875. Les reptiles du terrain Cr�tac�
du nord-est du Bassin de Paris. Bulletin scientifique, historique et litt�raire
du Nord. 6, 1-11.
Sauvage, 1876. Notes sur les reptiles fossiles no. 9. De la presence du type
dinosaurien dans le Gault du nord de la France. Bulletin de la Soci�t�
G�ologique de France. 4, 439-442.
Sauvage, 1882a. Sur les Reptiles trouv�s dans le gault de l'est de la
France [On the reptiles found in the Gault of eastern France]. Comptes Rendus
Hebdomadaires des Seances de l'Acad�mie des Sciences. 94, 1265-1266.
Sauvage, 1882b. Recherches sur les reptiles trouv�s dans le Gault de
l'est du bassin de Paris [Research on the reptiles found in the Gault of the
eastern Paris Basin]. M�moires de la Soci�t� G�ologique
de France, s�rie 3. 2(4), 1-42.
Huene, 1926a. The carnivorous Saurischia in the Jura and Cretaceous formations,
principally in Europe. Revista Museo de La Plata. 29, 35-167.
Huene. 1926b. On several known and unknown reptiles of the order Saurischia
from England and France. Annals and Magazine of Natural History.17, 473-489.
unnamed Averostra (Barrois, 1875)
Albian, Early Cretaceous
Grandpre, Ardennes, France
Material- (Mus�e de la Facult� des Sciences de Lille coll.)
two teeth, centrum (65 mm) (Barrois, 1875)
(Peron coll.) distal fibula(?), distal metatarsal(?) (Sauvage, 1882b)
Comments- Barrois (1875) and Sauvage (1876) described two teeth and a
centrum from Grandpre as Megalosaurus. Sauvage (1882b) referred the teeth
to Megalosaurus superbus, and described a supposed distal metatarsal
and proximal metapodial from Grandpre as also belonging to M. superbus.
Huene (1926a, b) retained all of these in that species, which he had moved to
the new genus Erectopus in 1923. He listed both limb bones as metatarsals
on page 43, but on page 79 stated the proximal metapodial was a distal fibula.
Huene (1932) later kept the teeth referred to superbus, while he kept
the limb elements with the Erectopus type material as E. sauvagei.
The illustrated tooth and limb bone fragment do not appear particularly diagnostic
and may belong to any ceratosaur, basal tetanurine, carnosaur or basal coelurosaur.
The centrum and supposed distal fibula may not even be theropodan.
References- Barrois, 1875. Les reptiles du terrain Cr�tac�
du nord-est du Bassin de Paris. Bulletin scientifique, historique et litt�raire
du Nord. 6, 1-11.
Sauvage, 1876. Notes sur les reptiles fossiles no. 9. De la presence du type
dinosaurien dans le Gault du nord de la France. Bulletin de la Soci�t�
G�ologique de France. 4, 439-442.
Sauvage, 1882b. Recherches sur les reptiles trouv�s dans le Gault de
l'est du bassin de Paris [Research on the reptiles found in the Gault of the
eastern Paris Basin]. M�moires de la Soci�t� G�ologique
de France, s�rie 3. 2(4), 1-42.
Huene, 1926a. The carnivorous Saurischia in the Jura and Cretaceous formations,
principally in Europe. Revista Museo de La Plata. 29, 35-167.
Huene. 1926b. On several known and unknown reptiles of the order Saurischia
from England and France. Annals and Magazine of Natural History.17, 473-489.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte.
Monographien zur Geologie und Palaeontologie. 4(1), 361 pp.
undescribed averostran (Dollo, 1909)
Hauterivian-Barremian, Early Cretaceous
Weald Clay, Belgium
Material- tooth
Comments- This tooth was referred to Megalosaurus dunkeri by Dollo
(1909), then Altispinax dunkeri by Huene (1926), but has not been described
in detail so cannot be compared to the holotype of the former. It may belong
to Altispinax, Valdoraptor, Neovenator, Calamosaurus, Eotyrannus or another
Wealden theropod.
References- Dollo, 1909. The fossil vertebrates of Belgium. Annals of
the New York Academy of Sciences. 19(4), 99-119.
Huene, 1926. The carnivorous Saurischia in the Jura and Cretaceous formations,
principally in Europe. Revista del Museo de La Plata. 29, 1-167.
unnamed averostran (Huene, 1966)
Toarcian, Early Jurassic
Schleswig-Holstein, Germany
Material- partial dorsal centrum (80 mm)
Comments- This centrum was described by Huene (1966) as a megalosaurid.
It has a convex anterior articular surface.
Reference- Huene, 1966. Ein Megalosauriden- Wirbel des lias aus norddeutschem
Geschiebe. Neues Jahrbuch f�r Mineralogie, Geologie und Pal�ontologie.
1966(5), 318-319.
undescribed averostran (Lubbe, Richter and Knotschke, 2009)
Kimmeridgian, Late Jurassic
Langenberg Quarry, Germany
Material- ?(DFMMh/FV 707.1) partial tooth (Lubbe, Richter and Kn�tschke,
2009)
Comments- While seven teeth assigned to Velociraptorinae by Lubbe et
al. (2009), Gerke and Wings (2014) found four of these were Neotheropoda indet. (presumably sensu Bakker),
megalosaurid and tyrannosauroid. Based on the information in Lubbe et al., FV
707.1 may be indet. (larger and only partially preserved).
References- Lubbe, Richter and Kn�tschke, 2009. Velociraptorine
dromaeosaurid teeth from the Kimmeridgian (Late Jurassic) of Germany. Acta Palaeontologica
Polonica. 54(3), 401-408.
Gerke and Wings, 2014. Characters versus morphometrics: A case study with isolated
theropod teeth from the Late Jurassic of Lower Saxony, Germany, reveals an astonishing
diversity of theropod taxa. Journal of Vertebrate Paleontology. Program and
Abstracts 2014, 137.
unnamed Averostra (Lanser and Heimhofer, 2015)
Late Barremian-Early Aptian, Early Cretaceous
Balve-Beckum quarry, Germany
Material- (LWL MN Ba 1; Morphotype B) partial lateral tooth (?x16.5x7.8
mm)
(LWL MN Ba 2; Morphotype B) partial lateral tooth (?x5.9x3.7 mm)
(LWL MN Ba 3; Morphotype B) partial lateral tooth
(LWL MN Ba 4; Morphotype B) lateral tooth (14.3x9.9x4.8 mm)
(LWL MN Ba 4a; Morphotype B) lateral tooth fragment
(LWL MN Ba 16; Morphotype A) lateral tooth fragment
(LWL MN Ba 18; Morphotype A) lateral tooth fragment
(LWL MN Ba 19; Morphotype A) lateral tooth fragment
(LWL MN Ba 20; Morphotype A) partial lateral tooth (?x8.3x4.1 mm)
(LWL MN Ba 22; Morphotype A) lateral tooth fragment
Reference- Lanser and Heimhofer, 2015. Evidence of theropod dinosaurs
from a Lower Cretaceous karst filling in the northern Sauerland (Rhenish Massif,
Germany). Pal�ontologische Zeitschrift. 89(1), 79-94.
unnamed averostran (Garilli, Klein, Buffetaut, Sander, Pollina, Galletti, Cillari and Guzzetta, 2009)
Late Aptian-Early Albian, Early Cretaceous
Section M, Pizzo Muletta succession, Sicily, Italy
Material- (uncollected) (subadult) partial ?humerus or ?femur
Comments- Discovered in 2005
and only visible in section, Garilli et al. (2009) conclude "the
combination of the open medullary cavity combined with laminar
fibrolamellar bone tissue suggests that the bone from Grotta Lunga
represents a theropod."
References- Garilli, Klein,
Buffetaut, Sander, Pollina, Galletti, Cillari and Guzzetta, 2009. First
dinosaur bone from Sicily identified by histology and its
paleobiogeographical implications. Neues Jahrbuch f�r Mineralogie,
Geologie und Pal�ontologie - Abhandlungen. 252(2), 207-216.
Randazzo, Di Stefano, Schlagintweit, Todaro, Cacciatore and Zarcone,
2021. The migration path of Gondwanian dinosaurs toward Adria:
New insights from the Cretaceous of NW Sicily (Italy). Cretaceous
Research. 126, 104919.
undescribed possible averostran (Codrea, Godefroit and Smith, 2012)
Maastrichtian, Late Cretaceous
Rusca Montana Basin, Romania
Material- (UBB NgTh1) partial tooth
Comments- Codrea et al. (2012)
state UBB NgTh1 "may be tentatively attributed to a troodontid-like
theropod because it displays the following characters (Currie et al.,
1990): the crown is less recurved than in teeth ascribed to
velociraptorines, and both the mesial and distal denticles are well
developed (six denticles per millimeter) and hooklike." Yet the
serrations are not nearly as large as derived troodontids, recurvature
is absent unlike serrated troodontid teeth, and the base is lacking so
that root constriction is unknown. The preserved tip is unusual in
being very low (height ~80% of FABL). This is here referred to
?Averostra.
Reference- Codrea, Godefroit and Smith, 2012. First discovery of Maastrichtian
(Latest Cretaceous) terrestrial vertebrates in Rusca Montana basin (Romania).
In Godefroit (ed.). Bernissart Dinosaurs and Early Cretaceous Terrestrial Ecosystems.
Indiana University Press. 570-581.
unnamed averostran (Codrea, Smith, Dica, Folie, Garcia, Godefroit and
Van Itterbeecke, 2002)
Late Maastrichtian, Late Cretaceous
Sinpetru Beds, Romania
Material- (IRSNB coll.) teeth (~3.1x2.5x? mm)
Comments-
Codea et al. (2002) describe an assemblage collected in 2001, stating
"some fragmentary teeth may be tentatively attributed to
troodontid-like theropods (Fig. 4k), as they display the following
characters: the crown is less recurved than in teeth ascribed to
velociraptorines, both the mesial and distal denticles are well
developed, the mesial denticles extend towards the base of the crown,
distal denticles are wider than long, oblique to the tooth axis and
often hook-like." The figured tooth does not preserve the base to
determine if it had constricted roots, so could easily be e.g. noasaurid instead considering biogeography.
Reference- Codrea, Smith, Dica, Folie, Garcia, Godefroit and Van Itterbeecke,
2002. Dinosaur egg nests, mammals and other vertebrates from a new Maastrichtian
site of the Hateg Basin (Romania). Comptes Rendus Palevol. 1(3), 173-180.
unnamed averostran (Hooijer, 1968)
Cenomanian-Santonian, Late Cretaceous
Qalamoun hill, Syria
Material- (cast AMNH 8254) distal tibia (110
mm transversely)
Comments- Found "some time before" 1965, Hooijer (1968) believed this to probably come from Carcharodontosaurus
or Erectopus, considering both to be megalosaurids. Carrano et al. (2012)
referred it to Tetanurae based on the distal compression (110x55 mm), but an equal amount
is present in Elaphrosaurus
and some abelisaurs. Note while no figures exist in the paper, a
photo in posterior view of the cast AMNH 8254 is on the AMNH online
collections. No mention of a repository exists in the paper,
although Hooijer is listed as working at the Rijksmuseum van
Natuurlijke Historie, which has since become the Naturalis Biodiversity
Center. Unfortunately, the latter has no record of ever acquiring
it (den Ouden, pers. comm. 5-2023), so its location is unknown.
Similarly, the AMNH has no record of the repository of the original,
date collected or dtate when the cast was received (Mehling, pers.
comm. 5-2023).
References- Hooijer, 1968. A Cretaceous dinosaur from the Syrian Arab
Republic. Koninklijke Nederlandse Akademie van Wetenschappen - Amsterdam, Proceedings
Series B. 71, 150-152.
AMNH 2007 online. http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=51261
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
undescribed Averostra (Kellner, Mirzaie Ataabadi, Dalla Vecchia, de
Paulo Silva and Pourbagheban, 2003)
Oxfordian-Hauterivian, Late Jurassic-Early Cretaceous
Ravar Formation, Ab Bid Syncline, Iran
Material-
?(Geological Survey of Iran coll.) limb bone shaft (Kellner, Dalla
Vecchia, Mirzaie Ataabadi, de Paula Silva and Khosravi, 2012)
(MN 7271-V) lateral tooth (36x17x7 mm) (Kellner, Mirzaie Ataabadi, Dalla Vecchia, de
Paulo Silva and Pourbagheban, 2003)
Comments- Note that while
Kellner et al. (2003) initially identified the stratigraphy as "the
uppermost part of the Bidou Formation" based on the 1995 Geological Map
of Iran, "according to the recent revision of the Jurassic stratigraphy
of the southern Tabas Block (Wilmsen et al. 2009) they come from the
Ravar Formation" (Kellner et al., 2012).
Discovered in September 2002, Kellner et al. (2012) noted that "At
least one of the specimens represents the cortex of a long bone that
due to its comparatively small thickness most likely belonged to a
medium-sized theropod dinosaur." The tooth has 3.5 serrations per
mm mesially and 3 serrations per mm distally. Serrations are
described as "straight, longer than wide, and chisel-like" with "blood
grooves ... present as faint impressions, curved towards the basal part
of the tooth." Besides excluding obviously apomorphic taxa like
spinosaurids, carcharodontosaurines and tyrannosaurids, Kellner et al.
(2012) could not specify its relationships further among
theropods. Given its size comparable to Allosaurus or Gorgosaurus
and age, comparisons should be made with ceratosaurids, megalosaurids,
non-carcharodontosaurine carnosaurs, megaraptorans and basal
tyrannosauroids.
References- Dalla Vecchia, Mirzaie
Aatabadi, Kellner, Jafarian, de Paula Silva, Seyfori, Medadi,
Pourbagheban and Khosravi, 2003. Ricerca di dinosauri in Iran. Abstract
Book Giornate di Paleontologia 2003, 12.
Kellner, Mirzaie Ataabadi, Dalla Vecchia, de Paulo Silva
and Pourbagheban, 2003. Theropod dinosaurs from Iran. Simposio Brasileiro de
Paleontologia de Vertebrados III, 34.
Mirzaie Ataabadi, Kellner, Dalla Vecchia and de Paula Silva, 2005. Palaeoichnology
and palaeontology of dinosaurs in north of Kerman, south east central Iran.
Geosciences. 14(54), 36-47.
Kellner, Dalla Vecchia, Mirzaie Ataabadi, de Paula Silva and Khosravi,
2012. Review of the dinosaur record from Iran with the description of
new material. Rivista Italiana di Paleontologia e Stratigrafia. 118(2),
261-275.
unnamed averostran (Schulp, Hanna, Hartman and Jagt, 2000)
Maastrichtian, Late Cretaceous
Al-Khawd, Al-Khod Conglomerate Formation, Oman
Material- (SQU-2-7) (~6-7 m) ~second caudal centrum (92 mm)
Comments- Discovered in 1997, Schulp et al. (2000) described this as much stouter than Majungasaurus, Spinosaurus, Allosaurus or Carcharodontosaurus.
The centrum is slightly amphicoelous, 106% taller than wide, with a
broad median ventral ridge and no pleurocoels. The authors based
the position and total length estimates off of Allosaurus.
Reference- Schulp, Hanna,
Hartman and Jagt, 2000. A Late Cretaceous theropod caudal vertebra from
the Sultanate of Oman. Cretaceous Research. 21, 851-856.
undescribed Averostra (Nessov, 1995)
Mid-Late Turonian, Late Cretaceous
Bissekty Formation, Uzbekistan
Material- (CCMGE 450/12457) partial braincase (Nessov, 1995)
several braincase fragments (Sues and Averianov, 2004)
Comments- Nessov (1995) figured CCMGE 450/12457 as a "relatively
large carnosaur". Sues and Averianov (2004) referred several braincases
to Itemirus, but these were not mentioned in Sues and Averianov's (2014)
resulting revision of the genus. Sues (pers. comm. 2014) stated these were fragments
that showed few diagnostic features once prepared, so their identification remains
uncertain. Nessov also tentatively referred a basisphenoid fragment (CCMGE 719/12457)
to a segnosaur, but Sues and Averianov (2016) stated it was "too fragmentary
for more precise taxonomic identification beyond Theropoda." Given the
age and location, all of these are probably coelurosaurs.
References- Nessov, 1995. Dinosaurs of Northern Eurasia: New data about
assemblages, ecology, and paleobiogeography. Institute for Scientific Research
on the Earth's Crust, St. Petersburg State University, St. Petersburg. 1-156.
Sues and Averianov, 2004. Dinosaurs from the Upper Cretaceous (Turonian) of
Dzharakuduk, Kyzylkum Desert, Uzbekistan. Journal of Vertebrate Paleontology.
24(3), 51A-52A.
Sues and Averianov, 2014. Dromaeosauridae (Dinosauria: Theropoda) from the Bissekty
Formation (Upper Cretaceous: Turonian) of Uzbekistan and the phylogenetic position
of Itemirus medullaris Kurzanov, 1976. Cretaceous Research. 51, 225-240.
Sues and Averianov, 2016 (online 2015). Therizinosauroidea (Dinosauria: Theropoda) from the
Upper Cretaceous of Uzbekistan. Cretaceous Research. 59, 155-178.
undescribed averostran (Godefroit, Sinitsa, Dhouailly, Bolotsky, Sizov,
McNamara, Benston and Spagna, 2014)
Aalenian, Middle Jurassic
Ukureyshaya Formation, Russia
Holotype-(INREC K coll.) (medium-sized) tooth
Comments- Godefroit et al. (2014) report that besides the numerous specimens
of Kulindadromeus preserved in the Kulinda bonebeds, only a single shed
theropod tooth is present. Two specimens were referred to Theropoda by Alifanov-
PIN 5435/51 (distal caudal vertebrae with associated scales) and a specimen
that consists of three distal metapodials, phalanges and scales. These were
later named Lepidocheirosaurus by Alifanov and Saveliev (2015), but one
and probably both are Kulindadromeus specimens (see entry).
References- Alifanov, 2014. The discovery of Late Jurassic dinosaurs
in Russia. Doklady Earth Sciences. 455(2), 365-367.
Godefroit, Sinitsa, Dhouailly, Bolotsky, Sizov, McNamara, Benston and Spagna,
2014. A Jurassic ornithischian dinosaur from Siberia with both feathers and
scales. Science. 345(6195), 451-455.
Alifanov and Saveliev, 2015. The most ancient ornithomimosaur (Theropoda, Dinosauria),
with cover imprints from the Upper Jurassic of Russia. Paleontological Journal.
49(6), 636-650.
undescribed Averostra (Averianov, Skutschas, Danilov, Krasnolutskii
and Martin, 2014)
Bathonian, Middle Jurassic
Upper Itat Formation, Russia
Material- teeth
Reference- Averianov, Skutschas, Danilov, Krasnolutskii and Martin, 2014.
