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 catalogd 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.
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Hu, 1993. A new Theropoda (Dilophosaurus sinensis sp. nov.) from
Yunnan, China. Vertebrata PalAsiatica. 31(1), 65-69.
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Xing, Bell, Rothschild, Ran, Zhang, Dong, Zhang and Currie, 2013. Tooth
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Xing, Paulina-Carabajal, Currie, Xu, Zhang, Wang, Burns and Dong, 2014.
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"Allosaurus" medius
Marsh, 1888
Etymology- "medius" is Latin for "moderate",
probably referring to its smaller estimated size of "ten or twelve feet
in length" compared to Allosaurus
fragilis which Marsh had estimated at twenty-five feet long a
decade beforehand (Creisler, pers. comm. 1-8-2024).
= Antrodemus medius (Marsh, 1888) Hay, 1902
= Dryptosaurus medius (Marsh, 1888) Gilmore, 1920
= Labrosaurus medius (Marsh, 1888) Kuhn, 1939
Late Aptian, Early Cretaceous
near Miurkirk USNM 41615, Arundel Formation, Prince George's
County, Maryland, US
Lectotype- (USNM 4972) tooth (~25.3x~14.7x7 mm)
Referred- ?(Goucher College 2521; lost) left pedal phalanx III-1
(110.0 mm) (Lull, 1911)
?(Goucher College 3121; = Charles Coffin coll.; lost?) tooth
(76x28.7x~16 mm) (Bibbins, 1895)
?(Goucher College coll.; = Johns Hopkins University coll.; lost)
incomplete right tibia (~894 mm) (Bibbins, 1895)
?(USNM 5693) incomplete tooth (?x~26x? mm) (Gilmore, 1920)
?(USNM 8446; not USNM 3446) tooth (~64x~27x? mm) (Gilmore, 1920 as 3446)
?(USNM 8447) apical tooth (Gilmore, 1920)
?(USNM 8502; = Goucher College 2534) first sacral centrum (90.0 mm)
(Lull, 1911)
?(USNM 8503; = Goucher College 2614) (juvenile or subadult) incomplete
proximal caudal centrum (107 mm) (Lull, 1911)
?(USNM 8504; = Goucher College 2536) proximal right pedal phalanx II-1
(Lull, 1911)
Diagnosis- Provisionally indeterminate relative to Acrocanthosaurus
atokensis.
Other diagnoses- Marsh (1888)
initially stated its non-measurement characters as "The teeth are
remarkably flat and trenchant, with the edges finely serrated, and the
surfaces very smooth. The limb bones, and even the phalanges, are
unusually hollow, and the latter have the articulations finely
finished", but these are typical of most theropods with the dental
characters matching most non-maniraptoriforms and dromaeosaurids.
Indeed the syntype limb elements have since been referred to
ornithomimosaurs.
As noted below, Lull (1911) distinguished Allosaurus medius from the
type A. fragilis based on
supposed postcranial differences, but these
elements (Gaucher College 2521, USNM 8502) cannot be referred to the
former species.
Comments- Marsh (1888) originally erected medius
for large theropod material from the Arundel Formation (with small
theropod material being named Coelurus
gracilis), "provisionally
referred to the genus Allosaurus"
for no stated reason and with no
explicit difference from A. fragilis.
He mentions teeth
(including USNM 4972) "and bones of the limbs and feet" of which only
an "astragalus ... 55 mm in width; and 50 mm in fore and aft diameter"
(USNM 5652) and "A first phalanx of the hind foot ... 90 mm in length"
(phalanx II-1- USNM 5453) are specified. The latter two elements
were considered cotypes of A. medius
by Lull (1911) who said "Among the
remains referred by Marsh to Allosaurus
medius, the tooth alone may be
with certainty referred to the Theropoda ; the “first phalanx of the
hind foot,” surely, and the astragalus, probably, may be relegated to
the ornithopod dinosaurs of the genus Dryosaurus,
and, together with
other material, become the cotypes of a new species." They thus
became syntypes of his new camptosaurid Dryosaurus grandis, which
became Ornithomimus affinis
in Gilmore's (1920) work so as not to
confuse it with the preexisting species Ornithomimus grandis. As
stated by ICZN Article 74.6, "the first author to have published before
2000 the assumption that the species-group taxon was based upon a
single type specimen is deemed to have designated that specimen as the
lectotype", thus Lull designated USNM 4972 as the lectotype of
Allosaurus medius. Chure
(2000) incorrectly includes USNM 5684
and 5704 as "Marsh material so transferred" (and mistypes USNM 5453 as
USNM 4553, which does not exist in their vertebrate paleontology
collection), but these remained syntypes of Dryosaurus grandis.
Almost none of the historic Arundel dinosaur specimens of Marsh, Lull
and Gilmore had recorded locality data, with most sources simply saying
"Near Muirkirk" (Lull, 1911; Gilmore, 1920). Hatcher (1903)
specified "The exact locality of the Marsh material was certain iron
ore mines on the farm of Mr. William Coffin, and especially in that one
locally known as “Swampoodle” and situated about 1 1/2 miles northeast
of Beltsville, on the Baltimore & Ohio Railway, some 13 miles from
Washington." As shown in Kranz's (1996) Figure 9, the Muirkirk
Iron
Works owned by Coffin were about 2.2 miles from the Swampoodle site, so
this description isn't really that specific and USNM Locality 41615
(where the lectotype, and USNM 5685, 5693 and 8503 are cataloged as
from) probably refers to at least this large area.
The lectotype- The type tooth
was discovered by Hatcher between October 17 and December 17 1887
(based on Appendix C of Kranz, 1996 and Marsh's publication
date). Marsh (1888) initially only says "The teeth are remarkably
flat and trenchant, with the edges finely serrated, and the surfaces
very smooth." He then states "One tooth has the crown 30 mm in
height; its antero-posterior diameter at base 15 mm; and its transverse
diameter 77 mm", which marks this specimen as the lectotype
tooth. Hay (1902) moved the species to Antrodemus, but
unexpectedly did not place Allosaurus
fragilis in that genus, instead
saying "The identification of Marsh's Labrosaurus
with Leidy's
Antrodemus ... has been made
by Mr. F. A. Lucas, of the U. S. National
Museum" and citing "Marsh, O. C. 1888 A, p. 93. (Labrosaurus.)" in
error as Marsh actually used Allosaurus.