Non-mammalian vertebrate assemblage from the Middle Jurassic of Berezovsk Quarry
in western Siberia. Journal of Vertebrate Paleontology. Program and Abstracts
2014, 83.
unnamed Averostra (Kurzanov, Efimov, and Gubin, 2003)
Callovian-Oxfordian, Middle Jurassic-Late Jurassic
Djaskoian Formation, Russia
Material- (PIN 4874/2) six teeth (10-15 mm)
Comments- These were referred to Allosaurus sp. by Kurzanov et
al. (2003) because they were said to be similar to "Labrosaurus"
stechowi and "Labrosaurus" (=Ceratosaurus) sulcatus, and
the type species of Labrosaurus (L. lucaris) is a junior synonym
of Allosaurus. However, the Djaskoian teeth appear to lack the distinctive
fluting found in "L." stechowi and C. sulcatus (which
are both ceratosaurids), while L. lucaris doesn't preserve teeth. Carrano
et al. (2012) believed they could not be identified past Theropoda indet..
References- Kurzanov, Efimov, and Gubin, 2003. New archosaurs from the
Jurassic of Siberia and Mongolia. Paleontological Journal. 37(1), 53-57.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
unnamed possible averostran (Riabinin, 1937)
Late Barremian-Mid Aptian, Early Cretaceous
Mogoito Member of Murtoi Formation, Russia
Material- distal metapodial or phalanx
Comments- A distal phalanx was discovered in 1931 and referred to Theropoda
indet. by Riabinin (1937). Nessov (1995) considered this to be a possibly sauropod
metapodial or phalanx. Averianov et al. (2003) referred it to Therizinosauridae
based on the unequally deep collateral ligament pits, but Zanno (2008) noted
both sides having well defined pits (albeit better developed on one side) is
unlike therizinosaurs. Whether this specimen is indeed a theropod at all is
still uncertain.
References- Riabin, 1937. A new discovery of dinosaurs in Transbaikalia.
Ezhegodnik Vsesoyuznogo Paleontologicheskogo Obshchestva. 11, 141-144.
Nessov, 1995. Dinosaurs of nothern Eurasia: New data about assemblages, ecology,
and paleobiogeography. Institute for Scientific Research on the Earth's Crust,
St. Petersburg State University, St. Petersburg. 1-156.
Averianov, Starkov and Skutschas, 2003. Dinosaurs from the Early Cretaceous
Murtoi Formation in Buryatia, Eastern Russia. Journal of Vertebrate Paleontology.
23(3), 586-594.
Zanno, 2008. A taxonomic and phylogenetic reevaluation of Therizinosauria (Dinosauria:
Theropoda): Implications for the evolution of Maniraptora. PhD Thesis. The University
of Utah. 329 pp.
unnamed averostran (Watabe, Tsogtbaatar, Uranbileg and Gereltsetseg, 2004)
Late Jurassic
Dariv Formation, Mongolia
Material- (IGM 107/5) (skull ~354 mm) maxillary fragments, dentary fragments, teeth (FABL 11.1, 9.8, 11, 12.7, 8.5, 10.5, 10 mm)
Comments- Watabe et al. (2004)
note "the articulated lower and upper jaws of theropod with teeth" were
found between Juily 25 to August 17 in the Dariv Formation. The
specimen was described by Watabe et al. (2006) as Theropoda indet.,
though they stated "the maxillary and dentary ziphodont teeth of this
specimen suggest that the specimen belongs to a basal tetanuran
theropod", comparing it favorably to Monolophosaurus and sinraptorids. Yet it is also similar to e.g. ceratosaurids, so might be more properly considered Averostra indet..
References- Watabe, Tsogtbaatar, Uranbileg and Gereltsetseg, 2004. Report on the Japan - Mongolia Joint Paleontological Expedition to the
Gobi desert, 2002. Hayashibara Museum of Natural Sciences Research
Bulletin. 2, 97-122.
Watabe, Tsogtbaatar and Barsbold, 2008. First discovery of a theropod
(Dinosauria) from the Upper Jurassic in Mongolia and its stratigraphy.
Paleontological Research. 12(1), 27-36.
undescribed averostran (Gubin and Sinitza, 1996)
Late Jurassic
Shar Teg Formation, Mongolia
Material- (PIN coll.) tooth
Reference- Gubin and Sinitza,
1996. Shar Teg: A unique Mesozoic locality of Asia. In Morales (ed.).
The Continental Jurassic. Museum of Northern Arizona Bulletin. 60,
311-318.
undescribed Averostra (Eberth, Kobayashi, Lee, Mateus, Therrien,
Zelenitsky and Norell, 2009)
Santonian-Campanian?, Late Cretaceous
Javkhlant Formation, Mongolia
Comments- Eberth et al. (2009) noted "small nonavian theropod dinosaur
skeletal remains" and "several taxa of variously-sized theropod dinosaurs"
from the Javkhlant Formation, of which only Albinykus has been described
so far. Based on their location and age, these are probably all coelurosaurs.
Reference- Eberth, Kobayashi, Lee, Mateus, Therrien, Zelenitsky and Norell,
2009. Assignment of Yamaceratops dorngobiensis and associated redbeds
at Shine Us Khudag (eastern Gobi, Dorngobi Province, Mongolia) to the redescribed
Javkhlant Formation. Journal of Vertebrate Paleontology. 29(1), 295-302.
Averostra indet. (Young, 1942)
?
Shaanxi or Gansu, China
Material- (IVPP V192) (~8 m) partial anterior dorsal centrum
(IVPP V204) partial jugal
(IVPP V209) postorbitals
Comments- Young (1942) in his
"description of other fragmentary bones" states "As mentioned above
there are a series of cranial and post-cranial bones, mostly
fragmentary, which cannot be attributed to a definite systematic
position with certainty", which seems to reference "A few fragments of
vertebrate fossils recovered also by us during the same trip in Shensi
and Kansu", now known as Shaanxi and Gansu. Thus there is no
stratigraphic data recorded.
Young (1942) describes IVPP V192 as "An anterior part of a centrum with
the breadth 81 mm., middle constriction 50 mm" and notes that "In
ventral aspect, there is a prominent ridge developed" which would make
it an anterior dorsal in most theropods. Its size is comparable to Neovenator's type (first dorsal 84 mm wide anteriorly and 50 mm wide at constriction). It was stated to "fit in size with Chienkosaurus ceratosauroides", but not explicitly referred and instead regarded as "Probably a dorsal vertebra of a Theropoda."
Young (1942) states that "One other much bigger bone suggests a part of
jugal" in reference to IVPP V204, the size comparison being to IVPP
V209.
For IVPP V209, Young (1942) states "Of the cranial element there are
two bones with the proximal end expanded and thick. The beam narrows
distally but the tip is broken. There is a ridge developed at the
proximal external part. They can be bestly regarded as the postorbital.
Size fits with the big Theropeda described above" [Szechuanosaurus and Chienkosaurus]. The ridge suggests a postorbital boss as in metriacanthosaurids, carcharodontosaurids and tyrannosaurids.
Reference- Young, 1942. Fossil vertebrates from Kuangyuan, N. Szechuan,
China. Bulletin of the Geological Society of China. 22(3-4), 293-309.
undescribed Averostra (Dong, Zhou and Zhang, 1983)
Aalenian, Middle Jurassic
Xintiangou Formation, Sichuan, China
Comments- Dong et al. (1983) stated "Vertebrate occurrences ... in
the Xintiangou Fm., as they consist merely of ... Carnosauria ...",
written long before the Laojun Village deposits yielding
"Yunyangosaurus" were discovered in 2016. This may refer to IVPP V893,
referred to Carnosauria indet. by Liu and Yeh (1957; here identified as
an afrovenatorine) and possibly from the contemporaneous Qianfoya
Formation.
References- Liu and Yeh, 1957. Two new species of Ceratodus from Szechuan, China. Vertebrata PalAsiatica. 1(4), 305-311.
Dong, Zhou and Zhang, 1983. Dinosaurs from the Jurassic of Sichuan. Palaeontologica
Sinica. Whole Number 162, New Series C, 23, 136 pp.
undescribed Averostra (Hao, Zhang, Peng and Ye, 2022)
Bajocian, Middle Jurassic
Qinglongshan, Xiashaximiao Formation, Sichuan, China
Material- (ZDM coll.) five teeth (~30x~15x? mm)
Comments- The Qinglongshan
fossils were excavated in March 1995, April 1999 and May-June
2019. Hao et al. (2022) note "scattered theropod teeth that are
laterally flat, distally curved along the crown, have a distal edge
thinner than the mesial edge, and are serrated along both edges."
They suggest "These characteristics suggest that the teeth are those of
an indeterminate carnivore theropod, and similar in size to those of Szechuanosaurus" and list a preliminary identification of "Szechuanosaurus?".
Reference- Hao, Zhang, Peng and
Ye, 2022. Discovery of a new Middle Jurassic dinosaur site in Sichuan,
China. Acta Geologica Sinica (English Edition). 96(1), 52-60.
unnamed averostran (Camp, 1935)
Middle Jurassic?
Jung-Hsein UCMP V1501, middle Chongqing Group, Sichuan, China
Material- (UCMP 32102) (~14.7 m) mesial dentary tooth (~69x~22x? mm), rib fragment, ischial fragment, femoral fragment (~1.33 m)
Comments-
The specimen was collected on August 30 1915 by Louderback. Note the
UCMP locality number is V1501 (as determined in their online
catalogue), not V151 as listed by Camp. Jung-Hsien is now called
Rongxian, a county in Zigong City. Camp stated "The beds in which they
occur have been called the Szechuan series", which was a term for the
stratigraphic section from the Early Jurassic Qianfuyan (= Tsienfuyan)
Formation and Ziliujing (= Tsuliuching, = Tzeliutsin) Formation to the
Cretaceous Chengqiangyan (= Chengtsiangyen) Group and Jiading (=
Chiating, = Tshiating) Group, depending on north versus south in the
Sichuan Basin. Rongxian is located in the south, so UCMP V1501 would be
part of the Ziliujing-Chongqing-Jiading sequence, and Young (1937; see
also Young et al., 1943) placed it above the Ziliujing Formation but
below the conglomerates of the Jiading Group, and thus within the
Middle-Late Jurassic Chongqing Group. Furthermore, Young (1937) stated
"The fossiliferous horizon discovered by Louderback lies probably
between our horizons 2 and 3, some 200 meters above horizon 2" which is
the type locality of Omeisaurus junghsiensis. As Omeisaurus
is generally recovered in the Xiashaximiao Formation and horizon 3 is
another 300 meters above where Young placed UCMP V1501 (so may be the
Penglaizhen or Suining Formation), UCMP 32102 may derive from the
Shangshaximiao Formation. Dong et al. (1983) listed it as deriving
from that formation, perhaps using the same logic although they did not
describe any explanation.
Camp (1935) initially referred the specimen to Megalosauridae because
histology "shows quite definitely that the relationship of the Chinese
form is with Allosaurus" instead of Tyrannosaurus. However, the plate shows this is because the sampled section of Tyrannosaurus femur (labeled AMNH 5886, but this is the Anatotitan paratype, and it is more probably Dynamosaurus holotype AMNH 5866 that is known to be histologically sampled) is composed of secondary osteons, while those of Allosaurus and UCMP 32102 are fibrolamellar bone. Yet Allosaurus
can develop secondary osteons where bone is redeveloped as well (e.g.
MHNG GEPI V2567a), so this isn't a real difference between these taxa.
While Camp wrote "a projection of the borders would indicate an
original total length of at least 90 mm" for the tooth, he also
repeated Osborn's 1906 statement that Tyrannosaurus
(CM 9380 and NHMUK R7994) teeth are up to 125 mm, which includes the
root. Combined with his statement the serrated distal carina "reaches
the base of the enamel" and serrations are not illustrated on the most
basal section, the actual crown length would have been about 69 mm.
Similarly, Camp wrote "At the base it is 17 mm. in longest diameter",
but scaling the figured tooth to the stated preserved length of 60 mm
results in a FABL of 22 mm instead. Young (1942) later wrote "The
general structure of [Szechuanosaurus campi
syntype] V236 with the way of serrations fits so well with the
Junghsien tooth, we feel that there is practically no doubt in
regarding them as identical" "and prefer to consider the Junghsien
tooth as belonging also to the new form" Szechuanosaurus. This despite previously stating UCMP 32102 "is bigger and straighter than all" Szechuanosaurus syntype teeth. Compared to Sinraptor dongi and Szechuanosaurus,
UCMP 32102 is larger (~69 vs. up to 63 vs. ~32 and ~47 mm), with a much
greater crown height/base ratio (~314% vs. up to 244% vs. ~224% and
~267%), making it less tapered. The crown section is similar to Allosaurus' fourth dentary tooth and the mid crown ratio of 53% is similar to dentary teeth in S. dongi and between S. campi IVPP V238B and 238C. As in mesial teeth of S. dongi,
the crown is slightly lingually curved and the mesial carina does not
reach the crown base. The same serration densities (mesial 15 per 5
mm, distal 6.7-10 per 5 mm) can be found in S. dongi as well, but are lower than S. campi
(distal ~12-19 per 5 mm). Dong et al. (1983) wrote "Camp's description
and the dentition size suggests it may be assignable to Yangchuanosaurus", but it is larger than even the magnus
type (up to 75 mm), and more elongate than the largest maxillary teeth
of the genotype (crown height/base ratio of 267%), but detailed dental
statistics of the genus have yet to be published. The femoral fragment
is notably large, Camp stating the shaft has "an enormous hollow cavity
about 125 mm. in the longest diameter of its ellipse. The greatest
diameter of this segment at its narrowest point is 20 cm." Based on
the absence of a fourth trochanter or medial narrowing, the section is
just distal to the former structure. Here large theropod femoral
shafts are wider than deep, so 200 mm would be the width and the
figured depth is then ~143 mm. Scaling from the largest
metriacanthosaurid, Yangchuanosaurus magnus,
results in a femoral length of ~1.33 meters, not far from Camp's
"estimated total length of about 140 cm." The ischium "consists of a
moderately hollow shaft spreading into a broader, solid plate", which
could describe most non-maniraptoran ischia, while the "tip of a large
rib" is not described. Notably, the ischium was found 37 meters from
the other material, so its association is less certain.
Given the above information, UCMP 32102 is different from the Szechuanosaurus syntypes and anteriorly straighter than Sinraptor
as well, and is perhaps the largest known Jurassic theropod. As no
characters are outside the range of ceratosaurids, it is considered
Averostra incertae sedis here.
References- Camp, 1935. Dinosaur remains from the province of Szechuan.
University of California Publications, Bulletin of the Department of Geological
Sciences. 23(14), 467-471.
Louderback, 1935. The stratigraphic relations of the Jung Hsien fossil
dinosaur in Szechuan red beds of China. University of California
Publications. Bulletin of the Department of Geological Sciences.
23(14), 459-466.
Young, 1937. New Triassic and Cretaceous reptiles in China (With some
remarks concerning the Cenozoic of China). Bulletin of the Geological
Society of China. 17(1), 109-120.
Young, 1942. Fossil vertebrates from Kuangyuan, N. Szechuan, China. Bulletin
of the Geological Society of China. 22(3-4), 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.
Dong, Zhou and Zhang, 1983. Dinosaurs from the Jurassic of Sichuan. Palaeontologica
Sinica. Whole Number 162, New Series C, 23, 136 pp.
undescribed Averostra (Young and Chow, 1953)
Middle-Late Jurassic(?)
Chongqing Group?, Sichuan, China
Material- (IVPP V712) (~3 m) two dorsal vertebrae (53 mm)
Middle-Late Jurassic
Chongqing Group, Jianyang, Sichuan, China
(IVPP V713) (~8 m) two femoral fragments (distal transverse width 210 mm) (~940 mm)
Comments- These were discovered
"during the construction of the Chengtu-Chungking Railway mainly in
1951" and/or "during the clearing away the building foundation in
Tatienwan, a suburb of Chungking." The latter is "generally
acccepted as
Chungking Series of Cretaceous age" according to Young and Chow (1953),
today called the Chongqing Group and considered of Middle-Late Jurassic
age. Most of the sediments exposed on the surface between
Chongqing and Chengdu are also Jurassic, so fossils discovered
constructing the railway between them are likely also from the
Chongqing Group. Young and Chow referred both to Carnosauria
indet..
Young and Chow called the two dorsals "A small form", stated "Exact
locality unknown" and "think more probable to regard these as belonging
to carnivorous dinosaur." Their length suggests an individual
about 3 meters long, so would probably be a young specimen if their
referral to Carnosauria sensu Huene is correct (i.e. a megalosauroid or
carnosaur), or it may be an elaphrosaurine or coelurosaur.
Young and Chow called the femoral fragments "A large form" and stated
"Its slenderness suggests a carnosaurian" and that the "Transversal
breadth of the distal end, 210 mm." The latter is very similar to
Sinraptor dongi's holotype
femur's distal width of 195 mm, which would make its length 943 mm if
scaled identically. This suggests a neoceratosaur, megalosauroid
or carnosaur. They also state one of the fragments "bears the
label indicating Chien Yang district", now known as Jianyang.
References- Young and Chow, 1953a. New fossil reptiles from Szechuan, China. Acta Palaeontologica Sinica. 1(3), 87-109.
Young and Chow, 1953b. New fossil reptiles from Szechuan, China. Acta Scientia Sinica. 2(3), 216-243.
unnamed Averostra (Dong, 1993)
Bathonian-Callovian, Middle Jurassic
Toutunhe Formation, Xinjiang, China
Material- (IVPP coll.) two teeth, several limb elements including pedal phalanx (?)I-1, two unguals including manual ungual (?)II
Comments- This material was discovered in the same quarry as Tianchisaurus
in 1974, with some of it figured by Dong (1993) as Megalosauridae
indet.. As Maisch and Matzke (2003) noted, "There appears little
reason for the assignment of this material to the Megalosauridae as
proposed by DONG (1993), because it could equally well represent an
allosauroid." Their identifications of the phalangeal elements as
listed here are plausible.
References- Dong, 1993. An ankylosaur (ornithischian
dinosaur) from the Middle Jurassic of the Junggar Basin, China.
Vertebrata PalAsiatica. 31, 257-266.
Maisch and Matzke, 2003. Theropods (Dinosauria, Saurischia) from the Middle
Jurassic Toutunhe Formation of the southern Junggar Basin, NW China. Palaeontologische
Zeitschrift. 77(2), 281-292.
Averostra indet. (Dong, 1973)
Middle-Late Juarssic
Kizinur Valley, Kuqa, Xinjiang, China
Material- (IVPP V3013; field number 993) tooth (45x~15x? mm)
Comments- Dong (1973) briefly
describes and figures this tooth, discovered in 1956. The photo shows
the typical recurved megalosaur-grade tooth, which Dong states has
small mesial and distal serrations. He says the tooth's features are
close to that of Chienkosaurus,
but merely refers it to Megalosauridae indet.. It is probably a
non-coelurosaur tetanurine, but may be ceratosaurid or tyrannosauroid
instead.
Reference- Dong. 1973. 新疆库车一肉食龙牙化石 [A fossil carnosaur tooth from Kuqa county, Xinjiang]. Vertebrata PalAsiatica. 11(2), 217.
undescribed probable averostran (Young, 1958)
Late Jurassic
Machiahukou IVPP locality 5672, Xiangtang Formation, Gansu, China
Material- (IVPP coll.) (small) vertebra
Comments- Discovered with Mamenchisaurus hochuanensis
paratype IVPP V946 in Summer 1956, Young (1958) mentions this as "a
small vertebra of probably a theropod from Machiahukou." As shown by
his Figure 2, this is located about 200 meters northwest of
Haishiwanzhen. Weishampel (1990) lists this as the Hantong Formation
in the Oxfordian, both stratigraphy and age credited to "Dong pers.
comm.", but Dong (1992) and most other sources list it as the Late
Jurassic Xiangtang Formation.
References- Young, 1958. New sauropods from China. Vertebrata Palasiatica. 2(1), 1-29.
Weishampel, 1990. Dinosaurian distribution. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria. University of California Press. 63-139.
Dong, 1992. Dinosaurian Faunas of China. Ocean Press/Springer-Verlag.
188 pp.
undescribed averostran (Xu, Zheng, Sullivan, Wang, Xing, Wang, Zhang,
O'Connor, Zhang and Pan, 2015)
Oxfordian, Late Jurassic
Tiaojishan Formation, Hebei, China
Material- (IVPP V20796) (small) skeleton
Reference- Xu, Zheng, Sullivan, Wang, Xing, Wang, Zhang, O'Connor, Zhang
and Pan, 2015. A bizarre Jurassic maniraptoran theropod with preserved evidence
of membranous wings. Nature. 521, 70-73.
unnamed Averostra (Maisch, Matzke, Pfretzschner, Ye and Sun, 2001)
Oxfordian, Late Jurassic
Qigu Formation, Xinjiang, China
Material- (GPIT SGP 2000/1) anterolateral tooth (43x18x12 mm) (Maisch, Matzke, Pfretzschner, Ye and Sun, 2001)
(GPIT SGP 2000/2) distal (?)ischium (Maisch, Matzke, Pfretzschner, Ye and Sun, 2001)
(GPIT SGP 2001/1) lateral tooth (~75x25.5x15.5 mm) (Maisch and Matzke, 2003)
(GPIT SGP 2001/2) lateral tooth (65.5x23.5x13.5 mm) (Maisch and Matzke, 2003)
(GPIT SGP 2001/3) incomplete anterolateral tooth (Maisch and Matzke, 2003)
(GPIT SGP 2001/4) anterior dentary tooth (~45x17x12.5 mm) (Maisch and Matzke, 2003)
(GPIT SGP 2001/5) incomplete tooth (16+x9.5x5 mm) (Maisch and Matzke, 2003)
(PMOL-SGP 2004/2) (multiple individuals) tooth fragments (Wings, Tutken, Fowler, Martin, Pfretzschner and
Sun, 2015)
(PMOL-SGP 2005/1) tooth (18 mm) (Wings, Tutken, Fowler, Martin, Pfretzschner and
Sun, 2015)
Comments- Maisch et al. (2001) figured GPIT SGP 2000/1 as a
carnosaur tooth. Maisch and Matzke (2003) described this and four teeth
from another locality as Carnosauria sensu lato indet., stating "there
are no striking features to indicate that they are derived from more
than one form." Discovered in 2000, Maisch et al. (2001) labeled
GPIT SGP 2000/2 the fibula of a coelurosaur sensu Huene. Maisch and
Matzke (2003) believed it was most similar to Sarcosaurus and
Coelophysis rhodesiensis
based on the anteroproximal inclination of the distal articular facet,
referring it to Coelophysoidea or basal Coelurosauria. However,
the
element is dissimilar from theropod fibulae in being significantly and
gradually expanded distally, so it is proposed here to be a distal
ischium instead (e.g. compare to Guanlong). The tooth GPIT SGP 2001/5
was assigned to Coelophysoidea by Maisch and Matzke based on the large
number of serrations (33-37/5mm) and otherwise plesiomorphic
morphololgy, but this is unlikely for a Late Jurassic taxon so it is
here retained as Averostra indet. pending study. Augustin et
al. (2020) revised their stratigraphic assignment from the Toutunhe
Formation to the overlying Qigu Formation, questionably referring the
large teeth to Metriacanthosauridae based on geography although they
did not distinguish them from other carnosaurs, megalosaurids or
ceratosaurids. They stated the small tooth "probably represents a
coelurosaur based on the geological age of the specimen."
References- Maisch, Matzke, Pfretzschner, Ye and Sun, 2001. The fossil
vertebrate faunas of the Toutunhe and Qigu Formations of the southern Junggar
Basin and their biostratigraphical and palecological implications. In Sun, Mosbrugger,
Ashraf and Wang (eds.). The Advanced Study of Prehistory Life and Geology of
Junggar Basin, Xinjiang, China. 83-94.
Maisch and Matzke, 2003. Theropods (Dinosauria, Saurischia) from the Middle
Jurassic Toutunhe Formation of the southern Junggar Basin, NW China. Palaeontologische
Zeitschrift. 77(2), 281-292.
Wings, Tutken, Fowler, Martin, Pfretzschner and Sun, 2015.
Dinosaur teeth from the Jurassic Qigu and Shishugou Formations of the Junggar
Basin (Xinjiang/China) and their paleoecologic implications. Palaontologische
Zeitschrift. 89(3), 485-502.
Augustin, Matzke, Maisch and Pfretzschner, 2020. A theropod dinosaur
feeding site from the Upper Jurassic of the Junggar Basin, NW China.
Palaeogeography, Palaeoclimatology, Palaeoecology. 560, 109999.
unnamed Averostra (Russell and Zheng, 1990)
Late Callovian-Early Oxfordian, Middle-Late Jurassic
Jiangjunmiao, Lower Shishugou Formation, Xinjiang, China
(IVPP coll.) (small) fragmentary skeleton (Dong, 1992)
Late Callovian-Early Oxfordian, Middle-Late Jurassic
Pingfengshan, Lower Shishugou Formation, Xinjiang, China
(IVPP coll.) (small) skeletons (Russell and Zheng, 1990)
Late Callovian-Early Oxfordian, Middle-Late Jurassic
Wucaiwan, Lower Shishugou Formation, Xinjiang, China
(IVPP V15858) incomplete tooth (~35x~15x8.0 mm) (Han, Clark, Xu, Sullivan, Choiniere and Hone, 2011)
Late Oxfordian, Late Jurassic
Giant's Tomb, Upper Shishugou Formation, Xinjiang, China
Material- (IVPP coll.; not collected?) (large) femur (Wings, Schwarz-Wings and Fowler, 2011)
(PMOL-SGP 2006/2) lateral tooth (46 mm) (Wings, Tutken, Fowler, Martin, Pfretzschner and Sun, 2015)
(PMOL-SGP 2006/3) lateral tooth (17 mm) (Wings, Tutken, Fowler, Martin, Pfretzschner and Sun, 2015)
(PMOL-SGP 2006/4) anterior tooth (15 mm) (Wings, Tutken, Fowler, Martin, Pfretzschner and Sun, 2015)
Late Oxfordian, Late Jurassic
Pingfengshan, Upper Shishugou Formation, Xinjiang, China
(IVPP coll.) elements including unguals (Russell and Zheng, 1990)
Late Oxfordian, Late Jurassic
TBB 2002, Wucaiwan, Upper Shishugou Formation, Xinjiang, China
?(IVPP V15302) specimen lacking skull and forelimbs but preserving
seventh cervical vertebra, four anterior dorsal vertebrae, several
posterior dorsal vertebrae, several dorsal ribs, sacrum,
first-~twenty-first caudal vertebrae, nine chevrons, ilium (~179 mm),
ischium and incomplete femur (Eberth, Xu and Clark, 2010)
Comments- Dong (1992) notes in
1988 before July, the Sino-Canadian expedition collected "small
theropods" in Xinjiang. Dong (1993) specified "In May 1989, a
party under X.-J. Zhao (IVPP), D. Brinkman (Royal Tyrrell Museum of
Palaeontology, RTMP), and D. Russell (Canadian Museum of Nature, CMN)
established a camp near the Pingfengshan locality in the Wucaiwan
region" and "Small theropod skeletons discovered the previous year were
collected." Similarly, Russell and Zheng (1990) said "The lower
part of the Shishugou Formation yielded ... the skeleton of a small
theropod discovered during the 1988 season" at Pingfengshan that was
excavated in 1989.
Russell and Zheng (1990) also reported "Isolated theropod bones,
including claws, were found in the higher levels of this formation"
collected at Pingfengshan in 1989.
Zhao and Currie (1994) mention "an unidentified small theropod" from the same horizon as Monolophosaurus,
which they called the Wucaiwan Formation but has since been joined with
the Shishugou Formation. This is probably the same as "A very
fragmentary specimen of a small theropod that could be new" excavated
by the Sino-Canadian expedition in 1987 mentioned by Dong (1992) and
listed as "Coelurosauria indet."
Eberth et al. (2010) note a skeleton discovered in 2002 as "a third,
unidentified taxon that is neither tyrannosauroid nor ceratosaur" and
thus not Guanlong or Limusaurus.
They call it "undescribed small theropod" and figure it in situ.
The low dorsal neural spines, squared postacetabular process, slender
and distally expanded ischium and slender unexpanded chevrons through
caudal eleven suggest something compsognathid grade, so perhaps this is
an adult Aorun. Notably
the distal caudal prezygapophyses are deep and ~80% of centrum length,
inviting comparison to coelophysoids and ornithomimosaurs when the
previous characters are considered. Either identification would
be notable temporally. Stiegler (2019) calls it "an as yet
unidentified theropod without a skull or forelimbs" and states "no
vertebrae anterior to C7 are preserved with V15302..."
Discovered by the Sino-American expeditions between 2001 and 2010, IVPP V15858 is called
Morphotype 3 by Han et al. (2011), who refer it to a ceratosaur, basal tetanurine
or basal tyrannosauroid.
Wings et al. (2011) reported that in April 2006 "our excavation was
cancelled prematurely after only nine field days and several partially
exposed bones (e.g., ... a large theropod femur) had to remain in the
field at the "Giant’s Tomb" site", but in Summer 2006 the quarry "has
been excavated by a field crew from the IVPP (Institute of Vertebrate
Paleontology and Paleoanthropology, Beijing), led by Xu Xing."
Whether the femur was excavated by the IVPP is unknown. From the
upper horizon, "theropod and sauropod teeth which will be published
elsewhere" noted in 2011 were catalogued as PMOL-SGP 2006/2-4 and
described as Theropoda Familia indet. by Wings et al. (2015).
PMOL SGP 2006/2 and 2006/4 have mesial serrations while in 2006/3 a
mesial carina "is not observed on PMOL-SGP 2006/3 (possibly
abraded)." "PMOL-SGP 2006/2-4 possesses 17, 27, and 13 denticles
per 5 mm of the distal carina, respectively", suggesting at least
2006/3 is a different taxon than the other two, but in the absence of
obvious enamel wrinkles or detailed metrics they are all assigned
Averostra here.
References- Russell and Zheng, 1990. The 1989 field season of the Dinosaur Project. Vertebrata PalAsiatica. 28(4), 322.
Dong, 1992. Dinosaurian Faunas of China. Ocean Press/Springer-Verlag.
188 pp.
Dong, 1993. The field activities of the Sino-Canadian Dinosaur Project
in China, 1987-1990. Canadian Journal of Earth Sciences. 30(10),
1997-2001.
Grady, 1993. The Dinosaur Project: The Story of the Greatest
Dinosaur Expedition Ever Mounted. Macfarlane
Walter & Ross and The Ex Terra Foundation. 261 pp.
Zhao and Currie, 1994 (as 1993). A large crested theropod from the Jurassic of Xinjiang,
People's Republic of China. Canadian Journal of Earth Sciences. 30(10), 2027-2036.
Eberth, Xu and Clark, 2010. Dinosaur death pits from the Jurassic of China. Palaios. 25(2), 112-125.
Han, Clark, Xu,
Sullivan, Choiniere and Hone, 2011. Theropod teeth from the
Middle-Upper Jurassic Shishugou Formation of northwest Xinjiang, China.
Journal of Vertebrate Paleontology. 31(1), 111-126.
Wings, Schwarz-Wings and Fowler, 2011. New sauropod material from the
Late Jurassic part of the Shishugou Formation (Junggar Basin, Xinjiang,
NW China). Neues Jahrbuch f�r Geologie und Pal�ontologie -
Abhandlungen. 262(2), 129-150.
Wings, Tutken, Fowler, Martin, Pfretzschner and Sun, 2015 (online 2014).
Dinosaur teeth from the Jurassic Qigu and Shishugou Formations of the Junggar
Basin (Xinjiang/China) and their paleoecologic implications. Palaontologische
Zeitschrift. 89(3), 485-502.
Stiegler, 2019. Anatomy, systematics, and paleobiology of noasaurid
ceratosaurs from the Late Jurassic of China. PhD thesis, The George
Washington University. 693 pp.
Averostra indet. (Young, 1963)
Late Jurassic-Early Cretaceous?
Tiekuling IVPP locality 16611, Jiangxi, China
Material- (IVPP V2691) incomplete tooth (~43x18x? mm)
Comments- Discovered in 1962 or
1963, while reported as being "from Tiekuling, Taiho district,
Chiangsi", this is now called Tiahe County, Jiangxi, and the only
Google result for 'Tiekuling' is this paper. Google translates the
locality from the Chinese section of text as "Que
Pi Ling Water Pond in Taihe County, Jiangxi Province." No
stratigraphic data are given, only that "Its geological age is regarded
as late Jurassic or early Cretaceous."
The
tooth is said to have ~14-15 serrations per 5 mm distally near the tip,
and a greater density near the base. While the text states "Total
length of the tooth is estimated to be 60 mm", the figure would
indicate an estimated length of ~43 mm from the preserved base to their
estimated tip. This makes it comparable in height and FABL to the
estimated tenth maxillary tooth of Sinraptor dongi,
and the thirteenth maxillary tooth of the latter has 15 serrations per
5 mm near the tip and 20 near the base, also matching IVPP V2691.
Young wrote "the tooth looks very similar to those of Szechuanosaurus
found in Kuangyuan", the syntypes of the genus, but its shape doesn't
match any of them although the distal serration density falls within
their range (~12-19 per 5 mm). He referred it to Carnosauria gen. et
sp. indet.. While the few known details are congruent with a
metriacanthosaurid posterior maxillary tooth, they also match
ceratosaurid dentition, so IVPP V2691 is placed in Averostra indet.
here.
Reference- Young, 1963. Note on a new locality of dinosaurian remains
from Taiho, Chiangsi, SE. China. Vertebrata PalAsiatica. 7(1), 48-51.
unnamed averostran (Young, 1935)
Early Cretaceous
Mengyin Formation, Shandong, China
Material- coracoid
Comments- This was initially
described by Young (1935) as Sauropoda gen. et sp. indet., but Wilson
and Upchurch (2009) suggested "it is quite narrow transversely and probably pertains to a large theropod dinosaur."
References- Young, 1935. Dinosaurian remains from Mengyin, Shantung. Bulletin of the Geological Society of China. 15, 519-533.
Wilson and Upchurch, 2009. Redescription and reassessment of Euhelopus zdanskyi (Dinosauria: Sauropoda) from the Early Cretaceous of China. Journal of Systematic Palaeontology. 7, 199-239.
unnamed Averostra (Cheng and Pang, 1996)
Late Cretaceous
Huiquanpu Formation, Shanxi, China
Material- many teeth
Comments- Discovered between
1989 and 1994, "dozens of teeth of Theropoda" were initially referred
to Megalosauridae indet. by Pang et al. (1995) and Cheng and Pang
(1996). The former stated they "are similar to cf. Szechuanosaurus campi
Young from the Upper Cretaceous Wangshi Group in Laiyang Basin,
Shandong Province", while Pang and Cheng (2001) referred them to cf. Szechuanosaurus campi. Wu et al. (2020) noted they have much
finer serrations (15.8 mesial and 20.5 distal per
5 mm) than Szechuanosaurus (8.5 mesial and less distal) or Young's Shandong teeth (12-13 per 5 mm), so that "the teeth of the KDLQ
of HDU also cannot be referred to S. campi." They also differ
from non-eutyrannosaur tyrannosauroid Jinbeisaurus from that formation,
which has 16-16.2 serrations per 5 mm on both carinae
References- Pang, Cheng, Yang,
Xie, Zhu and Luo, 1995. The principal characters and discussion on its
ages of dinosaur fauna in Tianzhen, Shanxi, China. Journal
of Hebei College of Geology. 18(supp.), 1-6.
Cheng and Pang,
1996. A new dinosaurian fauna from Tianzhen, Shanxi Province with its
stratigraphical significance. Acta Geoscientia Sinica. 17, 133-139.
Pang, Cheng, Yang, Xie, Zhu and Luo, 1996. The preliminary report on
Late Cretaceous dinosaur fauna expeditions in Tianzhen, Shanxi. Journal
of Hebei College of Geology. 19(3-4), 227-235.
Pang and Cheng, 2001. The Late Cretaceous dinosaur fauna and strata
from Tianzhen, Shanxi and Yangyuan, Hebei, China. In Deng and Wang
(eds.). Proceedings of the Eighth Annual Meeting of the Chinese Society
of Vertebrate Paleontology. China Ocean Press. 75-82.