Indeed, Hay (1908) later
states "By an inexplicable error the writerᵇ referred to Antrodemus the
remains described by Marsh as Allosaurus
medius." Lull says "the
only one of the original cotypes referable to it, the tooth, is one of
the least distinctive features of the skeleton", distinguishing it from
A. fragilis based on supposed
vertebral (USNM 8502) and phalangeal
(Gaucher College 2521) differences, although each of these elements was
misidentified. Gilmore (1920) wrote "The type tooth has been
carefully compared with teeth of Antrodemus
and Ceratosaurus, and with
those of Upper Cretaceous Theropods; but I have failed to find
characters that would distinguish it from any of those genera. In fact
on the basis of a single tooth I believe it is impossible to determine
the genus to which it pertains, and certainly there are not diagnostic
characters available for generic or specific differentiation. In so far
as the type specimen is concerned it will always remain a form of
doubtful affinities." He transferred it to Dryptosaurus "largely
on geographic considerations" as both are from the east coast of the US
unlike Allosaurus. Kuhn
(1939) explicitly used the combination
Labrosaurus medius, but
erroneously in reference to Marsh 1888,
seemingly repeating Hay's 1902 error. Carpenter et al. (1997) in
their Dryptosaurus
redescription stated "D. potens
and D. medius, are
based on fragmentary material and only superficially resemble
Dryptosaurus; they should be
considered nomina dubia" without
supporting evidence. Harris (1998) wrote "The tooth lacks an
adequate description, and its status is indeterminate." While
Lipka considered several more recently discovered Arundel teeth (USNM
497718, 497722-497726) to be Acrocanthosaurus,
at the time he said the
older specimens "have yet to be compared with the new material."
The lectotype has only ever been figured in labiolingual view, with the
best photo by far being that on the USNM online catalog. The
Crown Height ratio is 1.72 while the Crown Breadth Ratio is 0.48, which
is comparable to e.g. the tenth dentary tooth of Acrocanthosaurus NCSM
14345 (1.68, 0.51) that has a Crown Basal Length of 28.26 mm, very
similar to medius.
There are 13 serrations per 5 mm mesially and 14.6 serrations distally,
within the range of Acrocanthosaurus
(10-20 mesially, 11.5-17.6 distally).
Referred material- Lull (1911)
referred several additional remains to Allosaurus medius
seemingly because of their smaller size than his new Creosaurus potens,
"by far the largest carnivore known from the Arundel formation"
although it is possible locality played a role as the single element of
potens was discovered in
Washington DC instead. Gilmore (1920) stated "While there may be two
large carnivorous dinosaurs (Dryptosaurus?
potens (Lull) and D. medius
(Marsh) present in the Arundel fauna, I can
see no good reason at the present time for believing there is more than
one, even though, as a matter of expediency, both species are retained
in the present paper. Certainly the scattered teeth and other bones
assigned to D. medius
indicate an individual or individuals of
sufficient size to have had an anterior caudal of the dimensions of the
type of Creosaurus potens,
here referred to the genus
Dryptosaurus." By the
next year Gilmore (1921) specified "At this
time Dryptosaurus? medius
rests on a single tooth" and "The few
scattered bones referred to this species by Lull can be assigned, with
equal propriety, to Dryptosaurus?
potens." Huene (1932)
explicitly synonymized Creosaurus
potens with his Antrodemus
medius,
stating (translated) "Gilmore
(52, p. 121) considers it likely that the teeth and vertebrae, which
are found on the same horizon not far from each other, belong
together." Kranz (1998) having created the nomen nudum
"Capitalsaurus" for the C. potens
holotype listed in his Table 3 "large
carnivore" "teeth and various isolated postcrania" as "(possibly "Capitalsaurus"), implying a
possible synonymy but it should be noted medius has priority over potens.
Of the "several larger, better preserved teeth", Lull (1911) mentions
"The most perfect is one in the possession of the Hon. Charles E.
Coffin,¹ Muirkirk, Maryland (pl. xiv, figs. 1, 2). It is about 3 inches
(76 mm.) in length and 1 1/8 inches (28.7 mm.) in the anteroposterior
diameter. The crenulations of the margin cease about midway toward the
root on the anterior convex border, but extend the length of the crown
on the posterior edge." Superscript 1 is given as "Or No. 3121, Goucher
College." As the Goucher College collection was transferred to
the USNM in 1916 and no tooth matching Lull's figure exists in the USNM
catalog, it may be lost (unlike other former GC specimens, Gilmore
1920 does not provide a new USNM number either). This tooth is
first figured as "Allosaurus
tooth" by Bibbins (1895) who says it was
"taken from the “blue charcoal clay,” and lent for study by Hon.
Charles E. Coffin." Lull also says "one ... (No. 5685, Goucher
College) shows decided wear", but Gilmore (1920) correctly notes
this was "Wrongly attributed to Goucher College as this is the catalog
number
of the U. S. National Museum" and this remains true today. This
tooth has since been provisionally referred to Acrocanthosaurus sp. by Carrano
(2024). Gilmore (1920)
states that besides the lectotype, "Other teeth in the U. S. National
Museum Collections are Nos. 5693, 3446, and 8447." Note USNM 3446
is a typo for USNM 8446, which the USNM catalog indicates was
mentioned by Gilmore. It also indicates this and USNM 8447 were
collected by Bibbins in 1894, while USNM 5685 and 5693 were collected
by Hatcher (which according to Kranz' 1996 Appendix C would have been
between October 17 1887 and January 27 1888). Thus when Bibbins
says "In the same beds (“brown charcoal clay”) [as the mysterious
Allosaurus tibia noted below]
and at about the same level, were found
fragments of other bones of similar dimensions, and a tooth (Fig. G)³
probably belonging to the same individual" the tooth referenced is
probably USNM 8446 or 8447 (note the superscript 3 indicates "The tooth
shown in the figure is not the one taken from this bed" but is instead
Goucher College 3121). Carrano determined USNM 8446 to be "not
well-enough preserved for" two characters found by Hendrickx and Mateus
(2014) to be locally apomorphic for Acrocanthosaurus
"to be observed, and I consider them indeterminate to clade within
Theropoda."
A centrum "(No. 2534, G. C.), which seems to be a posterior presacral",
was regarded by Gilmore (1920) "as being the anterior vertebra of the
sacral series and the plane end (which appears to be sutural) as being
posterior. That it is a sacral is indicated not only by comparison with
the articulated sacral series of No. 4734 U.S.N.M. (Antrodemus valens),
but also by the character of the plane articular end for close
articulation with the centrum which followed it and the rapid widening
of the anterior half of the neural canal." Gilmore states this is
"Now Cat. No. 8502 U.S.N.M." Chure (2000) incorrectly lists this
specimen as "two posterior presacrals." The anterior face is
slightly concave, 85.0 mm tall and 105.0 mm wide, and based on the USNM
catalog photo there are no pleurocoels. It was collected by
Coffin in 1894.
"The anterior caudal vertebra (No. 2614 a, G. C.) is that of a young
individual, as the neural arch had not coossified with the centrum"
(Lull, 1911), and Gilmore notes this specimen is "Now Cat. No.
8503, U.S.N.M." It is slightly amphiplatyan and apneumatic, 92 mm
tall and 101 mm wide anteriorly and 93 mm wide posteriorly. "The
first phalanx of the second digit is represented by the proximal half
(No. 2536, G. C.)" (Lull, 1911) and as noted by Gilmore is "Now Cat.