Wu, Shi, Dong, Carr, Yi and Xu, 2020 (online 2019). A new tyrannosauroid from the
Upper Cretaceous of Shanxi, China. Cretaceous Research. 108, 104357.
unnamed Averostra (Hu, 1964)
Turonian, Late Cretaceous
Maortu, Ulanhsuhi Formation, Inner Mongolia, China
Material- (IVPP coll.; type 1) lateral tooth (~32x17x10 mm)
(IVPP coll.; type 1) mesiolateral tooth (~30x?x8 mm)
(IVPP coll.; type 1) lateral tooth (~21x12x7 mm)
(IVPP coll.; type 1) two partial teeth
(IVPP coll.; type 2) lateral tooth (~21x10x5 mm)
Turonian, Late Cretaceous
Tashuikou, Ulanhsuhi Formation, Inner Mongolia, China
Material- (IVPP V2884.8; lost) lateral tooth (45x22x11 mm)
Comments- Hu (1964) collected
these isolated teeth in 1960 and referred the Maortu specimens to two
morphotypes of Theropoda indet.. Note Hu's table gives "preserved
length", but this includes the root as demonstrated by Figures 13 and
14 so that crown length is given here instead. Type 1 have a
crown height ratio of 175-188%, a crown base ratio of 58-59%, 16
serrations per 5 mm on the distal(?) carina and finer serrations
mesially. The mesial tip is said to lack serrations, but this may
be due to wear. Hu states these "resemable [sic], in general form
and feature, to that of Szechuanosaurus"
but differ in that the latter is said to have "distinct anterior
serrations", but at least syntypes IVPP V235 and V236 are stated in
Young's description to have more fine mesial serrations as well.
Type 2 has a crown height ratio of 210%, a crown base ratio of 50%, and
"distinct serrations are on both anterior and posterior borders."
Hu again says "It is resemable [sic] to those teeth of Szechuanosaurus",
in the Chinese section specifically noting the measurements are similar
to syntype IVPP V239 - 25x12x7 mm vs. 29x12.5x7 mm. The shapes in
side and basal views are also quite similar, but given the time
difference of Kimmeridgian? vs. Turonian, Maortu specimens are highly
unlikely to be Szechuanosaurus.
Based on morphology, age and location both Maortu types are likely to
be carcharodontosaurids, pantyrannosaurs or eudromaeosaurs.
The Tashuikou tooth was associated with Chilantaisaurus
and said by Hu (1964) to be carnosaurian and "most probably belong to
the same species." Benson and Xu (2008) stated "The tooth
referred to C. tashuikouensis
was not found during the course of this study" and placed it in
Theropoda indet.. The crown height ratio is 205%, the crown base
ratio is 50% and "There are distinct serrations on front and back
borders." As in the Maortu specimens, it is likely to be
carcharodontosaurid, pantyrannosaurian or eudromaeosaurian.
References- Hu, 1964. Carnosaurian remains from Alashan, Inner Mongolia.
Vertebrata PalAsiatica. 8, 42-63.
Benson and Xu, 2008. The anatomy and systematic position of the theropod dinosaur
Chilantaisaurus tashuikouensis Hu, 1964 from the Early Cretaceous of
Alanshan, People’s Republic of China. Geological Magazine. 145(6), 778-789.
unnamed averostran (Hone et al., 2010)
Middle Santonian, Late Cretaceous
Majiacun Formation, Henan, China
Material- (XMDFEC V0010) tooth (52x15x9 mm)
Comments- Hone et al. (2010)
refer this to Baryonychinae, though without suggesting any characters
diagnostic for spinosaurids or subgroups. While isolated baryonychine
teeth can lack fluting, be this labiolingually compressed, or have
serrations this large, Kubota et al. (2017) point out that in the
absence of being spinosaurid-like in these ways and lacking enamel
sculpture, there is no reason to refer the tooth to Spinosauridae. It
is reassigned to Averostra.
References- Hone, Xu and Wang. 2010. A probable baryonychine (Theropoda:
Spinosauridae) tooth from the Upper Cretaceous of Henan Province, China. Vertebrata
PalAsiatica. 48, 19-26.
Kubota, Takakuwa and Hasegawa, 2017. Second discovery of a spinosaurid
tooth from the Sebayashi Formation (Lower Cretaceous), Kanna Town,
Gunma Prefecture, Japan. Bulletin of Gunma Museum of Natural History.
21, 1-6.
unnamed Averostra (Young, 1954a, b)
Late Campanian-Early Maastrichtian, , Late Cretaceous
Xigou, Jiangjunding Formation, Wangshi Group, Shandong, China
Material- (IVPP V756) tooth
(IVPP V757) tooth
(IVPP V758) tooth
(IVPP V759) tooth
(IVPP V760) tooth
(IVPP V761) tooth
(IVPP V764) tooth
(IVPP V765) tooth
(IVPP V766) tooth
Late Campanian-Early Maastrichtian, , Late Cretaceous
near Jingangkou Village, Jiangjunding Formation?, Wangshi Group, Shandong, China
(IVPP V783) tooth
(IVPP V784) tooth
(IVPP V785) tooth
Comments-
These specimens were discovered in 1951 and were first reported by
Young (1954) as "many teeth representing at least two forms" of
Theropoda from the "Wangshih series of the Laiyang region", currently
called the Wangshi Group. Young and Sun (1957) referred to them
as "teeth of Laiyang referred to Szechuanosaurus", later calling them
"tentatively referred to Szechuanosaurus" and noting their serration
density is similar to Kalaza dentary IVPP V903. Young (1958)
figured the teeth and described them in detail, most being from Hsikou
(now Xigou) (IVPP V757-761, V764-766; with V756 "from the place not far
from the opening of the valley and therefore probably derived from the
excavated locality") and three others from "some places near the
village Chingkankou" (now Jingangkou). and referred these to cf. Szechuanosaurus campi because they have mesial serrations unlike Prodeinodon and are larger than Chienkosaurus. Wu et al. (2020) note these teeth have finer serrations (12-13 per 5 mm) than Szechuanosaurus
(8.5 per 5 mm) so "cannot be assigned to S. campi of the Upper
Jurassic." Based on their age, these teeth are more likely
tyrannosauroid or dromaeosaurid.
References- Young, 1954a. Fossil reptilian eggs from Laiyang, Shantung, China. Acta Palaeontologica Sinica. 2(4), 371-388.
Young, 1954b. Fossil reptilian eggs from Laiyang, Shantung, China. Scientia Sinica. 3(4), 505-522.
Young and Sun, 1957. Note on a fragmentary carnosaurian mandible
from Turfan, Sinkiang. Vertebrata PalAsiatica. 1(2), 2027-2036.
Young, 1958. The dinosaurian remains of Liayang, Shantung.
Palaeontologia Sincia. 142(16), 1-138.
Wu, Shi, Dong, Carr, Yi and Xu, 2020 (online 2019). A new tyrannosauroid from the
Upper Cretaceous of Shanxi, China. Cretaceos Research. 108, 104357.
undescribed Averostra (Chow and Rozhdestvensky, 1960)
Middle-Late Campanian, Late Cretaceous
Iren Dabasu Formation, Inner Mongolia, China
Material- (AMNH 6376) phalanx II-1 (AMNH online)
(AMNH 6556) metatarsal II (AMNH online)
(AMNH 6744) four caudal vertebrae, 8 distal pedal elements (AMNH online)
(AMNH 6756) metatarsal (AMNH online)
(AMNH 6757) limb fragments, metapodials, phalanx, fragments (AMNH online)
(AMNH 21552) femur
(AMNH 21565) elements
(AMNH 21588)
(AMNH 21774) fibula
(AMNH 21775) pedal phalanx ?II-1
(AMNH 21776) four proximal pedal phalanges
(AMNH 21780) four unguals
(AMNH 21782) manual ungual
(AMNH 21784) four caudal vertebrae
(AMNH 30245) two metatarsal II or IV shafts (AMNH online)
(AMNH 30247) posterior dorsal rib fragment (AMNH online)
(AMNH 30248) proximal anterior rib (AMNH online)
(AMNH 30249) partial coracoid (AMNH online)
(AMNH 30250) distal femur (AMNH online)
(AMNH 30251) proximal femur (AMNH online)
(AMNH 30252) distal femur (AMNH online)
(AMNH 30253) proximal femur (AMNH online)
(AMNH 30254) distal femur (AMNH online)
(AMNH 30255) astragalus (AMNH online)
(AMNH 30256) proximal tibia (AMNH online)
(AMNH 30257) proximal femur (AMNH online)
(AMNH 30258) distal tibia (AMNH online)
(AMNH 30259) proximal metatarsal (AMNH online)
(AMNH 30260) proximal metatarsal (AMNH online)
(AMNH 30298) acetabular fragment (AMNH online)
(AMNH 30299) proximal ischium (AMNH online)
(AMNH 30360) metatarsal III shaft (AMNH online)
(AMNH 80277) distal humerus (AMNH online)
(IVPP or PIN coll.) (small) three partial skeletons (Chow and Rozhdestvensky, 1960)
(IVPP and PIN coll.) <400 specimens (Currie and Eberth, 1993)
Comments- The AMNH specimens
listed here are from the online catalogue, which generally lacks
identification for ranks between order and family so that Theropoda
indet. material is listed as Saurischia. Yet none of the
specimens are likely to be sauropods given Gilmore (1933) never
mentioned finding any and to this day only a few elements have been
reported (4 in the Erenhot Dinosaur Museum coll., 7 from the
Sino-Soviet expedition- Currie and Eberth, 1993; Sonidosaurus).
Most would have been found during the April 22 to May 25 1923 Central
Asiatic Expedition, AMNH 6556 on April 30. The online catalog
also specifies AMNH 6756 was discovered in AMNH site 141. Many of
these specimens (AMNH 6556, 30245, 30247-30260, 30298-30299, 30360,
80277) are listed as being from "8 mi. E. of station" which would place
them among Third Asiatic Expedition field sites 140-149, with AMNH 6757
listed as 9 miles east, so perhaps site 149. One exception is
AMNH 6744, stated as being found at Elephant Camp (12 miles NW of the
station) by de Chardin, who was only on the 1930 expedition.
Based on the elements preserved, specimen numbers and locality of "8
mi. E. of station", sevcral specimens (AMNH 30245, 30247-30260,
30298-30299, 30360) may belong to the two(+?) troodontid individuals
noted in the AMNH online catalog represented by specimen numbers AMNH
30261-30297, 30300-30318, 30320-30330 and 30336-30359.
Chow and Rozhdestvensky (1960) noted "three partially complete
skeletons of some small carnosaurian dinosaurs" discovered in the
June-July 1959 Sino-Soviet expedition, perhaps indicating
tyrannosauroids or dromaeosaurids. Currie and Eberth (1993)
stated "A rough tally of Sino-Soviet field identifications shows that
... 'theropods' (including large theropods, small theropods and
segnosaurs, but not ornithomimids) were more common (400 specimens)."
References- Gilmore, 1933. On the dinosaurian fauna of the Iren Dabasu
Formation. Bulletin American Museum of Natural History. 67, 23-78.
Chow and Rozhdestvensky, 1960. Exploration in Inner Mongolia - A
preliminary account of the 1959 field work of the Sino-Soviet
Plaeontological Expedition. Vertebrata PalAsiatica. 4(1), 1-10.
Rozhdestvensky and Chow, 1960. On the work of the Soviet-Chinese
paleontological expedition of the USSR and China Academy of Sciences in
1959. Palaeontological Journal. 1, 142-147.
Rozhdestvensky, 1961a. The field research of the Soviet-Chinese
palaeontological expedition of the USSR and China Academy of Sciences
in 1960. Palaeontological Journal. 1, 170-174.
Rozhdestvensky, 1961b. In Central Asia (Brief results of the two-year
investigations of the Soviet and Chinese palaeontologists). Vestnik
Akademii Nauk SSSR. 8, 85-90.
Dong, Currie and Russell, 1989. The 1988 field program of The Dinosaur Project. Vertebrata PalAsiatica. 27(3), 233-236.
Dong, 1992. Dinosaurian Faunas of China. China Ocean Press. 188 pp.
Currie and Eberth, 1993. Palaeontology, sedimentology and palaeoecology of the
Iren Dabasu Formation (Upper Cretaceous), Inner Mongolia, People’s Republic
of China. Cretaceous Research. 14, 127-144.
Dong, 1993. The field activities of the Sino-Canadian Dinosaur Project
in China, 1987-1990. Canadian Journal of Earth Sciences. 30(10),
1997-2001.
unnamed averostran (Mo and Xu, 2015)
Campanian-Maastrichtian, Late Creyaceous
Nanxiong Group, Jiangxi, China
Material- (NHMG 8500) maxillary tooth (91 x 45.2 x 21 mm)
Comments- Mo and Xu (2015) referred this tooth to Theropoda indet., stating
it was most likely carcharodontosaurid or tyrannosaurid. As it is far more labiolingually
compressed than tyrannosaurids, large and serrated without strong carcharodontosaurine
wrinkles, it seems most likely to be a basal carcharodontosaurid or megaraptoran.
Reference- Mo and Xu, 2015. Large theropod teeth from the Upper Cretaceous
of Jiangxi, southern China. Vertebrata PalAsiatica. 53(1), 63-72.
undescribed possible Averostra (Li and Gao, 2007)
Barremian-Albian, Early Cretaceous
Sinuiju Series, North Korea
Comments- Li and Gao (2007) reported theropods from the Sinuijiu Series,
while Gao et al. (2009) stated "possible theropod dinosaurs" had been
discovered there.
References- Li and Gao, 2007. Lower Cretaceous vertebrate fauna from
the Sinuiju basin, North Korea as evidence of geographic extension of the Jehol
biota into the Korean peninsula. Journal of Vertebrate Paleontology. 27(3),
106A.
Gao, Li, Wei, Pak and Pak, 2009. Early Cretaceous birds and pterosaurs from
the Sinuiju series, and geographic extension of the Jehol biota into the Korean
peninsula. Journal of the Paleontological Society of Korea. 25(1), 57-61.
undescribed Averostra (Manabe and Barrett, 2000)
Valanginian-Hauterivian, Early Cretaceous
Kuwajima Formation of the Tetori Group, Japan
Material- (SBEI-156) incomplete tooth (Matsuoka et al., 2002)
(SBEI-170) tooth (Matsuoka et al., 2002)
(SBEI-171) tooth (Matsuoka et al., 2002)
(SBEI-576) tooth (Matsuoka et al., 2002)
(SBEI-814) incomplete tooth (Matsuoka et al., 2002)
Comments- These small teeth were announced as velociraptorine in Barrett
and Manabe's (2000) abstract, but Matsuoka et al. (2002) note none have high
DSDIs, so merely refer to them as Theropod Type B.
References- Barrett and Manabe, 2000. The dinosaur fauna from the Earliest
Cretaceous Tetori Group of Central Honshu, Japan. Journal of Vertebrate Paleontology.
20(3), 28A-29A.
Matsuoka, Kusuhashi, Takada and Setoguchi, 2002. A clue to the Neocomian vertebrate
fauna: Initial results from the Kuwajima 'Kaseki-kabe' (Tetori Group) in Shiramine,
Ishikawa, central Japan. Memoirs of the Faculty of Science, Kyoto University,
Series of Geology and Mineralogy. 59(1), 33-45.
undescribed averostran (Azuma, Xu, Shibata,
Kawabe, Miyata and Imai, 2016)
Middle-Late Aptian, Early Cretaceous
Kitadani Dinosaur Quarry, Kitadani Formation of the Akaiwa Subgroup of the Tetori Group, Japan
Material- (FPDM uncatalogued; formerly part of FPDM-V-8461) distal caudal
vertebra
Comments- This was originally
assigned to the holotype of "Fukuivenator" by Azuma et
al. (2016), who identified thirty caudals before Hattori et al. (2021)
identified the fourth and twenty-ninth caudals while listing a distal
caudal as one of three "Withdrawn elements" without further
comment. Hattori (pers. comm., 3-4-2022) indicates this caudal is
theropodan but was incorrectly associated with the holotype block due
to similarities in its field number, but that it currently has no
specimen number.
References-
Azuma, Xu, Shibata, Kawabe, Miyata and Imai, 2016. A bizarre theropod from the
Early Cretaceous of Japan highlighting mosaic evolution among coelurosaurians.
Scientific Reports. 6, 20478.
Hattori, Kawabe, Imai, Shibata, Miyata, Xu and Azuma, 2021. Osteology of Fukuivenator paradoxus:
A bizarre maniraptoran theropod from the Early Cretaceous of Fukui,
Japan. Memoir of the Fukui Prefectural Dinosaur Museum. 20, 1-82.
undescribed Averostra (Ikegami, 2010)
Coniacian-Santonian, Late Cretaceous
Upper Formation of Mifune Group, Japan
Comments- Ikegami (2010) report two bonebeds, one containing isolated theropod
elements, and another preserving three theropod lineages.
Reference- Ikegami, 2010. Taphonomy and sedimentology of a bonebed from
the Upper Cretaceous Mifune Group in Kyushu, Japan. Journal of Vertebrate Paleontology.
Program and Abstracts 2010, 109A-110A.
undescribed Averostra (Hirayama, Takisawa, Sasaki, Sonoda, Yoshida,
Takekawa, Mitsuzuka, Kobayashi, Tsuihiji and Tsutsumi, 2015)
Early Santonian, Late Cretaceous
Tamagawa Formation of the Kuji Group, Japan
Material- limb elements
Reference- Hirayama, Takisawa, Sasaki, Sonoda, Yoshida, Takekawa, Mitsuzuka,
Kobayashi, Tsuihiji and Tsutsumi, 2015. Terrestrial vertebrates from the Late
Cretaceous (Santonian) of Iwate prefecture, eastern Japan. Journal of Vertebrate
Paleontology. Program and Abstracts 2015, 143-144.
unnamed Averostra (Tamara et al., 1991)
Late Cenomanian-Early Turonian, Late Cretaceous
Jobu Formation of Mifune Group, Japan
Material- (Kitakyusyu Museum of Natural History coll.) tooth
(Kitakyusyu Museum of Natural History coll.) two dorsal neural arches, fibula
(Mifune Board of Education laboratory coll.) four teeth, incomplete tibia, fibula,
distal metatarsal II, distal metatarsal III
Comments- These remains were stated to be like Allosaurus by Tamara
et al. (1991), though Chure (2000) thought they were different enough to be
excluded from Allosauridae. A femur was associated with the dorsal neural arches
and fibula, but Chure correctly notes it resembles pterosaurs more. The teeth
differ from Allosaurus in being taller with straight posterior margins.
The neural arches differ in having lower and thinner neural spines. The tibia
and fibulae are too damaged to be useful. The distal metatarsals differ in being
transversely wider, with larger rounder flexor depressions and less cranioproximally
extensive distal articular surfaces.
References- Tamara, Okazaki and Ikegami, 1991. Occurence of carnosaurian
and herbivorous dinosaurs from upper formation of Mifune Group, Japan, Memiors
of the Faculty of Education, Kumamoto University. 40, 31-45.
Chure, Manabe, Tanimoto and Tomida, 1999. An unusual theropod tooth from the
Mifune Group (Late Cenomanian to Early Turonian), Kumamoto, Japan. In Tomida,
Rich, and Vickers-Rich (eds.). Proceedings of the Second Gondwanan Dinosaur
Symposium. National Science Museum (Tokyo) Monographs. 15, 291-296.
Chure, 2000. A new species of Allosaurus from the Morrison Formation
of Dinosaur National Monument (Utah-Colorado) and a revision of the theropod
family Allosauridae. PhD thesis. Columbia University. 964 pp.
undescribed Averostra (Buffetaut and Suteethorn, 1998)
?Oxfordian-Early Valanginian, ?Late Jurassic-Early Cretaceous
Phu Kradung Formation, Thailand
Material- (?SM coll.) tooth fragments (Buffetaut and Suteethorn, 2007)
(SM coll.) teeth (Suteethorn et al., 2013)
teeth, astragalus (Buffetaut and Suteethorn, 1998a, b)
(small) material (Chanthasit, 2011)
Comments- Buffetaut and
Suteethorn (1998a, b) report "indeterminate theropods (isolated teeth,
an astragalus)" and "teeth and an astragalus of an indeterminate
theropod" respectively without specifying the locality.
Buffetaut and Suteethorn (2007) reported "fragments of theropod teeth" from the Kham Phok locality.
Chanthasit (2011) and Chanthasit et al. (2015) report on small theropod remains from the Phu Noi locality.
Suteethorn et al. (2013) reported "theropod teeth" found in 2004 from the Phu Dan Ma locality.
References-
Buffetaut and Suteethorn, 1998a. Jurassic dinosaurs from Thailand. EWVP 3. 81.
Buffetaut and Suteethorn, 1998b. Early Cretaceous dinosaurs from Thailand and
their bearing on the early evolution and biogeographical history of some groups
of Cretaceous dinosaurs. In Lucas, Kirkland and Estep (eds.). Lower and Middle
Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History Bulletin.
14, 205-210.
Buffetaut and Suteethorn, 2007. A sinraptorid theropod (Dinosauria:
Saurischia) from the Phu Kradung Formation of northeastern Thailand. Bulletin
de la Societe Geologique de France. 178, 497-502.
Chanthasit, 2011. New theropod remains from the Phu Kradung Formation
of Kalasin Province and a review of Late Jurassic theropod record in
Thailand. World Conference on Paleontology and Stratigraphy. Program
and Abstracts, 34.
Suteethorn, Le Loeuff, Buffetaut, Suteethorn and Wongko, 2013. First
evidence of a mamenchisaurid dinosaur from the Upper Jurassic-Lower
Cretaceous Phu Kradung Formation of Thailand. Acta Palaeontologica
Polonica. 58(3), 459-469.
Chanthasit, Suteethorn and Suteethorn, 2015. Dinosaur assemblage from
Phu Noi fossil site in Kalasin Province, northeastern Thailand. 2nd
International Symposium on Asian Dinosaurs 2015 Bangkok, p. 23.
undescribed Averostra (Buffetaut, 1983)
Late Barremian?, Early Cretaceous
Ban Ao Kalang, Sao Khua Formation?, Thailand
(PRC MR45) incomplete tooth (14x9x5 mm) (Cuny, Laojumpon, cheychiw and Lauprasert, 2010)
Late Barremian, Early Cretaceous
Kalasin 11, Sao Khua Formation, Thailand
(SM K11-0168) tooth (~38x~16x? mm) (Suteethorn, Martin, Buffetaut, Triamwichanon and Chaimanee, 1995)
Late Barremian, Early Cretaceous
Khok Doo 1, Sao Khua Formation, Thailand
(SM-KD-1 coll.) teeth (Martin, Suteethorn and Buffeaut, 1999)
Late Barremian, Early Cretaceous
Phu Din Daeng, Sao Khua Formation, Thailand
Material- (PRC148) incomplete tooth (~67x~27x? mm) (Tong, Buffetaut, Suteethorn, Suteethorn,
Cuny, Cavin, Deesri, Martin, Wongko, Naksri and Claude, 2019)
(PRC coll.) several teeth, incomplete caudal vertebra (Tong, Buffetaut, Suteethorn, Suteethorn,
Cuny, Cavin, Deesri, Martin, Wongko, Naksri and Claude, 2019)
Late Barremian, Early Cretaceous
Phu Sung, Sao Khua Formation, Thailand
tooth, manual unguals (Chanthasit, Suteethorn, Naksri, Tong, Wongko and Sonoda, 2019)
Late Barremian, Early Cretaceous
Phu Wiang 1A, Sao Khua Formation, Thailand
(SM-TF coll.) teeth (~60x~23x~15.5 mm) (Buffetaut and Ingavat, 1983)
Late Barremian, Early Cretaceous
Phu Wiang 3, Sao Khua Formation, Thailand
(SM-PW-3 coll.) about ten teeth (?x~14x? mm) (Buffetaut and Suteethorn, 1989)
(?SM coll.) teeth (Buffetaut, 1983)
Late Barremian, Early Cretaceous
Phu Wiang 5, Sao Khua Formation, Thailand
(SM-PW-5 coll.) teeth and vertebrae (Martin, Suteethorn and Buffeaut, 1999)
Late Barremian, Early Cretaceous
Phu Wiang 5B, Sao Khua Formation, Thailand
(SM-PW-5B coll.) remains (Martin, Suteethorn and Buffeaut, 1999)
Late Barremian, Early Cretaceous
Phu Wiang 7, Sao Khua Formation, Thailand
(SM-PW-7 coll.) remains (Martin, Suteethorn and Buffeaut, 1999)
Late Barremian, Early Cretaceous
Phu Wiang 9A, Sao Khua Formation, Thailand
(SM-PW-9A coll.) teeth (Martin, Suteethorn and Buffeaut, 1999)
Late Barremian, Early Cretaceous
Phu Wiang 11, Sao Khua Formation, Thailand
(SM-PW-11) teeth (Martin, Suteethorn and Buffeaut, 1999)
Late Barremian, Early Cretaceous
Sakhon Nakhon 1, Sao Khua Formation, Thailand
(SM-SN-1 coll.) teeth (Martin, Suteethorn and Buffeaut, 1999)
Comments- Buffetaut (1983)
reported "additional remains of ... theropods" found in 1981 "at Phu
Wieng", which was clarified to be "Theropoda indet. : teeth" from Phu
Wiang 3 by Martin et al. (1999).
Buffetaut and Ingavat (1983) reported teeth "of the usual carnosaur
type : strongly compressed blade-like teeth with smooth enamel and
serrated cutting edges" in addition to what would be described as Siamosaurus
by Buffetaut and Ingavat (1986) from the Phu Wiang 1A locality.
The latter reference figures one such tooth as "a "normal" carnosaur
from the same locality."
Buffetaut and Suteethorn (1989) noted about ten teeth found in 1987 associated with what would be described as the holotype of Phuwiangosaurus
in 1994, at Phu Wiang 3. These "exhibit the characteristic
morphology of carnosaur teeth: the crown is strongly compressed
laterally, recurved, with sharp, finely serrated cutting edges" and
one tooth is photographed.
Suteethorn et al. (1995) reported a tooth associated with a Phuwiangosaurus
skeleton (SM K11-0001 to SM K11-0167) from Phu Wiang 11, which was
figured when that specimen was described by Suteethorn et al.
(2009).
Martin et al. (1999) listed several occurances of Theropoda indet. from various localities in the Sao Khua Formation.
Chanthasit et al. (2019) report "a large isolated tooth with serration
and manual claws of indeterminate theropods" from the Phu Sung locality.
Tong et al. (2019) reported a "large caudal vertebra lacking the neural
spine" and "several blade-shaped, laterally compressed teeth with
serrated carinae" from Phu Din Daeng, one of the latter which was
figured.
While it's been suggested some of these teeth belong to Siamotyrannus (e.g. Buffetaut and Suteethorn, 1998), Phuwiangvenator and Vayuraptor have since been described from the formation and likely had similar teeth.
References- Buffetaut, 1983. Mesozoic vertebrates from Thailand: A review. Acta Palaeontologica Polonica. 28(1-2), 43-53.
Buffetaut and Ingavat, 1983. Vertebrates from the continental Jurassic of Thailand. CCOP Technical Bulletin. 16, 68-75.
Buffetaut and Ingavat, 1986. Unusual theropod dinosaur teeth from the Upper
Jurassic of Phu Wiang, northeastern Thailand. Revue de Pal�obiologie.
5, 217-220.
Buffetaut and Suteethorn, 1989. A sauropod skeleton associated with
theropod teeth in the Upper Jurassic of Thailand: Remarks on the
taphonomic and palaeoecological significance of such associations.
Palaeogeography, Palaeoclimatology, Palaeoecology. 73, 77-83.
Suteethorn, Martin, Buffetaut, Triamwichanon and Chaimanee, 1995. A new
dinosaur locality in the Lower Cretaceous of northeastern Thailand.
Comptes Rendus de l’Academie des Sciences, Serie IIa. 321, 1041-1047.
Buffetaut and Suteethorn, 1998. Early Cretaceous dinosaurs from Thailand and
their bearing on the early evolution and biogeographical history of some groups
of Cretaceous dinosaurs. In Lucas, Kirkland and Estep (eds.). Lower and Middle
Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History Bulletin.
14, 205-210.
Martin, Suteethorn and Buffeaut, 1999. Description of the type and referred material of Phuwiangosaurus sirindhornae Martin, Buffetaut and Suteethorn, 1994, a sauropod from the Lower Cretaceous of Thailand. Oryctos. 2, 39-91.
Suteethorn, Le Loeuff, Buffetaut, Suteethorn, Talubmook and Chonglakmani, 2009. A new skeleton of Phuwiangosaurus sirindhornae
(Dinosauria, Sauropoda) from northeastern Thailand. In Buffetaut, Cuny,
Le Loeuff and Suteethorn (eds.). Late Palaeozoic and Mesozoic
Ecosystems of Southeast Asia. Geological Society London Special
Publication. 315, 189-215.
Cuny, Laojumpon, Cheychiw and Lauprasert, 2010. Fossil vertebrate
remains from Kut Island (Gulf of Thailand, Early Cretaceous).
Cretaceous Research. 31, 415-423.
Chanthasit, Suteethorn, Naksri, Tong, Wongko and Sonoda, 2019. New
vertebrate fossil site from the Early Cretaceous Sao Khua Formation,
Sakon Nakhon Province, northeastern Thailand. Open Journal of Geology.
9, 619-622.
Tong, Buffetaut, Suteethorn, Suteethorn, Cuny, Cavin, Deesri, Martin,
Wongko, Naksri and Claude, 2019. Phu Din Daeng, a new Early Cretaceous
vertebrate locality on the Khorat Plateau, NE Thailand. Annales de
Pal�ontologie. 105(3), 223-237.
undescribed Averostra (Buffetaut, 1983)
Aptian, Early Cretaceous
Khok Kruat Formation, Thailand
Material- quadrate, tooth (Buffetaut, 1983)
(multiple taxa and localities) teeth (Buffetaut, Suteethorn, Le Loeuff, Khansubha, Tong and Wongko, 2005)
Comments- Buffetaut (1983)
initially reported "a theropod tooth and a quadrate which probably also
belongs to a theropod (P. Taquet, pers. comm.)" discovered in
1978. Buffetaut and Ingavat (1986) stated "a few incomplete bones
can be referred to a theropod dinosaur", while Buffetaut and Suteethorn
(1992) said "scanty remains of theropod dinosaurs (including teeth and
fragmentary bones)" were known. These reports may all refer back
to Buffetaut's tooth and quadrate and are from the Ban Khok Kruat
locality.
Buffetaut et al. (2005) wrote "isolated teeth are relatively common at
several localities. At Khok Pa Suam (Ubon Ratchathani Province), a
number of blade-like teeth, with serrations on both margins, have been
collected. Differences in size and morphology strongly suggest that
several taxa are present, with at least a large form and a smaller one."
References- Buffetaut, 1983. Mesozoic vertebrates from Thailand: A review. Acta Palaeontologica Polonica. 28(1-2), 43-53.
Buffetaut and Ingavat, 1986. The succession of vertebrate faunas in the
continental Mesozoic of Thailand. Bulletin of the Geological Society of
Malaysia. 19, 167-172.
Buffetaut and Suteethorn, 1992. A new species of the ornithischian
dinosaur Psittacosaurus from the Early Cretaceous of Thailand.
Palaeontology. 35(4), 801-812.
Buffetaut, Suteethorn, Le Loeuff, Khansubha, Tong and Wongko, 2005. The dinosaur
fauna from the Khok Kruat Formation (Early Cretaceous) of Thailand. In Wannakao,
Youngme, Srisuk and Lertsirivorakul (eds.). Proceedings of the International
Conference on Geology, Geotechnology and Mineral Resources of Indochina. 575-581.
undescribed Averostra (Glut, 1982)
Middle Jurassic
Paikasigudem, Kota Formation, India
Material-
(DUGF/J19-23, 25-37, 39, 45-46, 52-56, 60-65, 68) thirty-three teeth (Prasad and Manhas, 2002)
Middle Jurassic
Yamanpalli, Kota Formation, India
(GSI coll.) elements including braincase (Glut, 1982)
Comments- Glut (1982) reported
"Two "carnosaurs," one of which includes a preserved brain cavity, from
the Lower Jurassic Kota Formation near Yamanpalli, Andra Pradesh,
India, excavated by Yadagiri in 1979." Olshevsky (1991) says one
of these is Dandakosaurus, described by Yadagiri in 1982, which does not preserve a braincase. The other, Olshevsky lists as "Genus:
[To be described from the Kota Formation of India; Yadagiri, 1979]"
under Megalosauridae. Despite a thorough search of the literature
involving Olshevsky, Ford and Molnar, no Yadagiri 1979 work referencing
theropods has been located (note no theropoids are mentioned in "Yadagiri, 1979. Observations on Kota Formation of Pranhita Godavari
valley, south India. Geological Survey of India, Miscellaneous
Publications. 45, 73-79."). Glut also mentions "two
sauropod skulls, generically different because of the position of the
frontals, parietals, supratemporal fossae and paroccipitals, discovered
by Yadagiri in 1979" and "a stegosaur, discovered by Yadagiri in 1979",
which may indicate these are all personal communications or unpublished
presentations from Yadagiri as opposed to anything published by
Yadagiri in 1979. In this case Olshevsky was mistaken to present
it as a reference, but it would explain its absence from the literature
otherwise.
Prasad and Manhas (2002) note undescribed "theropod and ornithischian
dinosaur teeth" from the Paikasigudem microvertebrate site.
References- (?) Yadagiri, 1979. [title] [journal] [volume, pp]
Glut, 1982. The New Dinosaur Dictionary. Citadel Press. 288 pp.
Olshevsky, 1991. A revision of the parainfraclass Archosauria Cope, 1869, excluding
the advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.
Prasad and Manhas, 2002. Triconodont mammals from the Jurassic Kota Formation of India. Geodiversitas. 24(2), 445-464.
unnamed Averostra (Lasseron, 2020)
Early Bathonian, Middle Jurassic
GEA 3, Guelb el Ahmar,
Anoual Formation, Morocco
(MNHN GEA3-23; Theropoda gen. et sp. indet. morphotype III) lateral tooth (3.18x2.15x.99 mm) (Lasseron, 2020)
(MNHN GEA3-24; Coelurosauria gen. et sp. indet. morphotype II) lateral tooth (16.76x5.42x6.20 mm) (Lasseron, 2020)
Early Bathonian, Middle Jurassic
GEA 6, Guelb el Ahmar,
Anoual Formation, Morocco
(MNHN GEA6-4; Theropoda gen. et sp. indet. morphotype III) lateral tooth (4.04x2.13x1.52 mm) (Lasseron, 2020)
Early Bathonian, Middle Jurassic
GEA 7, Guelb el Ahmar,
Anoual Formation, Morocco
(MNHN GEA7-16; Coelurosauria gen. et sp. indet. morphotype II) lateral tooth (8.31x4.13x4.11 mm) (Lasseron, 2020)
Early Bathonian, Middle Jurassic
Guelb el Ahmar,
Anoual Formation, Morocco
(MNHN GEA coll.) 46 teeth (Lasseron, 2020)
Comments-
Discovered in 2010, Haddoumi et al. (2015) stated "A few nearly
complete teeth showing a labiolingually compressed crown and serrated
mesial and distal carinae can be assigned to a small, indeterminate
theropod." The figured tooth (MNHN GEA 2-5) was placed in
Lasseron's
(2020) 'Theropoda gen. et sp. indet. morphotype I' in that work, which
is listed under Coelurosauria indet. here. No other theropod
teeth from GEA 2 were mentioned by Lasseron, meaning the others were
probably fragments among the 46 additional dental specimens catalogued
by Lasseron. The FSAC Theropoda
gen. et sp. indet. morphotype III and Coelurosauria gen. et sp. indet.
morphotype II of Lasseron have both mesial and distal serrations
References- Haddoumi, Allain, Meslouh, Metais, Monbaron, Pons, Rage, Vullo,
Zouhri and Gheerbrant, 2016 (online 2015). Guelb el Ahmar (Bathonian, Anoual Syncline, eastern
Morocco): First continental flora and fauna including mammals from the Middle
Jurassic of Africa. Gondwana Research. 29(1), 290-319.
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 possible averostran (Lapparent, 1955)
Early Bathonian, Middle Jurassic
Oued Botane, El Mers Formation, Morocco
Material- (MNHN ELM coll.; syntype of Megalosaurus mersensis) two or three incomplete teeth (<50 mm)
Comments- Excavated in 1941, these were made a syntype of Lapparent's (1955) new theropod species Megalosaurus mersensis,
along with a partial vertebral column now considered a teleosauroid
(Chabli, 1986) and a dorsal vertebra supposedly similar to the latter
specimen and so also placed in Teleosauroidea here. Unfortunately
the teeth were not illustrated and were only described as "three teeth,
unfortunately very incomplete. Their flattened form, in the slightly
arched blade of a saber, makes them belong to Megalosaurus. The enamel is ornamented with fine, regular longitudinal striations, as in Megalosaurus bucklandi,
but the serrations on the edges are not visible. Their size does not
exceed 5 cm in length." The teeth are here provisionally retained
as Averostra indet., as teleosaurid teeth are at best slightly
compressed labiolingually and Lapparent describes teleosaurid teeth in
the same publication (as the "Steneosaurus
from El Mers") so would be expected to know the difference (although he
didn't for the vertebrae, so this opinion is not made
confidently). As Lapparent compared the fine longitudinal
striations to Megalosaurus bucklandii,
they are unlikely to be strong enough to be fluting, and this leaves us
with no data to distinguish them from e.g. Jurassic ceratosaurs,
megalosauroids or carnosaurs. Note Lapparent's "List of principal
vertebrate localities in the Jurassic of El Mers" gives the number of
teeth as two, and while they were found two years after the vertebral
column, his statement "The presence of theropod carnivores at El Mers
was revealed to us initially by three teeth" could easily mean the
vertebrae were not (incorrectly) recognized as theropod until after the
teeth were discovered. The specimen numbers are assumed to be
MNHN ELM based on "Cetiosaurus" mogrebiensis material described by Lapparent in the same publication and redescribed by Lang (2008).