No. 8503,
U.S.N.M.". Its proximal articular surface is 73 mm tall and 58 mm
wide, and as Gilmore says "belongs to the right hind foot." These
two specimens were collected by Bibbins in 1894, and USNM 8503 is
probably what Bibbins (1895) referred to when he wrote "Half a mile
distant [from what is probably USNM 8446] in a bed of similar character
a single vertebra was found which is probably referable to the same
species." Lull also notes "The first phalanx of the third digit
(No. 2521, G. C.) (fig. 2) is entire, most excellently preserved, and
presents a decided similarity to the type of Allosaurus fragilis",
although he states it differs "in being more depressed proximally,
especially in the broader, flatter under surface. Distally, the present
type is not so broad relatively as that of A. fragilis, and the
articular face is more concave transversely." Unfortunately,
Gilmore states "When the Goucher College collection was desposited in
the U.S. National Museum this specimen was missing" although Lull's
figure remains. Lull finally states "There are also two distal
caudals, one No. 5701, the other unnumbered [eventually numbered USNM
6116 as noted by Gilmore], both of the U. S. National Museum
collection", but Gilmore states "their close resemblance to the caudals
of Ornithomimus appears to
indicate their Ornithomimid affinities" and
he referred them to Ornithomimus
affinis. Chure (2000)
incorrectly lists USNM 5701 as the number for both caudals.
Bibbins' (1895) mentions "a tibia, probably of Allosaurus, which
measures ten inches in width, and thirty-two inches in length,
exclusive of the ends, which are lacking." There is no other
record of an Arundel dinosaurian tibial shaft from this period, let
alone one over 813 mm in length without preserved ends (longer than
even the 635 mm type of Pleurocoelus
altus), so exactly what became of
this is unknown. Kranz (online) says "Last seen at Johns Hopkins
University June 1909. The nearly 3 ft long section of dinosaur limb
bone was excavated by workers in Charles Coffin's iron mines (1895?).
It was given a Women's College (Goucher) #3121, but this may be no
longer be on the bone" and includes a photo which he cannot recall the
source of (pers. comm., 1-19-2024). Notably, Goucher College 3121
is the number of the tooth figured by Bibbins, and as it is unlikely
both were given the same number, a mistake has probably occured
somewhere. According to the Internet Archive Wayback Machine,
Kranz's page with quote and photo go back to at least 2009, which is
how they are referenced here. The photo indicates this is a right
tibia based on the lateral curvature of Mapusaurus' proximal tibia and
the abrupt shadowing proximally indicating the cnemial crest is facing
the viewer. Based on comparison to Mapusaurus, the tibia is
~91% complete and would thus be ~894 mm when complete. Not much
more can be ascertained from the photo except it appears more robust
and medially bowed than Mapusaurus
MCF-PVPH-108.68.
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.
Bibbins, 1895. Notes on the paleontology of the Potomac Formation.
Johns Hopkins University Circulars. 15(121), 17-20.
Hay, 1902. Bibliography and catalog of the Fossil Vertebrata of North
America. Bulletin of the United States Geological Survey. 179, 868 pp.
Hatcher, 1903. Discovery of remains of Astrodon (Pleurocoelus) in the Atlantosaurus Beds of Wyoming. Annals
of the Carnegie Museum of Natural History. 2, 9-14.
Hay, 1908. On certain genera and species of carnivorous
dinosaurs, with special reference to Ceratosaurus nasicornis
Marsh. Proceedings of the United States National Museum. 35(1648),
351-366.
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.
Gilmore, 1921. The fauna of the Arundel Formation of Maryland.
Proceedings of the United States National Museum. 59, 581-594.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung
und Geschichte. Monographien zur Geologie und Palaeontologie. 4(1), 361
pp.
Kuhn, 1939. Beitrage zur Keuperfauna von Halberstadt. Palaeontologische
Zeitschrift. 21, 258-286.
Kranz, 1996. Notes on the sedimentary iron ores of Maryland and their
dinosaurian fauna. Maryland Geological Survey Special Publication. 3,
87-111.
Carpenter, Russell, Baird and Denton, 1997. Redescription of the
holotype of Dryptosaurus aquilunguis (Dinosauria: Theropoda)
from the Upper Cretaceous of New Jersey. Journal of Vertebrate
Paleontology. 17(3), 561-573.
Harris, 1998. Large, Early Cretaceous theropods in North America. In
Lucas, Kirkland and Estep (eds.). Lower and Middle Cretaceous
Terrestrial Ecosystems. New Mexico Museum of Natural History and
Science Bulletin. 14, 225-228.
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.
Lipka, 1998. The affinities of the enigmatic theropods of the Arundel
Clay facies (Aptian), Potomac Formation, Atlantic coastal plain of
Maryland. In Lucas, Kirkland and Estep (eds.). Lower and Middle
Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History
and Science Bulletin. 14, 229-234.
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.
Kranz, 2009 online. https://terpconnect.umd.edu/~gdouglas/wanted/index.html
Carrano, 2024 (online 2023). First definitive record of Acrocanthosaurus
(Theropoda: Carcharodontosauridae) in the Lower Cretaceous of eastern
North America. Cretaceous Research. 157, 105814.
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 catalog 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. Beiträge 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" Means, 1990
"C." potens (Lull, 1911) Molina-Perez and Larramendi,
2019
= Creosaurus potens Lull, 1911
= Dryptosaurus potens (Lull, 1911) Gilmore, 1920
Late Aptian, Early Cretaceous
Arundel Formation, Washington DC, US
Holotype- (USNM 3049) (7-10 m) proximal caudal centrum (140.0 mm)
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. Gilmore (1920) referred "The median portion of a very
large ungual (No. 8505 U.S.N.M.) collected by Arthur Bibbins from the
Arundel formation near Contee, Maryland" to this taxon "on account of
its great size". However due to its low curvature it is here
provisionally assigned to Ornithomimosauria.
When discussing the potens
specimen in a popular magazine article, Means (1990) wrote "For the
moment the beast is labeled a "dryptosaurus," but it may prove to be a
unique member of the dinosaur family. Local paleontologist Peter Kranz
would like it to be known as "capitalsaurus," Washington's very own
giant reptile." Kranz (1998) latere 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 1998
publication,
Means is discussing "a large vertebra and other bone fragments of a
twenty-foot-long carnivorous dinosaur" discovered in Washington DC in
1898 and Kranz (pers. comm. 8-6-2005) informs 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
establish the name "Capitalsaurus"
potens
in the literature, the genus 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.
Means, 1990. The great outdoors. The Washingtonian. 25(7), 124-160.
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
Cretacéo Argentino. Anales del Museo de La Plata (series 3). 3, 1-196.
Olshevsky, DML 1999. https://web.archive.org/web/20200720012936/http://dml.cmnh.org/1999Nov/msg00507.html
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 Ignère) 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 Rémada: 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 Ignère, 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 Ignère 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. Découverte de Dinosauriens,
associés à une faune de Reptiles et de Poissons, dans le Crétacé
inférieur de l'Extrême Sud tunisien [Discovery of dinosaurs associated
with a reptile and fish fauna in the Lower Cretaceous of extreme
southern Tunisia]. Comptes Rendus de l'Académie des Sciences à 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. Géologie et Paléontologie du Gisement de Gadoufaoua
(Aptien du Niger) [Geology and Paleontology of the Gadoufaoua Locality
(Aptian of Niger)]. Cahiers de Paléontologie, 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 Wüsten Ägyptens. II. Wirbeltierreste der
Baharije-Stufe (unterstes Cenoman). 13. Dinosauria. Abhandlungen der
Bayerischen Akademie der Wissenschaften
Mathematisch-naturwissenschaftliche Abteilung, Neue Folge. 22, 1-79.
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.
"Mangahouanga"
Molina-Perez and Larramendi, 2019
Campanian-Maastrichtian, Late
Cretaceous
Mangahouanga Stream, Mata Series,
North Island, New Zealand
Material- (NZGS CD1) incomplete
mid caudal vertebra (~50 mm)
Comments- Molnar (1980)
described this vertebra as a probable theropod, further stating that
"The proximal caudals of Poikilopleuron
bucklandii
... closely approach this, with elevated transverse processes and an
inclined spine located far posteriorly" unlike the less inclined neural
spine in Elaphrosaurus and Allosaurus. He believed "The
Mangahouanga vertebra would represent a spine placed farther back than
the P . bucklandii
caudals, but otherwise are very similar having even the ventral
sulcus", but stated "it is obvious that a single, incomplete vertebra
is hardly a substantial basis for an identification, and although I
think it unlikely, it is possible that the vertebra is of an
ornithopod." It was most recently briefly reviewed by Agnolin et
al. (2010) who stated "This material is phylogenetically uninformative
and is considered here as Theropoda indet." Molina-Perez and
Larramendi (2019) listed it under Megaraptora without rationale,
although this would make sense biogeographically. It is retained
as Averostra here pending further study.
Lindsay's (2001) Spanish translation of his 1991 popular book states
"Este hueso de la cola es suficiente para indicarnos que el dinosaurio Mangahouanga
fue probablemente un predador, un terópodo" next to an illustration of
NZGS CD1 in posterior view. While the italicization and a naive
reading might suggest "... the dinosaur Mangahouanga
was ..." and thus qualify this as a nomen nudum, the reversed
adjective-noun order in Spanish leads to a perfect translation of the
original 1991 English version's sentence "This
tailbone is enough to tell us that the Mangahouanga dinosaur was
probably a predator, a theropod." Indeed, Mangahouanga is
correctly indicated as a locality on the New Zealand map next to the
illustration with the specimen merely labeled "Theropod
vertebra". Thus Lindsay never intended Mangahouanga to be a genus
name, and while the translator of the Spanish version or an editor
apparently confused it for a genus name and italicized it, the text
doesn't read as if it's a genus. Thus the accidental
italicization of a word would not count as a nomen nudum here, except
that
Molina-Perez and Larramendi (2019) explicitly listed ""Mangahouanga"
(n.n.)" in their list of "Megaraptora similar to Megaraptor",
referencing both NZGS CD1 and Molnar, 1980. They say "its name
was created by accident. Lambert 1983", but while the specimen was
discussed on page 237 of that book as "Late Cretaceous creature from
New Zealand's North Island" and "probably a theropod", it was never
even accidentally named there and the Mangahouang site isn't even
referenced. Molina-Perez (pers. comm., 10-2021) indicates this
was a mistake and he was thinking of Lindsay (2001), but this does make
Molina-Perez and Larramendi (2019) the first reference to explicitly
use "Mangahouanga" as a name, so they are the authors credited
here. It's not listed as "Lambert, 1983 vide Molina-Perez and
Larramendi, 2019" because they never explicitly tie Lambert to the
name, with their format more logically (though incorrectly) implying
Lambert was the one to say its name was created by accident. In
any case "Mangahouanga" is a nomen nudum, as it was not stated to be
new (ICZN Article 16.1- "Every new name published after 1999 ... must
be explicitly indicated as intentionally new"), listed as a nomen nudum
(Article 11.5- "a name must be used as valid for a taxon when
proposed") and lacks a species (Article 13.3- " every new genus-group
name published after 1930 ... must ... be accompanied by the fixation
of a type species in the original publication").
References- Molnar, 1980. A
dinosaur from New Zealand. In Cresswell and Vella (eds.). Gondwana
Five: Selected Papers and Abstracts of Papers Presented at the Fifth
International Gondwana Symposium. A. A. Balkema. 91-96.
Lambert, 1983. A Field Guide to Dinosaurs. Avon Books. 256 pp.
Lindsay, 1991. The Great Dinosaur Atlas. Dorling Kindersley Limited. 64
pp.
Lindsay, 2001. Atlas Visual de los Dinosaurios. Editoral Diana. 63 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.
Molina-Perez and Larramendi, 2019. Dinosaur Facts and Figures: The
Theropods and Other Dinosauriformes. Princeton University Press. 288
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 catalogd 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. catalog 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 für Geologie und
Paläontologie - Abhandlungen. 280(2), 215-219.
"Megalosaurus" insignis
Eudes-Delongchamps and Lennier vide Lennier, 1870
= Streptospondylus insignis (Eudes-Deslongchamps and Lennier
vide Lennier, 1870) Depéret and Savornin, 1928
= Erectopus insignis (Eudes-Deslongchamps and Lennier vide
Lennier, 1870) Stromer, 1931
Early Kimmeridgian, Late Jurassic
Saint-Adresse, Cap de La Hève, 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 Crèche (Musée de
Boulogne coll.), three teeth from Portel (Beaugrand coll.), a sacral
fragment and pedal ungual from Châtillon (Musée 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 Châtillon
(Musée 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 espèce de Chélonien
fossile d'un genre nouveau, trouvé dans la craie du cap la Hève par M.
Lennier, du Havre, et décrit par M. A. Valenciennes. Compte Rendu
des Séances de l'Académie des Sciences. 46(8), 317-322.
Lennier, 1870. Études Géologiques et Paléontologiques 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 Eugène Costey, Havre. 1-245.
Sauvage, 1874. Mémoire 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]. Mémoires de
la Société Géologique de France, série 2. 10(2), 1-57.
Lennier, 1887. Études paléontologiques. Description des fossiles du Cap
de la Hève. Bulletin de la Société Géologique de Normandie. 12, 17-98.
Lydekker, 1888. catalog 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 supérieur du
Boulonnais. Bulletin de la Société Géologique de France, 3e serie. 22,
465-470.
Sauvage, 1898. Les Reptiles et les Poissons des terrains Mésozoïques du
Portugal [The reptiles and fishes from the Mesozoic terrains of
Portugal]. Bulletin de la Société Géologique de France, 3e série. 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.
Depéret and Savornin, 1928. La faune de Reptiles et de Poissons albiens
de Timimoun (Sahara algérien) [The Albian reptile and fish fauna of
Timimoun (Algerian Sahara)]. Bulletin de la Societé Géologique de
France, 4e série. 27, 257-265.
Stromer, 1931. Ergebnisse der Forschungsreisen Prof. E. Stromers in den
Wüsten Ägyptens. II. Wirbeltier-Reste der Baharîjestufe (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)]. Mémoires de la Société
Géologique de France (Nouvelle Série). Mémoire 21(47), 1-21.