References- Lapparent, 1955. �tude pal�ontologique des vert�br�s du Jurassique d'El
Mers (Moyen Atlas). Notes et M�moires du Service G�ologique du Maroc. 124, 1-36.
Chabli, 1986. Donn�es nouvelles sur un "Dinosaurien" Jurassique Moyen du Maroc: Megalosaurus mersensis
Lapparent 1955, et sur les Megalosaurides en g�n�ral. In Taquet and
Sudre (eds.). Les Dinosaures de la Chine � la France. Mus�e d'Histoire
Naturelle de Toulouse. 66-72.
Lang, 2008. Les cetiosaures (Dinosauria, Sauropoda) et les sauropodes du Jurassique
moyen: Revision systematique, nouvelles decouvertes et implications phylogenetiques.
Doctoral Thesis. Museum National d’Histoire Naturelle. 638 pp.
undescribed Averostra (Monbaron, Russell and Taquet, 1999)
Bathonian, Middle Jurassic
Wawmda, Gres de Guettioua Formation, Morocco
Material- (Mus�e des sciences de la Terre de Rabat coll.) teeth
Comments- Excavated in Autumn 1980, Monbaron et al. (1999) noted "theropod teeth were found in association with the" holotype of Atlasaurus.
Although initially identified as the Tilougguit Formation by Monbaron
et al., Allain and Aquesbi (2008) later placed it in the overlying Gres
de Guettioua Formation.
References- Monbaron, Russell and Taquet, 1999. Atlasaurus imelakei,
n. g., n. sp., a brachiosaurid-like sauropod from the Middle Jurassic
of Morocco. Comptes Rendus de l’Acad�mie des Sciences. 329, 519-526.
Allain and Aquesbi, 2008. Anatomy and phylogenetic relationships of Tazoudasaurus naimi (Dinosauria, Sauropoda) from the late Early Jurassic of Morocco. Geodiversitas. 30(2), 345-424.
unnamed Averostra (Knoll and Ruiz-Omenaca, 2009)
Berriasian, Early Cretaceous
KM 1983, Ksar Metlili, Ksar Metlili Formation, Morocco
Material- (MNHN SA 2004/2A; lost) tooth fragment (Knoll and Ruiz-Omenaca, 2009)
(MNHN SA 2004/2C; lost) tooth fragment (Knoll and Ruiz-Omenaca, 2009)
(MNHN SA 2004/2D; lost) tooth fragment (Knoll and Ruiz-Omenaca, 2009)
(MNHN SA 2004/2E; lost) tooth fragment (Knoll and Ruiz-Omenaca, 2009)
(MNHN SA 2004/2F; lost) tooth fragment (Knoll and Ruiz-Omenaca, 2009)
(MNHN SA 2004/2G; lost) partial tooth (Knoll and Ruiz-Omenaca, 2009)
(MNHN SA 2004/3B; lost) tooth fragment (4.90x?x1.60 mm) (Knoll and Ruiz-Omenaca, 2009)
(MNHN SA 2004/4A; lost) tooth fragment (Knoll and Ruiz-Omenaca, 2009)
(MNHN SA 2004/5C; lost) partial tooth (Knoll and Ruiz-Omenaca, 2009)
(MNHN SA 2004/5D; lost) tooth fragment (Knoll and Ruiz-Omenaca, 2009)
(MNHN SA mcm 167; lost) tooth fragment (4.00x?x1.16 mm) (Knoll and Ruiz-Omenaca, 2009)
Beriassian, Early Cretaceous
KM-A1, Ksar Metlili, Ksar Metlili Formation, Morocco
(FSAC-KM-A1-9) incomplete manual ungual (Lasseron, 2020)
(FSAC-KM-A1-11) incomplete manual ungual, pedal ungual, three unguals (Lasseron, 2020;
Lasseron, Allain, Gheerbrant, Haddoumi, Jalil, M�tais, Rage, Vullo and
Zouhri, 2020)
(FSAC-KM-A1-13) thirteen teeth (Lasseron, Allain, Gheerbrant, Haddoumi, Jalil, M�tais, Rage, Vullo and
Zouhri, 2020)
(FSAC-KM-A1-55) incomplete manual ungual (Lasseron, 2020)
(FSAC-KM-A1-56) pedal ungual (~56 mm) (Lasseron, 2020)
Beriassian, Early Cretaceous
KM-A2, Ksar Metlili, Ksar Metlili Formation, Morocco
(FSAC-KM-A2-10) two teeth (Lasseron, Allain, Gheerbrant, Haddoumi, Jalil, M�tais, Rage, Vullo and
Zouhri, 2020)
Beriassian, Early Cretaceous
KM-B', Ksar Metlili, Ksar Metlili Formation, Morocco
(FSAC-KM-B'-21) incomplete manual ungual (Lasseron, 2020)
(FSAC-KM-B'-23) ten teeth (Lasseron, Allain, Gheerbrant, Haddoumi, Jalil, M�tais, Rage, Vullo and
Zouhri, 2020)
(FSAC-KM-B'-24) four teeth (Lasseron, Allain, Gheerbrant, Haddoumi, Jalil, M�tais, Rage, Vullo and
Zouhri, 2020)
(FSAC-KM-B'-25; Coelurosauria gen. et sp. indet. morphotype II) anterior tooth (1.51x.97x.80 mm) (Lasseron, 2020)
(FSAC-KM-B'-26; Theropoda gen. et sp. indet. morphotype III) lateral tooth (.89x.81x.54 mm) (Lasseron, 2020)
(FSAC-KM-B'-26 [3]; Coelurosauria gen. et sp. indet. morphotype III) lateral tooth (3.18x1.65x.83 mm) (Lasseron, 2020)
(FSAC-KM-B'-94) pedal ungual (Lasseron, 2020)
Beriassian, Early Cretaceous
KM-C, Ksar Metlili, Ksar Metlili Formation, Morocco
(FSAC-KM-C-1) manual(?) ungual (Lasseron, 2020)
Beriassian, Early Cretaceous
KM-D1, Ksar Metlili, Ksar Metlili Formation, Morocco
(FSAC-KM-D1-9) eighteen teeth (Lasseron, Allain, Gheerbrant, Haddoumi, Jalil, M�tais, Rage, Vullo and
Zouhri, 2020)
Beriassian, Early Cretaceous
KM-D2, Ksar Metlili, Ksar Metlili Formation, Morocco
(FSAC-KM-D2-9) manual(?) ungual (Lasseron, 2020)
(FSAC-KM-D2-10) twelve teeth (Lasseron, Allain, Gheerbrant, Haddoumi, Jalil, M�tais, Rage, Vullo and
Zouhri, 2020)
Comments- The MNHN specimens were collected in 1983, 1986 and 1999, while the
FSAC specimens were collected in 2010, 2015 and 2018 (Lasseron, 2020).
Lasseron (2020) noted the "dinosaur remains (Knoll, 2000; Knoll & Ruiz-Ome�aca, 2009) ... were taken out of the MNHN and lost" (translated). Perhaps
first mentioned by Evans and Sigogneau-Russell (1997) as "small ...
theropod dinosaurs", Sigogneau-Russell et al. (1998) noted "We have
been able to distinguish 11 dinosauromorph tooth varieties (8
theropod") based on very small teeth (FABL 1-3 mm)" which were later
described in detail by Knoll and Ruiz-Omenaca (2009). The MNHN
specimens are referred to Theropoda indet. by Knoll and
Ruiz-Omenaca (2009) and are mostly fragments that could not be
evaluated for their tooth morphotypes, except MNHN SA 2004/5C which is
the second largest tooth in the sample (preserved length 11 mm) and has
a low DSDI (.77-.91) unlike their other morphotypes which lack mesial
serrations or have a high DSDI. The FSAC Theropoda gen. et sp.
indet. morphotype III of Lasseron has both mesial and distal
serrations, as do the Coelurosauria gen. et sp. indet. morphotypes II
and III. Lasseron et al. lists all four teeth under FSAC-KM-B'-26 as Dromaeosauridae.
Lasseron refers all the manual and pedal unguals from Ksar Metlili to Coelurosauria because "The
presence of a single groove per side identifies these claws as
belonging to coelurosaurs, and not to abelisauroids, which have two
grooves on each side", but some ceratosaurs like Limusaurus
(IVPP V15304, V20096 and V15923 in part) have single grooves and they
could also be juvenile e.g. spinosaurids or carcharodontosaurids
instead of coelurosaurs, so they are all assigned to Averostra
here. Unequivocal manual unguals show "strong mediolateral compression ... the proximal section of the claws indicates that it was dorsoventrally high. Their cross section is oval. They are strongly curved ventrally." the "others
(KM-C-1, KMD2-9) are very poorly preserved, but their strong lateral
compression suggests that they are claws from the hands." He
notes FSCA-KM-A1-55 differs from most of the manual unguals in that "the
side grooves are less extended proximally, but are underlined by a
small bony plateau, which protrudes medially and laterally from the
proximal contour of the claw." Pedal unguals "have a dorsoventrally larger articular surface, slightly concave and smooth. Their
transverse section is subtriangular and their ventral surface is very
slightly curved, almost flat." Lasseron et al. listed FSAC-KM-A1-9 and FSAC-KM-A1-55 as "Claw (pedal?)"
References-
Evans and Sigogneau-Russell, 1997. New sphenodontians (Diapsida:
Lepidosauria: Rhynchocephalia) from the Early Cretaceous of North
Africa. Journal of Vertebrate Paleontology. 17(1), 45‑51.
Sigogneau-Russell, Evans, Levine and Russell, 1998. The Early
Cretaceous microvertebrate locality of Anoual, Morocco: A glimpse at
the small vertebrate assemblages of Africa. In Lucas, Kirkland and
Estep (eds.). Lower and Middle Cretaceous Terrestrial Ecosystems. New
Mexico Museum of Natural History and Science. New Mexico Museum of
Natural Histroy and Science Bulletin. 14, 177‑182.
Knoll and Ruiz-Omenaca, 2009. Theropod teeth from the basalmost
Cretaceous of Anoual (Morocco) and their palaeobiogeographical significance.
Geological Magazine. 146(4), 602-616.
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.
Lasseron, Allain, Gheerbrant, Haddoumi, Jalil, M�tais, Rage, Vullo and
Zouhri, 2020 (online 2019). New data on the microvertebrate fauna from
the Upper Jurassic or lowest Cretaceous of Ksar Metlili (Anoual
Syncline, eastern Morocco). Geological Magazine. 157, 367‑392.
unnamed Averostra (Lavocat, 1954)
Cenomanian, Late Cretaceous
Kem Kem beds, Morocco
Material- (CMN 41770; bone taxon N) metatarsal IV (430 mm) (Russell,
1996)
(CMN 41806; bone taxon E) coracoid fragment (Russell, 1996)
(CMN 41863; bone taxon D) partial distal caudal vertebra (86 mm) (Russell, 1996)
(CMN 50382; bone taxon M) (3.78 kg, juvenile) femur (118 mm)
(CMN 50797; bone taxon D) incomplete distal caudal vertebra (65 mm) (Russell,
1996)
(CMN 50807; bone taxon A) (adult) frontals (50 mm), parietals (Russell, 1996)
(CMN 50808; bone taxon A) (adult) frontals (65 mm) (Russell, 1996)
?(CMN 50810; bone taxon B) (~4 m) (subadult) axis (45 mm) (Russell, 1996)
(CMN 50826; bone taxon P) partial pedal ungual (Russell, 1996)
(CMN 50830; bone taxon O) metatarsal V (187 mm) (Russell, 1996)
(CMN 50839a-e; bone taxon K) manual ungual fragments (~20-50 mm) (Russell, 1996)
(CMN 50842a; bone taxon K) manual ungual fragment (Russell, 1996)
(CMN 50842b; bone taxon L) partial manual ungual (Russell, 1996)
(CMN 50987; bone taxon P; cast) incomplete pedal ungual (Russell, 1996)
(MNHN coll.; Gara Sbaa) teeth (Lavocat, 1954)
(MNHN coll.; Kouah Trick) tibia (630 mm) (Lavocat, 1954)
(MNHN coll.; Kouah Trick) teeth (Lavocat, 1954)
(MNHN coll.; Tabroumit) several teeth (Lavocat, 1954)
(ROM 65779) (>5 year old adult) incomplete femur (Evans, Barrett, Brink and
Carrano, 2015)
Comments- Lavocat (1954) mentioned a coelurosaur (sensu Huene) tibia which "looks very much like the tibia of Elaphrosaurus bambergi" and "also resembles very much the two tibiae figured by STROMER (1934, pl. III, fig. 1, 2) as cf. Elaphrosaurus.
The relative orientation of the proximal and distal extremities is
exactly the same as in the figure 1 b of STROMER. Certain detailed
differences, notably in the development of the proximal antero-external
crest, permit one to think that pertains to a distinct species."
As Stromer's tibiae (IPHG 1912 VIII 76 and 1912 VIII 192) have since
been referred to Tetanurae, the placement of Lavocat's tibia in more
generalized Averostra is uncertain, although the similar size and
stratigraphic
placement could suggest it is closely related.
Russell (1996) proposed multiple informal 'bone taxa' for isolated Kem
Kem theropod elements. Bone taxa G, I, J and M (in part) are here
referred to Spinosaurus
based on Ibrahim et al. (2014) for the first three and Chiarenza et al. (2016)
for the last, bone taxon F to a Xuanhanosaurus-like tetanurine, and bone
taxon H to a noasaurid. McFeeters (2013) could only identify CMN 50810 to Saurischia
incertae sedis, though he noted it was possibly a non-abelisauroid ceratosaur
if theropod (so maybe a juvenile Deltadromeus). Chiarenza and Cau referred
bone taxon C to Abelisauroidea based on similarity to Libyan abelisaur PRC.NF.1.21.
References-
Lavocat, 1954. Sur les dinosauriens du Continental Intercalaire des Kem-Kem
de la Daoura. Comptes Rendus 19th Intenational Geological Congress, 1952.
1, 65-68.
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.
Ibrahim, Sereno, Dal Sasso, Maganuco, Fabbri, Martill, Zouhri, Myhrvold and
Iurino, 2014. Semiaquatic adaptations in a giant predatory dinosaur. Science.
345(6204), 1613-1616.
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.
Chiarenza and Cau, 2016. A large abelisaurid (Dinosauria, Theropoda) from Morocco
and comments on the Cenomanian theropods from North Africa. PeerJ. 4:e1754.
undescribed Averostra (Bardet, Pereda Suberbiola, Jouve, Bourdon, Vincent,
Houssaye, Rage, Jalil, Bouya and Amaghzaz, 2010)
Late Maastrichtian, Late Cretaceous
Couche 3, Morocco
Material- two teeth
Comments- Supposed to be described in Pereda-Suberbiola et al. (in prep.).
Reference- Bardet, Pereda Suberbiola, Jouve, Bourdon, Vincent, Houssaye,
Rage, Jalil, Bouya and Amaghzaz, 2010. Reptilian assemblages from the Latest
Cretaceous - Palaeogene phosphates of Morocco: From Arambourg to present time.
Historical Biology. 22(1), 186-199.
undescribed Averostra (Mahammed, Lang, Mami, Mekhali, Benhamou, Bouterfa, Kacemi, Cherief, Chaouati and Taquet, 2005)
Callovian, Middle Jurassic
Rouis El Djir Mount, Algeria
Material- (Centre de Recherche et Developpement of the SONATRACH Company coll.) teeth
Comments- Discovered between 1999 and 2005, Mahammed et al. (2005) noted that the holotype remains of Chebsaurus "were found associated with other indeterminate sauropod bones and theropod teeth."
Reference- Mahammed, Lang,
Mami, Mekhali, Benhamou, Bouterfa, Kacemi, Cherief, Chaouati and
Taquet, 2005. The 'Giant of Ksour', a Middle Jurassic sauropod dinosaur
from Algeria. Comptes Rendus Palevol. 4, 707-714.
undescribed Averostra (Bassoullet and Iliou, 1967)
Albian, Early Cretaceous
Continental Intercalaire, Oued Boudjihane, Algeria
Material- (small?) teeth
(large) ?cervical vertebra
Comments- Bassoullet and Iliou (1967) reported (translated) "numerous teeth of carnivorous theropods
some of which are attributable to the genus Carcharodontosaurus, others to coelurosaurids" and "1
vertebra (probably cervical) of a large theropod." Bassoullet
worked at the Facult� des sciences de Paris, but this was dissolved in
1970 so that if the remains were stored there they may have been moved
to one of the University of Paris campuses or the University of Orleans.
References- Bassoullet and
Iliou, 1967. D�couverte de Dinosauriens associ�s � des Crocodiliens et
des Poissons dans le Cr�tac� inf�rieur de l'Atlas saharien (Alg�rie).
Compte Rendu Sommaire des S�ances de la Soci�t� G�ologique de France.
7, 294-295.
unnamed Averostra (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- (MGGCTUN26; Morphotype 7) (juvenile?) lateral tooth (20x11x7 mm)
(MGGCTUN33; Morphotype 1) anterior tooth (60x27x17 mm)
(MGGCTUN77; Morphotype 7) (juvenile?) lateral tooth (14x11x7 mm)
(MGGCTUN112; Morphotype 1) anterior tooth (68x27x22 mm)
Comments- Fanti et al. (2014)
recovered Morphotype 1 as a non-abelisaurian abelisauroid using
Hendrickx's theropod dental matrix, but noted this is poorly
supported. They further "refrain from referring Morphotype 1 to
as premaxillary teeth of a tetanuran, as the former does not display a
U-shaped cross-section nor a lingual migration of the anterior carina
as commonly observed in large tetanurans ... , and disparity in the
denticles density between the carinae." Thus perhaps these are
mesialmost dentary teeth of a more generic averostran outside
Megalosauria and Allosauroidea. Fanti et al. also noted three
teeth from the Elrhaz Formation (MNHN GRD553a, GRD553b and GAD600) have
this same morphotype. The authors referred Morphotype 7 to
Abelisauridae or Carcharodontosauridae, considering them "posterior
lateral teeth or possibly juvenile lateral teeth."