Lapparent and Zbyszewski, 1957. Les dinosauriens du Portugal. Mémoires
des Services Géologiques du Portugal, nouvelle série. 2, 1-63.
Mateus, 1999. Upper Jurassic dinosaurs from Lourinhã 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- (Musée 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
rencontrés dans le Crétacé supérieur de la Belgique [Note on the
dinosaur remains found in the Upper Cretaceous of Belgium]. Bulletin du
Musée Royal d'Histoire Naturelle de Belgique. 2, 205-221.
Dollo, 1903. Les dinosauriens de la Belgique. Comptes Rendus de l'
Académie 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 Paläoherpetologen. Extended Abstracts. Duesseldorf,
Germany. 33-34.
Windolf, 1998. Dinosaurierfunde in Niedersachsen [Dinosaur finds in
Lower Saxony]. Arbeitskreis Paläontologie 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. Mémoires des Services Géologiques du Portugal, nouvelle
série. 2, 1-63.
,
,
and ,
2016.
Systematic
review of Late Jurassic sauropods from the Museu Geológico collections
(Lisboa, Portugal).
Journal of Iberian Geology. 42, 227-250.
Mocho,
Royo-Torres, Escaso, Malafaia, de Miguel Chaves, Narváez, Pérez-García,
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
über Zirkonzwillinge. Zeitschrift der deutschen Geologischen
Gesellshaft. 36, 664-665.
Koken and Kayser, 1885. Über 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. Über 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-Brañas,
Espinosa-Chávez, Maccracken, Gutiérrez-Blando, de León-Dávila and
Ventura, 2020
P. normalensis
Serrano-Brañas, Espinosa-Chávez, Maccracken, Gutiérrez-Blando, de
León-Dávila 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-Brañas 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-Brañas,
Espinosa-Chávez, Maccracken, Gutiérrez-Blando, de León-Dávila 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 recatalogd 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 Société géologique 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
Paläoherpetologie/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. Paläontologische
Zeitschrift. 56, 265-275.
Vadet
and Rose, 1986. catalog commente des types et figures de dinosauriens,
ichthyosauriens, sauropterygiens, pterosauriens et cheloninens du Musée
d'Histoire Naturelle de Boulogne-sur-Mer.
In E. Buffetaut, Rose and
Vadet (eds.). Vértébrés Fossiles du Boulonnais. Mémoires de la Société
Académique 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 Société académique
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 Paléobiologie. 7(vol. spéc.),
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 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 catalog 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.
undescribed Averostra
(Martin and Brett-Surman, 1992 unpublished)
Late Aptian, Early Cretaceous
Dinosaur Park / Cherokee-Sanford Brick Clay Pit / Muirkirk Clay Pit
USNM 41614, Arundel Formation, Prince George's County, Maryland,
US
Material- (NHRD-AP 2010.v.014) tooth (Dinosaur Fund online)
(NHRD-AP 2013.v.678) tooth (Dinosaur Fund online)
(NHRD-AP 2014.s.094) tooth fragment (Dinosaur Fund online)
(NHRD-AP 2014.s.156) metatarsal (Dinosaur Fund online)
(NHRD-AP 2014.v.685) caudal vertebra (Dinosaur Fund online)
(USNM 437629) tooth (Martin and Brett-Surman, 1992 unpublished)
(USNM 442406) three teeth (Martin and Brett-Surman, 1992 unpublished)
(USNM 442442) tooth (Martin and Brett-Surman, 1992 unpublished)
(USNM 442521) left pedal phalanx II-2 (~64.8 mm) (USNM online)
(USNM 451937; phalanx = USNM 442510 in Martin and Brett-Surman, 1992
unpublished) (associated?) two teeth, partial pedal ungual (Carrano,
2024)
(USNM 451991) tooth (Martin and Brett-Surman, 1992 unpublished)
(USNM 466055) (associated?) incomplete tooth, proximal ?ischium (USNM
online)
(USNM 497726) partial tooth (~80x>40x~14.5 mm) (Lipka, 1998)
(USNM 497731) proximal left femur (shaft ~78 mm wide trans) (USNM
online)
(USNM 497732; = USNM 491732 of Carrano, 2024) partial tooth (?x~23x?
mm) (Frederickson et al., 2018)
(USNM 497737) (associated?) partial dorsal or proximal caudal centrum
(~52 mm tall), two vertebral fragments (USNM online)
(USNM 497750) partial tooth (Frederickson, Lipka and Cifelli, 2018)
(USNM 497751) partial tooth (Frederickson, Lipka and Cifelli, 2018)
(USNM 497752) partial tooth (?x~3.7x? mm) (Frederickson, Lipka and
Cifelli, 2018)
(USNM 497753) partial tooth (Frederickson, Lipka and Cifelli, 2018)
(USNM 497754) fragmentary bone (Frederickson, Lipka and Cifelli, 2018)
(USNM 497755) fragmentary bone (Frederickson, Lipka and Cifelli, 2018)
(USNM 497756) fragmentary bone (Frederickson, Lipka and Cifelli, 2018)
(USNM 497757) fragmentary bone (Frederickson, Lipka and Cifelli, 2018)
(USNM 497758) fragmentary bone (Frederickson, Lipka and Cifelli, 2018)
(USNM 497759) fragmentary bone (Frederickson, Lipka and Cifelli, 2018)
(USNM 534099; marked USNM 534018) proximal ?metatarsal IV (~35.6 mm
?anteropost prox) (USNM online)
(USNM 540723) posterior distal caudal vertebra (~9.3 mm tall post)
(USNM online)
(USNM 540728) partial distal caudal vertebra (~18.3 mm tall) (USNM
online)
(USNM 540764) proximal left pedal phalanx II-2 (~28.4 mm tall post)
(USNM online)
(USNM 544050) proximal left metatarsal IV (~33 mm anteropost prox)
(USNM online)
(USNM 544051) dorsal centrum (~42.1 mm) (USNM online)
(USNM 544054) right tibial shaft (midhsaft ~10.5 mm anteropost) (USNM
online)
(USNM 544055) incomplete mid caudal vertebra (~77.9 mm) (USNM online)
(USNM 544133) partial distal caudal centrum (~18.2 mm tall) (USNM
online)
(USNM 604957) incomplete tooth (USNM online)
(USNM 604986) limb bone fragment (shaft ~8.8 mm) (USNM online)
Late Aptian, Early Cretaceous
Arundel Formation, Prince George's County, Maryland, US
(USNM 8507) proximal tibia (~189 mm anteropost) (USNM online)
(USNM 425732) phalanx (~61.3 mm) (USNM online)
(USNM 535490 in part) ?ilial fragment (USNM online)
Comments- USNM 8507 is a
proximal tibia discovered in 1887 and identified by Gilmore as Dryptosaurus sp. (USNM online),
although it was not mentioned in his 1920 work on USNM theropods
referring medius and potens to the genus.