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.
undescribed possible Averostra (Nessov, Zhegallo and Averianov, 1998)
Cenomanian, Late Cretaceous
'unnamed' beds, Mizdah Formation, Draa Ubari, Libya
Material- (ZIN PC coll.) teeth, fragments
Comments- Nessov et al. (1998) mentioned "bone fragments and teeth of theropod (?) dinosaurs"
References- Nessov, Zhegallo and Averianov, 1998. A new locality of Late
Cretaceous snakes, mammals and other vertebrates in Africa (western Libya).
Annales de Pal�ontologie. 84(3-4), 265-275.
Rage and Cappetta, 2002. Vertebrates from the Cenomanian, and the geological
age of the Draa Ubari fauna (Libya). Annales de Pal�ontologie. 88, 79-84.
unnamed Averostra (Stromer, 1934)
Early Cenomanian, Late Cretaceous
Baharija Formation, Egypt
Material- (IPHG 1911 XII 31; destroyed) proximal caudal vertebra (105 mm)
(IPHG 1912 VIII 60; destroyed) skull fragment, centrum (~140 mm)
....(IPHG 1912 VIII 60; paratype of Deltadromeus agilis; destroyed) partial
scapula, coracoid
....(IPHG 1912 VIII 60a; destroyed) proximal caudal vertebra (~100 mm)
....(IPHG 1912 VIII 60b; destroyed) proximal caudal vertebra (125 mm)
....(IPHG 1912 VIII 60c; destroyed) proximal caudal vertebra (130 mm)
....(IPHG 1912 VIII 60d; destroyed) partial proximal caudal vertebra (120 mm)
....(IPHG 1912 VIII 60e; destroyed) proximal caudal vertebra (120 mm)
....(IPHG 1912 VIII 60f; destroyed) proximal caudal vertebra (~125 mm)
....(IPHG 1912 VIII 60g; destroyed) partial distal caudal vertebra (115 mm)
....(IPHG 1912 VIII 60h; destroyed) distal caudal vertebra (120 mm)
(IPHG 1912 VIII 63; destroyed) proximal ?metatarsal
....(IPHG 1912 VIII 63a; destroyed) proximal caudal centrum (~110 mm)
....(IPHG 1912 VIII 63b; destroyed) proximal caudal vertebra (120 mm)
....(IPHG 1912 VIII 63c; destroyed) incomplete proximal caudal vertebra (123
mm)
....(IPHG 1912 VIII 63d; destroyed) proximal caudal vertebra (119 mm)
....(IPHG 1912 VIII 63e; destroyed) distal caudal centrum (87 mm)
....(IPHG 1912 VIII 63f; destroyed) distal caudal vertebra (78 mm)
....(IPHG 1912 VIII 63g; destroyed) distal caudal centrum (71 mm)
(IPHG 1912 VIII 70; paratype of Deltadromeus agilis; destroyed) fibula
(1.04 m)
(IPHG 1912 VIII 73; destroyed) incomplete astragalus (180 mm wide)
(IPHG 1912 VIII 75; destroyed) metatarsal IV (430 mm)
(IPHG 1912 VIII 77; destroyed) incomplete scapula, coracoid fragment
(IPHG 1912 VIII 78; paratype of Deltadromeus agilis; destroyed) proximal
tibia
(IPHG 1912 VIII 81; paratype of Bahariasaurus ingens; paratype of Deltadromeus
agilis; destroyed) pubes (550 mm)
(IPHG 1912 VIII 82; destroyed) ischia (325 mm)
(IPHG 1912 VIII 85; destroyed) distal femur
(IPHG 1912 VIII 88a; destroyed) (juvenile) incomplete dorsal vertebra (70 mm)
....(IPHG 1912 VIII 88b; destroyed) centrum (55 mm)
....(IPHG 1912 VIII 88c; destroyed) partial dorsal rib
....(IPHG 1912 VIII 88d; destroyed) distal pubis
(IPHG 1912 VIII 89; destroyed) pedal phalanx (58 mm)
(IPHG 1912 VIII 90; destroyed) incomplete ?manual ungual
(IPHG 1912 VIII 92; destroyed) three incomplete chevrons
(IPHG 1912 VIII 177; destroyed) metatarsals II (523, 543 mm)
(IPHG 1912 VIII 182; destroyed) incomplete humeri or metatarsals
(IPHG 1912 VIII 190; destroyed) (juvenile?) distal tibia
(IPHG 1912 VIII 193; destroyed) incomplete humerus
Comments- This material was mostly referred to Theropoda gen. et sp.
indet. by Stromer (1934), and is retained together as Averostra here for
the moment, although study of individual specimens will no doubt allow referral
to more precise clades. As with all of Stromer's Baharija material, the fossils
themselves were destroyed in 1944.
The pubis IPHG 1912 VIII 81 was referred to Bahariasaurus by Stromer
and stated to be "generally very similar to it", but differs several
ways. Bahariasaurus has a less conspicuous and more proximally placed
lateral flaring (15% down the shaft, compared to 21%). The distal end is not
flared laterally. There is an extensive separation of the pubic shafts distally,
and the interpubic foramen is more distally placed (80% down the shaft, vs.
71%). The proximally placed interpubic foramen suggests this is tetanurine,
while the absent obturator foramen is like spinosaurids, allosaurians and most
coelurosaurs. Sereno et al. (1996) referred it to Deltadromeus, but that
genus should have an interpubic foramen placed distally in the pubic boot itself
as in other ceratosaurs, making the referral unlikely. The specimens listed
under IPHG 1912 VIII 60, 69 and 70 were tentatively referred to Bahariasaurus.
IPHG 1912 VIII 60a-h is a series of caudals which Stromer wrote of as belonging
to a single individual. a is separated proximally, and g-h are located far more
distally. The centra are amphicoelous to amphiplatyan, g-h much broader than
tall, a-d have a ventral groove which is missing in g-h, and a-c have pleurocoels
but d-h do not. All vertebrae have neural spines, but g-h lack transverse processes.
Stromer wrote these were associated with the pectoral girdle, though he doesn't
state whether the skull fragment or larger centrum were associated more exactly
than being from the same layer. The scapula IPHG 1912 VIII 60 is missing most
of the acromion and the distal end, but was at least 7.6 times longer than the
minimum blade width. There is a slight midshaft expansion anteriorly, and the
glenoid faces completely ventrally. The associated coracoid is 84% as deep as
it is long, excluding the well developed posteroventral process that is missing
its distal end. No obvious coracoid tuber is present, nor is there a subglenoid
fossa. Stromer states the pectoral girdle IPHG 1912 VIII 77 is very similar
in size and shape, and shows the distal end of the scapula was unexpanded. He
declined to officially refer the material to a named species because the pectoral
girdle of Spinosaurus was unknown, and remains so today. Interestingly,
the coracoid is very similar to Baryonyx, differing only in the more
posteriorly positioned coracoid foramen and the lesser concavity below the glenoid.
The scapulae are also similar, though Baryonyx's is expanded slightly
at the distal end and has a less projected glenoid region. Note that the caudals
are unlike spinosaurids' however, as the latter lack pleurocoels. Stromer tentatively
referred femur IPHG 1912 VIII 69 to Bahariasaurus based on differences
from Spinosaurus and Carcharodontosaurus and similarity in size
with referred Bahariasaurus specimen IPHG 1912 VIII 62 which was discovered
nearby. Fibula IPHG 1912 VIII 70 was referred to Bahariasaurus for the
same reasons, though note Stromer considered it to belong to a larger individual
than the femur although they were discovered in the same locality. The femur
was referred to Deltadromeus by Sereno et al., which is followed here. The fibula's deep
proximomedial fossa indicates it is probably from a ceratosaur or avetheropod,
so it's possible they belong to the same taxon. Sereno et al. also referred the scapulocoracoid
IPHG 1912 VIII 60 and fibula IPHG 1912 VIII 70 discussed
above to Deltadromeus. The scapulocoracoid is more similar to Baryonyx,
but the fibula is rather similar to Deltadromeus and may belong to that genus.
IPHG 1912 VIII 78, 85 and 190 were referred to aff. Erectopus sauvagei
by Stromer, but none of the elements (distal femur, proximal and distal tibia) are especially similar to
Erectopus. They were all found in different deposits and may belong to
different taxa. Sereno et al. referred the proximal tibia to Deltadromeus,
and it is very similar to that genus so this may be correct.
Stromer misidentified IPHG 1912 VIII 82 as pubes, but the strong transverse
tapering distally, obturator process and lack of an interpubic foramen all resemble
ischia more. IPHG 1912 VIII 63 was identified as proximal femur, but resembles
a metatarsal more. Similarly, IPHG 1912 VIII 177 was identified as a humerus
but is a metatarsal II.
References- Stromer, 1934. Ergebnisse der Forschungsreisen Prof. E. Stromers
in den W�sten �gyptens. II. Wirbeltierreste der Baharije-Stufe (unterstes
Cenoman). 13. Dinosauria. Abhandlungen der Bayerischen Akademie der Wissenschaften
Mathematisch-naturwissenschaftliche Abteilung, Neue Folge. 22, 1-79.
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.
undescribed Averostra (Churcher and Russell, 1992)
Early-Middle Campanian, Late Cretaceous
lungfish site 2 km SE of Baris, Quseir Formation, Kharga Oasis, Egypt
Material- (ROM? coll.) two teeth
Comments- Churcher and Russell
(1992) indicates in March 1991 and/or 1992 "two large theropod teeth"
were located in the Baris Formation. The locality was described
later in Churcher (1995) who used "the terminology of the Geological
Survey of Egypt and refer to them as the Quseir Formation" as did
future studies, although note the Spinosaurus and Carcharodontosaurus tooth he mentions there were found in 1993 at a different locality (Atrun).
Reference- Churcher and
Russell, 1992. Terrestrial vertebrates from Campanian strata in Wadi
el-Gedid (Kharga and Dakhleh Oases), Western Desert of Egypt. Journal
of Vertebrate Paleontology. 12(3), 23A.
Churcher, 1995. Giant Cretaceous lungfish Neoceratodus tuberculatus
from a deltaic environment in the Quseir (=Baris) Formation of Kharga
oasis, Western Desert of Egypt. Journal of Vertebrate Paleontology.
15(4), 845-849.
unnamed averostran (Salem, O'Connor, Gorscak, El-Sayed, Sertich, Seiffert and Sallam, 2021)
Middle Campanian, Late Cretaceous
Tineida, El Hindaw Member, Quseir Formation, Dakhla Oasis, Egypt
Material- (MUVP 199) (subadult or adult) partial ~third caudal vertebra (69.3 mm)
Comments- Discovered in 2008 or
2010 (Sallam et al., 2016), this is an amphicoelous caudal centrum 14%
taller than wide anteriorly and with no pleurocoels. The ventral
surface lacks a median groove or keel. At 8% longer than tall,
proportions are closest to the third caudal in Carnotaurus. Salem et al. (2021) merely list it as Theropoda indet..
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 Averostra (Gemmellaro, 1921)
Late Campanian, Late Cretaceous
Duwi Formation, Al Sharauna, Gebel Duwi and/or Gebel Nakheil, Egypt
Material- (MGUP coll.; lost) teeth
Comments- Gemmellaro (1921) referred several remains to Megalosaurus crenatissimus
(Abelisauridae indet. here) from three localities in eastern
Egypt. These included a smaller tooth 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. The MGUP only has three teeth in their collection
now (Di Patti, pers. comm. 6-2023), none of which match Figure 14.
Reference- Gemmellaro, 1921. Rettili ma�strichtiani di Egitto. Giornale
di Scienze Naturali ed Economiche. 32, 339-351.
unnamed Averostra (Lapparent, 1960)
Aptian, Early Cretaceous
Elrhaz Formation, Niger
Material- (MNHN GAD600; Morphotype 1) anterior tooth (58x22x13 mm) (Fanti, Cau, Martinelli and Contessi, 2014)
(MNHN GDF coll.) posterior skull fragment, dentary fragment,
dozen teeth, vertebrae, manual ungual (~290 mm), ilium, proximal femur, several
metatarsals, phalanges, three unguals (Taquet, 1976)
(MNHN GRD553 a; Morphotype 1) anterior tooth (62x21x13 mm) (Fanti, Cau, Martinelli and Contessi, 2014)
(MNHN GRD553 b; Morphotype 1) anterior tooth (82x32x18 mm) (Fanti, Cau, Martinelli and Contessi, 2014)
(MNHN Td. 2250) anterior tooth (70 mm) (Lapparent, 1960)
Comments- Taquet (1976) notes the tooth MNNHN Td. 2250 is not referrable
to Carcharodontosaurus, contra Lapparent.
Fanti et al. (2014) list three teeth as "Large theropod indet." and
state these Gadoufaoua specimens are referrable to their Morphotype 1
from Kem Kem, which are placed in Averostra indet. here.
References- Lapparent, 1960. Les dinosauriens du "Continental intercalaire"
du Sahara central. Memoirs of the Geological Society of France. 88A, 1-57.
Taquet, 1976. G�ologie et Pal�ontologie du Gisement de Gadoufaoua
(Aptien du Niger). Cahiers de Pal�ontologie, Centre National de la Recherche
Scientifique, Paris. 191 pp.
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 possible Averostra (Werner, 1994)
Cenomanian, Late Cretaceous
Wadi Milk Formation, Sudan
Material- (Vb-715) pedal ungual
(Vb-716) pedal ungual
Comments- These two unguals were referred to Ornithomimosauria by Werner
(1994), but differ from known taxa in being more dorsoventrally compressed and
broader, with no lateral grooves or flanges, almost no posterodorsal process,
a low flexor tubercle instead of a fossa, and paired fossae posterior to this
with a foramen in each. They may not be theropod.
Reference- Werner, 1994. Die kontinentale Wirbeltierfauna aus der unteren
Oberkreide des Sudan (Wadi Milk Formation). In Kohring and
Martin (eds.). Miscellanea Palaeontologica 3: Festschrift Bernard Krebs. Berliner
Geowissenschaften Abhandlungen, Reihe E. 13, 221-249.
undescribed averostran (Hall and Goodwin, 2007)
Tithonian, Late Jurassic
Mugher Mudstone, Ethiopia
Material- tooth
Reference- Hall and Goodwin, 2007. A preliminary analysis of dinosaur
teeth from the Mugher Mudstone of Ethiopia. Journal of Vertebrate Paleontology.
27(3), 86A.
unnamed Averostra (Rauhut, 2011)
Late Jurassic
Tendaguru Formation, Tanzania
Material- (NHMUK coll.) few teeth
Comments- Rauhut (2011) stated
"Although theropod remains were probably also recovered during the
British Tendaguru Expeditions (see Maier 2003), only a few teeth have
been prepared and are currently available in the collections of the
NHMUK (pers. obs.)."
Reference-
Rauhut, 2011. Theropod dinosaurs from the Late Jurassic of Tendaguru (Tanzania).
Palaeontology. 86, 195-239.
unnamed Averostra (Janensch, 1925)
Late Tithonian, Late Jurassic
Upper Dinosaur Member of the Tendaguru Formation, Tanzania
Material- (HMN MB R 1934; = MW 5) distal caudal centrum (119 mm)
(HMN MB R 1935; = MW 3) (~7.8 m) partial proximal caudal centrum
(HMN MB R 1940; = TL 45) proximal caudal vertebra (105 mm)
(HMN MB R 2160; = MW 2) incomplete quadrate (transverse condylar width 140 mm)
(HMN MB R lost; = EH 103) (adult) incomplete posterior dorsal vertebra (80 mm)
(HMN MB R lost; = St 297) cervical neural arch
Comments- MB R 1934, 1935 and 2160 were syntypes of Ceratosaurus roechlingi
(Janensch, 1925), but the latter is too large to belong to the lectotype individual
and none of the specimens is diagnostic beyond Theropoda (Rauhut, 2011).
Rauhut noted the caudal MB R 1940 shows similiarities to his new carcharodontosaurid
Veterupristisaurus, but it derives from higher sediments than that genus.
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, 2011. Theropod dinosaurs from the Late Jurassic of Tendaguru (Tanzania).
Palaeontology. 86, 195-239.
undescribed averostran (Bond and Bromley, 1970)
Early Cretaceous
Gokwe Formation, Zimbabwe
Material- (University of Zimbabwe coll.) tooth (~37x~20x? mm)
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, but the second is narrower and more recurved, so referred
to Averostra
indet. here. Fine serrations are present at least distally (~7
per 5 mm), which are not as tall as in the abelisaurid teeth.
Reference-
Bond and Bromley, 1970. Sediments with the remains of dinosaurs near
Gokwe, Rhodesia. Palaeogeography, Palaeoclimatology, Palaeoecology.
8(4), 313-327.
unnamed Averostra (Mateer, 1987)
Late Jurassic
Enon Conglomerate, South Africa
Material- (SAM 643) tooth (28 mm)
(SAM 649) tooth (>32 mm)
Comments- These teeth lack mesial serrations, and have 18-20 serrations
per 5 mm on the distal carina. As no other details are available besides general
shape in lateral view and absence of vertical striations (unlike some ceratosaurids),
it is unlikely these teeth can be classified more precisely based on published
data.
Reference- Mateer, 1987. A new report of a theropod dinosaur from South
Africa. Palaeontology. 30(1), 141-145.
unnamed averostran (Mateer, 1987)
Berriasian-Barremian, Early Cretaceous
Sundays River Formation, South Africa
Material- (SAM K1475) pedal ungual (94 mm)
Comments- Mateer (1987) referred this ungual to either Megalosauridae
or Allosauridae, while Nessov (1995) referred it to Therizinosauria or "groups
most closely related to them" (which in his opinion consisted of spinosaurids
and dryptosaurids). Neither of these opinions was made in the context of a modern
understanding of theropod phylogeny however. Provisional comparisons suggest
it more closely resembles pedal unguals of Sinraptor and Poekilopleuron
than those of any therizinosaurs (Beipiaosaurus, Alxasaurus, Nothronychus,
Erlikosaurus), which tend to be deeper and more curved.
References- Mateer, 1987. A new report of a theropod dinosaur from South
Africa. Palaeontology. 30(1), 141-145.
Nessov, 1995. Dinosaurs of nothern Eurasia: new data about assemblages, ecology,
and paleobiogeography. Institute for Scientific Research on the Earth's Crust,
St. Petersburg State University, St. Petersburg. 1-156.
unnamed possible averostran (Huene, 1934)
Barremian-Santonian, Early Cretaceous-Late Cretaceous
Guichon Formation, Uruguay
Material- (CA coll.; tooth B; lost) premaxillary tooth (9.3x5.2x3.3 mm)
Comments- The tooth is similar to tyrannosaurid premaxillary
teeth in being narrow and D-shaped with both carinae on the lingual
edge, with at loeast one carina having 4.5 serrations per mm. The
surface between carinae is deeply concave.