Martin and Brett-Surman (1992 unpublished) report "A small collection
of theropod teeth has been recovered from the Muirkirk clay pit
locality, and presents some interesting characteristics. The collection
includes both premaxillary and maxillary/dentary teeth, with many
crowns intact to the tip. The most consistent feature found in the
maxillary/dentary teeth is the restriction of anterrior serrations to
the top third of the tooth crown. The smaller teeth also exhibit this
feature, but have a carina reaching close to the base of the crown. In
the larger teeth the serrations of the anterrior [sic] surface
terminate abruptly, without any observable decrease in the size of the
individual denticles, and are not followed by a carina." They
determined "A brief analysis of the Arundel clay pit theropod
maxillary/dentary teeth was made using methods outlined in Farlow, et.
al. (1991)" which revealed "that the Arundel theropod teeth cluster
most closely with those of an unidentified Comanchean theropod,
dromaeosaurids, and Alectrosaurus
(in the figures by Farlow, et. al., 1991). The larger teeth also
correlate with those of an unidentified theropod from the Morrison
Formation." Unfortunately, the existing supplementary document to
Farlow et al.'s paper does not indicate the identity of the Comanchean
(Early Cretaceous), Morrison or supposed Alectrosaurus
teeth, and the pdf of Martin and Brett-Surman's paper doesn't show the
icon for Arundel material, so any further analysis is impossible.
Martin and Brett-Surman figure a pedal ungual in Figure 9 as "Theropod
ungual (USNM 442510), side and dorsal views", but 442510 doesn't
include an ungual and is actually figured on the next page incorrectly
as USNM 442521 (based on the painted label and matching the USNM online
catalog). You might assume the figure captions were switched, but
the
online catalog shows USNM 442521 is a non-ungual phalanx. The
ungual shown in Figure 9 is actually USNM 451937 (Carrano, pers. comm.
9-2024). Among specimens figured in this
work, USNM 437629, 442406 and 442442 were all found in 1989.
Frederickson et al. (2018) note regarding USNM 497750-497759 "Theropod
teeth that were highly weathered and abraded were indeterminate, based
on their incompleteness. All of the teeth are small (less than 15 mm
long) and are missing most of their enameled surface. Multiple
fragmentary and abraded metapodial ends and undiagnostic other elements
are also present in the Arundel sample. These specimens are referred to
Theropoda based on their hollow interiors and well-developed collateral
ligament pits."
Carrano (2024) stated the presence of "(1) the mesial margin of
the rounded denticles on the mesial carina is subrectangular, with a
flattened surface [character 89], and (2) the mesiodistal axis of the
mid-crown denticles on the distal carina is inclined apically from the
distal margin in lateral view [character 96]" that were found to be
locally diagnostic to the genus by Hendrickx and Mateus (2014)
"indicates that several isolated teeth from the Arundel Clay could
pertain to Acrocanthosaurus",
but "Other large teeth in the collection (USNM 8446, 451937, 491732,
497726) are not well-enough preserved for these features to be
observed, and I consider them indeterminate to clade within
Theropoda." However, according to the USNM online catalog USNM
491732
is a specimen of the crinoid Uintacrinus
socialis from the Turonian Carlile Shale of Kansas and it was a
typo for tooth USNM 497732 assigned to Acrocanthosaurus by Frederickson et
al. (2018) (Carrano, pers. comm. 9-2024). USNM 8446 was
previously referred to Allosaurus
medius by Lull (1911) and USNM 497726 was referred to Acrocanthosaurus by Lipka
(1998). USNM 451937 was found in 1991.
USNM 534099 is in the USNM online catalog as "proximal end of
metapodial" and photographed in distal view, but the broad rounded
outline of one edge of the proximal end is only a good match for the
lateral edge of metatarsal IV among theropods, but is 1.5 times larger
than Deinonychus AMNH
3015. However given a proximal depth versus total length ratio
similar to Arkansaurus it
would result in a metatarsal length of ~245 mm, well within the range
of ornithomimosaurs and thus possibly referrable to grandis.
Note the associated photo has the number 534108 painted on it, but USNM
534108 is a collection of teeth from the cricetid muskrat Ondatra annectens from the
Pleistocene of West Virginia. USNM 534099 was discovered in 1995.
Among other USNM specimens listed above, USNM 425732 was found in 1988,
442521 in 1989, 451991 in 1991, 466055 in 1992, 497731 and 487737 in
1998, 535490 in 1999, 540723 and 604986 in 2010, 604957 in 2012, and
540728, 540764 544050, 544051, 544054, 544055 and 544133 from
2010-2013. For NHRD-AP specimens, NHRD-AP 2010.v.014 was
found in
2010, 2013.v.678, 2014.s.156 and 2014.v.685 in 2013, and 2014.s.094 in
2014.
References- Lull, 1911.
Systematic paleontology of the Lower Cretaceous deposits of Maryland:
Vertebrata. Maryland Geological Survey. Lower Cretaceous, 183-211.
Martin and Brett-Surman, 1992 unpublished. A preliminary report on new
dinosaur material from the Arundel Clay (Lower Cretaceous, Late
Aptian-Early Albian) of Maryland. Smithsonian Institution. 31 pp.
Lipka, 1998. The affinities of the enigmatic theropods of the Arundel
Clay facies (Aptian), Potomac Formation, Atlantic coastal plain of
Maryland. In Lucas, Kirkland and Estep (eds.). Lower and Middle
Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History
and Science Bulletin. 14, 229-234.
Frederickson, Lipka and Cifelli, 2018. Faunal composition and
paleoenvironment of the Arundel Clay (Potomac Formation; Early
Cretaceous), Maryland, USA. Palaeontologia Electronica. 21.2.31A, 1-24.
Carrano, 2024 (online 2023). First definitive record of Acrocanthosaurus
(Theropoda: Carcharodontosauridae) in the Lower Cretaceous of eastern
North America. Cretaceous Research. 157, 105814.
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. Boletín Geológico 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. Boletín Geológico 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-Rodríguez, 2006. Terópodos del Cretácico
Superior del Estado de Coahuila, México. MS thesis, Universidad
Nacional Autónoma de México. 91 pp.
Monroy-Mújica, 2009. Microvertebrados fósiles Cretácicos Tardíos
(Campaniano Tardío) de la Formación Aguja en el Noroeste de Coahuila,
México. Tesis Licenciatura. Universidad Nacional Autónoma de México.
111 pp.
Ramirez-Valesco and Hernandez-Rivera, 2015. Diversity of Late
Cretaceous dinosaurs from Mexico. Boletín Geológico 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) (Rodríguez-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 (Rodríguez-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- Rodríguez-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.
Rodríguez-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 Cretácicos Tardíos del área de El
Rosario, Baja California, México. MS thesis, Universidad Nacional
Autónoma de México. 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. Boletín Geológico 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. Boletín Geológico y Minero.
126(1), 63-108.
unnamed possible Averostra (Lindgren, Currie, Rees, Siverson,
Lindström 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, Lindström 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, Milàn
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. catalog 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 Arqueológico-Paleontológico
Salense (ed.). Actas de las III Jornadas sobre Dinosaurios y su
Entorno. Salas de los Infantes (Burgos, España). 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. catalog 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 für Geologie und Paläontologie 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. catalog 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. catalog 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. catalog 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. catalog 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]. Mémoires des Services Géologiques
du Portugal, nouvelle série. 2, 1-63.
unnamed possible Averostra (Lapparent and Zbyszewski, 1957)
Callovian-Oxfordian, Middle-Late Jurassic
Fervença, 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]. Mémoires des Services Géologiques
du Portugal, nouvelle série. 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]. Mémoires des Services Géologiques
du Portugal, nouvelle série. 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]. Mémoires des Services Géologiques
du Portugal, nouvelle série. 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 Mésozoïques du Portugal [The reptiles and fishes from the
Mesozoic terrains of Portugal]. Bulletin de la Société Géologique de
France, 3e série. 26, 442-446.
undescribed averostran (Rauhut, 2000)
Kimmeridgian, Late Jurassic
Guimarota, Alcobaça 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
Alcobaça 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 supérieur du Boulonnais. Bulletin de la Société Géologique
de France, 3e serie. 22, 465-470.
Sauvage, 1898. Vertébrés fossiles du Portugal: Contributions à l'étude
des poissons et des reptiles du jurassique et du crétacique. Direction
des Travaux Geologiques du Portugal. Memoires. Comissão do Serviço
Geológico 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]. Mémoires des Services Géologiques
du Portugal, nouvelle série. 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]. Mémoires des Services Géologiques
du Portugal, nouvelle série. 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]. Mémoires des Services Géologiques
du Portugal, nouvelle série. 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]. Mémoires des Services Géologiques
du Portugal, nouvelle série. 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]. Mémoires des Services Géologiques
du Portugal, nouvelle série. 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]. Mémoires des Services Géologiques
du Portugal, nouvelle série. 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]. Mémoires des Services Géologiques
du Portugal, nouvelle série. 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. Vertébrés fossiles du Portugal:
Contributions à l'étude des poissons et des reptiles du jurassique et
du crétacique. Direction des Travaux Geologiques du Portugal. Memoires.
Comissão do Serviço Geológico de Portugal. 1-46.
Lapparent and Zbyszewski, 1957. Les dinosauriens du Portugal [The
dinosaurs of Portugal]. Mémoires des Services Géologiques du Portugal,
nouvelle série. 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. Vertébrés fossiles du Portugal:
Contributions à l'étude des poissons et des reptiles du jurassique et
du crétacique. Direction des Travaux Geologiques du Portugal. Memoires.
Comissão do Serviço Geológico de Portugal. 1-46.
Lapparent and Zbyszewski, 1957. Les dinosauriens du Portugal [The
dinosaurs of Portugal]. Mémoires des Services Géologiques du Portugal,
nouvelle série. 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) (Suñer, Santisteban and
Galobart, 2005)
(GAOO/CL/14) tooth (Barco and Ruiz-Omeñaca 2001a)
(LP-1) partial tooth (?x19.4x15 mm) (Suñer, Santisteban and Galobart,
2005)
(LV-1) tooth (67.5x24.7x11.1 mm) (Suñer, Santisteban and Galobart, 2005)
(MPZ01/97) proximal caudal centrum (Barco and Ruiz-Omeñaca 2001b)
(PM-1) partial pedal ungual (Suñer, Santisteban and Galobart, 2005)
tooth (Abella and Suñer, 2004)
three teeth, one bone (Suñer and Martín, 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-Omeñaca and Canudo (2003).
References- Casanovas-Cladellas, Santafé-Llopis and
Santisteban-Bové, 1993. First dinosaur teeth from the Lower Cretaceous
of Benicatazara (Aras de Alpuente, Valencia). Revue de Paléobiologie.
Special volume 7, 37-44.
Barco and Ruiz-Omeñaca, 2001a. Primeros dientes de terópodos
(Dinosauria, Saurischia) en la Formación Villar del Arzobispo
(Titónico-Berriasiense): Yacimientos Cuesta Lonsal y Las Cerradicas 2
(Galve, Teruel). Publicaciones del Seminario de Paleontología de
Zaragoza. 5, 239-246.
Barco and Ruiz-Omeñaca, 2001b. Primeros restos postcraneales de
terópodos (Dinosauria, Saurischia) en la Formación Villar del Arzobispo
(Titónico-Berriasiense): Un centro vertebral caudal del yacimiento
Carretera (Galve, Teruel). Publicaciones del Seminario de Paleontología
de Zaragoza. 5, 247-254.
Ruiz-Omeñaca and Canudo, 2003. Dinosaurios (Saurischia, Ornithischia)
en el Barremiense (Cretácico inferior) de la Península Ibérica. In
Pérez-Lorente (Ed.). Dinosaurios y otros reptiles Mesozoicos en España.
Ciencias de la Tierra. 26, 269-312.
Canudo and Ruiz-Omenaca, 2003. Los restos directos de dinosaurios
teropodos (excluyendo Aves) en Espana. In Pérez-Lorente (Ed.).
Dinosaurios y otros reptiles Mesozoicos en España. Ciencias de la
Tierra. 26, 347-373.
Abella and Suñer, 2004. Un nuevo diente aislado de terópodo del
yacimiento de El Chopo (Alpuente, Los Serranos, Valencia). Libro de
Resúmenes del II Encuentro de Jóvenes Investigadores en Paleontología.
87-88.
Suñer, Santisteban and Galobart, 2005. Nuevos restos de Theropoda del
Jurásico Superior-Cretácico Inferior de la Comarca de Los Serranos
(Valencia). Revista Española de Paleontologia. N. Extra X, 93-99.
Suñer and Martín, 2009. Un nuevo yacimiento del tránsito
Jurásico-Cretácico de Alpuente (Los Serranos, Valencia, España):
Resultados preliminares. Paleolusitana. 1, 441-447.
Gascó, Cobos, Royo-Torres, Mampel and Alcalá, 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)]. Mémoires
de la Société Géologique de France (Nouvelle Série). Mémoire 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
supérieur du Boulonnais. Bulletin de la Société Géologique de France,
3e serie. 22, 465-470.
unnamed Averostra (Sauvage, 1874)
Late Kimmeridgian, Late Jurassic
Châtillon, Pas-de-Calais, France
Material- (Musée de Boulogne coll.) two fused sacral centra
(Musée de Boulogne coll.) pedal ungual
(Musée 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. Mémoire sur les dinosauriens et les
crocodiliens des terrains jurassiques de Boulogne-sur-Mer. Mémoires de
la Société Géologique de France, série 2. 10(2), 1-57.
Sauvage, 1894. Les dinosauriens du terrain jurassique supérieur du
Boulonnais. Bulletin de la Société Géologique 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 supérieur du Boulonnais. Bulletin de la Société Géologique
de France, 3e serie. 22, 465-470.
Sauvage, 1900. catalog 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 supérieur du Boulonnais. Bulletin de la Société Géologique
de France, 3e serie. 22, 465-470.