Huene (1934) identified this tooth as an ornithomimid, but contra Soto et al. (2011) was unjustified in doing so by then as Struthiomimus had a described edentulous skull as of 1916 even ignoring Dromiceiomimus samueli
in 1928. Huene's identification was based on a tooth described by Lambe (1902) provisionally as Ornithomimus altus
(CMN coll.; plate XIV figure 12-13), but which can now be identified as
an albertosaurine premaxillary tooth. Soto et al. suggested Mones
(1997) might have tentatively
identified this tooth as a sebecosuchian and also noted Soto and
Cambiaso (2006) reidentified the element as Theropoda indet..
Ford (online
2015) placed it in Troodontidae without comment. Huene states the
tooth was discovered by Aznarez with the type of Uruguaysuchus,
which is in the private Colecci�n Azn�rez according to Soto et al.
(2011) , who also state the teeth described by Huene are lost.
References- Lambe, 1902. New genera and species from the
Belly River series (Mid-Cretaceous). Geological Survey of Canada
Contributions to Canadian Palaeontology. 3(2), 25-81.
Huene, 1934. Neue Saurier-Z�hne aus der Kreide von Uruguay. Centralblatt f�r Mineralogie,
Geologie und Pal�ontologie, Abteilung B: Geologie und Pal�ontologie.
1934(4),183-189.
Mones, 1997. Los vertebrados mesozoicos del Uruguay y sus relaciones
con los de �reas vecinas. In Arroyo Cabrales and Polaco (eds.).
Homenaje al Profesor Ticul �lvarez. Mexico D.F., Mexico: Instituto
Nacional de Antropolog�a e Historia, Colecci�n Cient�fica. 357, 205-222.
Soto and Cambiaso, 2006. Reinterpretaci�n de los dientes de dinosaurios
de la Formaci�n Guich�n: Iguanodontes basales y ter�podos no
ornitom�midos. Ameghiniana. 43 (Suppl.), R55-R56.
Soto, Pol and Perea, 2011. A new specimen of Uruguaysuchus aznarezi
(Crocodyliformes: Notosuchia) from the middle Cretaceous of Uruguay and
its phylogenetic relationships. Zoological Journal of the Linnean
Society. 163, S173-S198.
Ford, online 2015. http://www.paleofile.com/Dinosaurs/Theropods/Troodonincertae.asp
undescribed Averostra (Kellner, Azevedo, Carvalho, Henriques and
Costa, 2004)
Campanian-Maastrichtian, Late Cretaceous
Parecis Group, Brazil
Holotype- teeth and possible elements
Comments- These were initially noted as being from the Bauru Group, but
Bittencourt and Langer (2011) reassigned the locality to the later Parecis Group.
References- Kellner, Azevedo, Carvalho, Henriques and Costa, 2004. Bones
out of the jungle: On a dinosaur locality from Matio Grosso, Brazil. Journal
of Vertebrate Paleontology. SVP abstracts. 150-151.
Bittencourt and Langer, 2011. Mesozoic dinosaurs from Brazil and their biogeographic
implications. Anais da Academia Brasileira de Ci�ncias. 83(1), 23-60.
undescribed Averostra (Bertini, 1993)
Early Maastrichtian, Late Cretaceous
Adamantina Formation, Bauru Group, Brazil
Material- (LF-019-R) tooth (15 mm) (Geroto and Bertini, 2014) D
(LF-020-R) tooth (12 mm) (Geroto and Bertini, 2014) E
(LF-023-R) tooth (21 mm) (Geroto and Bertini, 2014) H
(LF-026-R) tooth (10 mm) (Geroto and Bertini, 2014) K
(LF coll.; V17 provisionally) tooth (11.5 mm) (Geroto and Bertini, 2014) N
(UFRJ-DG 375-Rd) tooth (Candeiro, Abranches, Abrantes, Avilla, Martins,
Moreira, Torres, Bergqvist, 2004)
(UFRJ-DG 376-Rd) tooth (Candeiro, Abranches, Abrantes, Avilla, Martins,
Moreira, Torres, Bergqvist, 2004)
(UFRJ-DG 377-Rd) tooth (Candeiro, Abranches, Abrantes, Avilla, Martins,
Moreira, Torres, Bergqvist, 2004)
Comments- Bertini and
Franco-Rosas (2001) state "Troodontidae teeth were recognized" among
over 200 specimens, probably originating with Bertini's (1993) and
Franco's (1999) theses. Geroto and Bertini (2014) described five such
teeth (LF-019-R, 020-R, 023-R, 026-R and provisionally labeled V17)
serrated both mesially and distally with low DSDIs and transversely
compressed and recurved. Considering the lack of South American
troodontids and the absence of clear troodontid characters, these may
belong to e.g. noasaurids instead and are placed in Averostra here.
References-
Bertini, 1993. Paleobiologia do Grupo Bauru, Cret�ceo Superior
continental da Bacia do Paran�, com �nfase em sua fauna de amniotas.
PhD thesis, Universidade Federal do Rio de Janeiro. 493 pp.
Franco, 1999. Dentes de teropodomorfos do Cret�ceo Superior da Bacia do
Paran�. An�lise em Microscopia Eletr�nica de Varredura. Masters thesis,
Universidade Estadual Paulista. 113 pp.
Bertini and Franco-Rosas, 2001. Scanning electron microscope
analysis on Maniraptoriformes teeth from the Upper Cretaceous of southeastern
Brazil. Journal of Vertebrate Paleontology. 21(3), 33A.
Franco Rosas, 2002. Methodological parameters for identification and
taxonomic classification of isolated theropodomorph teeth. Anais da
Academia Brasileira de Ci�ncias. 74(2), 367.
Candeiro, Abranches, Abrantes, Avilla, Martins, Moreira, Torres,
Bergqvist, 2004. Dinosaur remains from western Sao Paulo State, Brazil (Bauru
Basin, Adamantina Formation, Late Cretaceous). Journal of South American Earth
Sciences. 18, 1-10.
Geroto and Bertini, 2014. New records of fossil vertebrates from the
Upper Cretaceous Adamantina Formation (Bauru Group), southeastern
Brazil. Revista do Instituto Geol�gico, S�o Paulo. 35(2), 39-56.
unnamed Averostra (Bertini, 1993)
Late Maastrichtian, Late Cretaceous
Serra da Galga Formation of the Bauru Group, Brazil
Material- teeth (Bertini, 1993)
(CPP 128, 243, 271, 371) four teeth (Candeiro, Bergqvist, Novas and Currie, 2004)
(CPP 374) tooth (Candeiro, Currie and Bergqvist, 2012)
Comments- In 2021 the Serra da Galga and Ponte Alta Members of the Marilia Formation were recognized as the Serra da Galga Formation.
Bertini and
Franco-Rosas (2001) state "Troodontidae teeth were recognized" among
over 200 specimens, probably originating with Bertini's (1993) and
Franco's (1999) theses. Candeiro et al. (2012) describes one tooth
(CPP 374) initially listed by Candeiro et al. (2004) resembling the Adamantina Formation troodontid morph of Bertini, and
like those teeth these are here placed more generally as Averostra.
References- Bertini, 1993. Paleobiologia do Grupo Bauru, Cret�ceo Superior
continental da Bacia do Paran�, com �nfase em sua fauna de amniotas.
PhD thesis, Universidade Federal do Rio de Janeiro. 493 pp.
Franco, 1999. Dentes de teropodomorfos do Cret�ceo Superior da Bacia do
Paran�. An�lise em Microscopia Eletr�nica de Varredura. Masters thesis,
Universidade Estadual Paulista. 113 pp.
Bertini and Franco-Rosas, 2001. Scanning electron microscope
analysis on Maniraptoriformes teeth from the Upper Cretaceous of southeastern
Brazil. Journal of Vertebrate Paleontology. 21(3), 33A.
Franco Rosas, 2002. Methodological parameters for identification and
taxonomic classification of isolated theropodomorph teeth. Anais da
Academia Brasileira de Ci�ncias. 74(2), 367.
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, Currie and Bergqvist, 2012. Theropod teeth from the Mar�lia
Formation (Late Maastrichtian) at the paleontological site of Peir�polis
in Minas Gerais State, Brazil. Revista Brasileira de Geoci�ncias. 42(2),
323-330.
undescribed averostran (Pol and Rauhut, 2012)
Callovian, Middle Jurassic
Canadon Asfalto Formation, Chubut, Argentina
Material- three cervical vertebrae
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.
unnamed averostran (Lydekker, 18__ in Huene, 1929)
Cretaceous?
Neuquen, Argentina
Material- dorsal vertebra (100 mm)
References- Lydekker, 18__.
Huene, 1929. Los saurisquios y ornitisquios del Cretac�o Argentino. Anales
del Museo de La Plata (series 3). 3, 1-196.
unnamed Averostra (Huene, 1929)
Cenomanian, Late Cretaceous
Mata Amarilla Formation (= Pari Aike Formation), Santa Cruz, Argentina
Material- (Ameghino coll.) partial tooth (26 mm)
(Ameghino coll.) distal manual phalanx
(Ameghino coll.) long bone epiphysis
Reference- Huene, 1929. Los saurisquios y ornitisquios del Cretac�o
Argentino. Anales del Museo de La Plata (series 3). 3, 1-196.
unnamed averostran (Huene, 1929)
Late Coniacian, Late Cretaceous
Plottier Formation of the Rio Neuquen Group, Neuquen, Argentina
Material- (MLP coll.) incomplete tooth (16 x 9.5 x 6.3 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.
Huene described the Plottier tooth under the heading "tooth of a carnivorous
saurischian", stating it was not his intention to decide whether it was
a coelurosaur or carnosaur. The mesial edge is more convex than the distal edge,
especially basally. There are 12.5 serrations per 5 mm, stated to be on both
mesial and distal carinae. The tooth seems to be from a somewhat anterior position,
as it has an asymmetrical section, with one side generally more convex except
for a distally placed concavity. What are described as growth lines on the labial
and lingual surfaces may be enamel wrinkles, which are apically concave in the
distal half of one side. This tooth is difficult to place without further study
of Gondwanan small theropod dentitions and is here assigned to Averostra.
References- Huene, 1929. Los saurisquios y ornitisquios del Cretac�o
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
undescribed averostran (Filippi, Martinelli and Garrido, 2015)
Santonian, Late Cretaceous
Bajo de la Carpa Formation of the Rio Colorado Subgroup, Neuquen, Argentina
Material- (MAU-Pv-N 496/1) tooth
Reference- Filippi, Martinelli and Garrido, 2015. Una nueva asociaci�n
de dientes vertebrados para la Formaci�n Bajo de la Carpa (Santoniense,
Cret�cico Superior) en Rinc�n de los Sauces, Neuqu�n, Argentina.
Revista Espa�ola de Paleontolog�a. 30(2), 223-238.
undescribed averostran (Gasparini, Sterli, Parras, O'Gorman, Salgado,
Varela and Pol, 2015)
Campanian-Maastrichtian, Late Cretaceous
La Colonia Formation, Chubut, Argentina
Material- (MPEF-PV 10829) incomplete metatarsal III, partial metatarsal
IV, distal metatarsal
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 averostran (Long and Cruickshank, 1996)
Hauterivian-Barremian, Early Cretaceous
Birdrong Sandstone, Western Australia
Material- (WAM 96.5.1) (~5 m) incomplete mid caudal vertebra (~63 mm)
Comments- Long and Cruickshank (1996) referred this to Tetanurae without
justification, and Carrano et al. (2012) believed it to be Theropoda indet..
References- Long and Cruickshank, 1996. First record of an Early Cretaceous
theropod dinosaur bone from Western Australia. Records of the Western Australian
Museum. 18, 219-222.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
unnamed Averostra (Woodward, 1906)
Late Barremian-Early Aptian, Early Cretaceous
Wonthoggi Formation of the Strzelecki Group, Victoria, Australia
Material- (NMV P10058) incomplete ?pedal ungual (Woodward, 1906)
(NMV P186054) incomplete manual phalanx (90 mm) (Benson, Rich, Vickers-Rich
and Hall, 2012)
(NMV P186151) partial pedal phalanx (49 mm) (Cleeland, 2004)
?(NMV P186175) vertebra (Cleeland, 2004)
?(NMV P186218) (large) vertebra (Cleeland, 2004)
(NMV P199050) manual phalanx (95 mm) (Benson, Rich, Vickers-Rich and Hall, 2012)
(NMV P199085) incomplete ?manual ungual (~34 mm) (Benson, Rich, Vickers-Rich
and Hall, 2012)
(NMV P203700) distal caudal vertebra (29 mm) (Benson, Rich, Vickers-Rich and
Hall, 2012)
(NMV P208512) incomplete manual ungual (Benson, Rich, Vickers-Rich and Hall,
2012)
(NMV P209990) incomplete metatarsal ?IV (Benson, Rich, Vickers-Rich and Hall,
2012)
(NMV P210090) incomplete distal caudal vertebra (32 mm) (Benson, Rich, Vickers-Rich
and Hall, 2012)
(NMV P212806) distal caudal vertebra (29 mm) (Benson, Rich, Vickers-Rich and
Hall, 2012)
(NMV P212840) mid caudal neural arch (Cleeland, 2004)
(NMV P216642) incomplete distal caudal vertebra (24 mm) (Benson, Rich, Vickers-Rich
and Hall, 2012)
(NMV coll.) frontal (Benson, Rich, Vickers-Rich and Hall, 2012)
Comments- NMV P199085 is probably the ungual mentioned by Pigdon (online
2000) as being ornithomimosaurian on his Timimus page. Benson et al.
(2012) described it as weakly curved with a low flexor tubercle and flattened
ventral surface which is broader than the dorsal surface. The authors referred
it to Theropoda indet., and there is no evidence it belonged to Timimus.
Rich (2003) reported identifying small theropod skull fragments, but Benson
et al. (2012) state the only cranial element known is an unidentified frontal.
Cleeland (2004) reported NMV P186175 and P186218, though neither are mentioned
by Benson et al..
Kool (2004) reported two small recurved serrationless theropod teeth.
The Dinosaur Dreaming 2007 update notes several theropod teeth and a dorsal
centrum were discovered.
References- Woodward, 1906. On a tooth of Ceratodus and a dinosaurian
claw from the Lower Jurassic of Victoria, Australia. Annals and Magazine of
Natural History, 7th series. 103, 1-3.
Pigdon, online 2000. http://home.alphalink.com.au/~dannj/timimus.htm
Rich, 2003. Research update. Dinosaur Dreaming 2003 Report. 22-23.
Cleeland, 2004. A summary of the status of known Cretaceous vertebrate fossil
localities on the Strzelecki Coast, Victoria, Australia. Dinosaur Dreaming 2004
Field Report. 10-13.
Benson, Rich, Vickers-Rich and Hall, 2012. Theropod fauna from Southern Australia
indicates high polar diversity and climate-driven dinosaur provinciality. PLoS
ONE. 7(5), e37122.
unnamed Averostraa (Currie, Vickers-Rich and Rich, 1996)
Late Aptian-Early Albian, Early Cretaceous
Eumeralla Formation of the Otway Group, Victoria, Australia
Material- (NMV P180856) incomplete metatarsal ?II (Benson, Rich, Vickers-Rich
and Hall, 2012)
(NMV P180899) distal manual phalanx (Benson, Rich, Vickers-Rich and Hall, 2012)
(NMV P185858) distal caudal vertebra (43 mm) (Benson, Rich, Vickers-Rich and
Hall, 2012)
(NMV P186386) surangular (Currie, Vickers-Rich and Rich, 1996)
(NMV P199783) distal caudal vertebra (44 mm) (Benson, Rich, Vickers-Rich and
Hall, 2012)
(NMV P223063) distal caudal vertebra (23 mm) (Benson, Rich, Vickers-Rich and
Hall, 2012)
(NMV P229456) distal caudal vertebra (25 mm) (Benson, Rich, Vickers-Rich and
Hall, 2012)
Comments- Though NMV P186386 was identified as an oviraptorosaur by Currie
et al. (1996) based on its medially projecting coronoid process, Agnolin et
al. (2010) noted the convex margin with no obvious external mandibular fenestra
and lack of articular fusion differed from that clade.
References- Currie, Vickers-Rich and Rich, 1996. Possible oviraptorosaur
(Theropoda, Dinosauria) specimens from the Early Cretaceous Otway Group of Dinosaur
Cove, Australia. Alcheringa. 20(1-2), 73-79.
Agnolin, Ezcurra, Pais and Salisbury, 2010. A reappraisal of the Cretaceous
non-avian dinosaur faunas from Australia and New Zealand: Evidence for their
Gondwanan affinities. Journal of Systematic Palaeontology. 8(2), 257-300.
Benson, Rich, Vickers-Rich and Hall, 2012. Theropod fauna from Southern Australia
indicates high polar diversity and climate-driven dinosaur provinciality. PLoS
ONE. 7(5), e37122.
unnamed averostran (Brougham, Smith and Bell, 2020)
Early Cenomanian, Late Cretaceous
Wallangulla Sandstone Member of Griman Creek Formation, New South Wales, Australia
Material- (LRF 3054) ulna
Comments- Brougham et al. (2020) mention "a possible indeterminate theropod ulna."
Reference-
Brougham, Smith and Bell, 2020. Noasaurids are a component of the
Australian 'mid'-Cretaceous theropod fauna. Scientific Reports. 10:1428.
unnamed averostran (Long, 1995)
Cenomanian-Early Turonian, Late Cretaceous
Molecap Greensand, Western Australia
Material- (WAM 92.7.1) (~4 m) pedal phalanx IV-1 (40.8 mm)
Comments- Long (1995) thought this was probably a carnosaur, but Carrano
et al. (2012) believed it to be Theropoda indet.
References- Long, 1995. A theropod dinosaur bone from the Late Cretaceous
Molecap Greensand, Western Australia. Records of the Western Australian Museum.
17, 143-146.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
undescribed Averostra (Rejcek, 2011)
Early Maastrichtian, Late Cretaceous
Cape Lamb Member of the Snow Hill Island Formation, Vega Island, Antarctica
Material- (MLP 15-I-7-2) pedal phalanx III-1 (Coria, O'Gorman, Cardenas,
Mors, Chornogubsky and Lopez, 2015)
tooth (Rejcek, 2011)
References- Rejcek, 2011. Big haul: Paleontologists return from Antarctic
expedition with about 200 fossils. The Antarctic Sun. 11-16-2011.
Coria, O'Gorman, Cardenas, Mors, Chornogubsky and Lopez, 2015. New dinosaur
records from the Upper Cretaceous of Vega Island, Antarctica. XXIX Jornadas
Argentinas de Paleontolog�a de Vertebrados, resumenes. Ameghiniana. 52(4)
suplemento, 13.