Sauvage, 1914. catalog des reptiles jurassiques du Boulonnais. Bulletin
de la Société académique 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 Crèche, Pas-de-Calais, France
Material- (Musée de Boulogne coll.) tooth (110 mm)
(Musée de Boulogne coll.) incomplete tooth (~70 mm)
(Musée de Boulogne coll.) incomplete tooth
(Musée de Boulogne coll.) tooth (>17 mm)
Comments- These were referred to Megalosaurus insignis.
Reference- Sauvage, 1874. Mémoire sur les dinosauriens et les
crocodiliens des terrains jurassiques de Boulogne-sur-Mer. Mémoires de
la Société Géologique de France, série 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. catalog 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 trouvés
dans le Gault de l'est du bassin de Paris [Research on the reptiles
found in the Gault of the eastern Paris Basin]. Mémoires de la Société
Géologique de France, série 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 supérieur du
Boulonnais. Bulletin de la Société Géologique 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- (Musée de la Faculté 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 Crétacé du
nord-est du Bassin de Paris. Bulletin scientifique, historique et
littéraire 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
Société Géologique de France. 4, 439-442.
Sauvage, 1882a. Sur les Reptiles trouvés 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'Académie des Sciences. 94,
1265-1266.
Sauvage, 1882b. Recherches sur les reptiles trouvés dans le Gault de
l'est du bassin de Paris [Research on the reptiles found in the Gault
of the eastern Paris Basin]. Mémoires de la Société Géologique de
France, série 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- (Musée de la Faculté 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 Crétacé du
nord-est du Bassin de Paris. Bulletin scientifique, historique et
littéraire 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
Société Géologique de France. 4, 439-442.
Sauvage, 1882b. Recherches sur les reptiles trouvés dans le Gault de
l'est du bassin de Paris [Research on the reptiles found in the Gault
of the eastern Paris Basin]. Mémoires de la Société Géologique de
France, série 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 für Mineralogie, Geologie und
Paläontologie. 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
Knötschke, 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 Knötschke, 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). Paläontologische 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 für Mineralogie,
Geologie und Paläontologie - 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 left 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). A 3D model was recently uploaded to MorphoSource
(AMNH online 2024).
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.
AMNH 2024 online. https://www.morphosource.org/concern/media/000599253
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 catalog),
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 catalogd 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 für Geologie und Paläontologie -
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 catalog, 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 uncatalogd; 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
Paléobiologie. 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
Paléontologie. 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, Carpenter, Chatterjee, Ford, Khosla, Molnar and
Prasad, no Yadagiri 1979 work referencing
theropods has been located (note no theropods 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 catalogd
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. Étude paléontologique des vertébrés
du Jurassique d'El Mers (Moyen Atlas). Notes et Mémoires du Service
Géologique du Maroc. 124, 1-36.
Chabli, 1986. Données nouvelles sur un "Dinosaurien" Jurassique Moyen
du Maroc: Megalosaurus mersensis
Lapparent 1955, et sur les Megalosaurides en général. In Taquet and
Sudre (eds.). Les Dinosaures de la Chine à la France. Musée 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- (Musée 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’Académie 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, Métais,
Rage, Vullo and
Zouhri, 2020)
(FSAC-KM-A1-13) thirteen teeth (Lasseron, Allain, Gheerbrant, Haddoumi,
Jalil, Métais, 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, Métais, 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, Métais, Rage, Vullo and
Zouhri, 2020)
(FSAC-KM-B'-24) four teeth (Lasseron, Allain, Gheerbrant, Haddoumi,
Jalil, Métais, 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, Métais, 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, Métais, 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-Omeñaca, 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, Métais, 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 Faculté 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. Découverte de Dinosauriens associés à des Crocodiliens et
des Poissons dans le Crétacé inférieur de l'Atlas saharien (Algérie).
Compte Rendu Sommaire des Séances de la Société Géologique 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 Paléontologie. 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
Paléontologie. 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 Wüsten Ägyptens. II. Wirbeltierreste der
Baharije-Stufe (unterstes Cenoman). 13. Dinosauria. Abhandlungen der
Bayerischen Akademie der Wissenschaften
Mathematisch-naturwissenschaftliche Abteilung, Neue Folge. 22, 1-79.
Sereno, Dutheil, Iarochene, Larsson, Lyon, Magwene, Sidor, Varricchio
and Wilson, 1996. Predatory dinosaurs from the Sahara and Late
Cretaceous faunal differentiation. Science. 272(5264), 986-991.
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 maëstrichtiani 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. Géologie et Paléontologie du Gisement de Gadoufaoua
(Aptien du Niger). Cahiers de Paléontologie, 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 Colección Aznárez 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-Zähne aus der Kreide von Uruguay.
Centralblatt für Mineralogie, Geologie und Paläontologie, Abteilung B:
Geologie und Paläontologie. 1934(4),183-189.
Mones, 1997. Los vertebrados mesozoicos del Uruguay y sus relaciones
con los de áreas vecinas. In Arroyo Cabrales and Polaco (eds.).
Homenaje al Profesor Ticul Álvarez. Mexico D.F., Mexico: Instituto
Nacional de Antropología e Historia, Colección Científica. 357, 205-222.
Soto and Cambiaso, 2006. Reinterpretación de los dientes de dinosaurios
de la Formación Guichón: Iguanodontes basales y terópodos no
ornitomímidos. 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 Ciências.
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, Cretáceo Superior
continental da Bacia do Paraná, com ênfase em sua fauna de amniotas.
PhD thesis, Universidade Federal do Rio de Janeiro. 493 pp.
Franco, 1999. Dentes de teropodomorfos do Cretáceo Superior da Bacia do
Paraná. Análise em Microscopia Eletrônica 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 Ciências. 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 Geológico, São 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,
Cretáceo Superior
continental da Bacia do Paraná, com ênfase em sua fauna de amniotas.
PhD thesis, Universidade Federal do Rio de Janeiro. 493 pp.
Franco, 1999. Dentes de teropodomorfos do Cretáceo Superior da Bacia do
Paraná. Análise em Microscopia Eletrônica 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 Ciências. 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 Marília
Formation (Late Maastrichtian) at the paleontological site of
Peirópolis in Minas Gerais State, Brazil. Revista Brasileira de
Geociências. 42(2), 323-330.
undescribed averostran (Pol and Rauhut, 2012)
Middle Toarcian, Early 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 Cretacéo 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
Cretacéo 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
Cretacéo Argentino. Anales del Museo de La Plata (series 3). 3, 1-196.
Olshevsky, DML 1999. https://web.archive.org/web/20200720012936/http://dml.cmnh.org/1999Nov/msg00507.html
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
asociación de dientes vertebrados para la Formación Bajo de la Carpa
(Santoniense, Cretácico Superior) en Rincón de los Sauces, Neuquén,
Argentina. Revista Española de Paleontología. 30(2), 223-238.
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 Paleontología de Vertebrados, resumenes.
Ameghiniana. 52(4) suplemento, 13.