Ceratosauria
As I said before, I don't
support the
idea that the "neoceratosaurians" and the coelophysoids form a natural group,
Ceratosauria. Instead, the "neoceratosaurians" form my
Ceratosauria. Indeed, some workers are suggesting that this reduced
Ceratosauria may be paraphyletic, with Ceratosaurus and the abelisauroids
as basal relatives to the tetanurans.
Ceratosaurians are most commonly found in the continents that made up
Gondwana (South America, Africa, Antarctica, Australia, India, and Madagascar), in late Cretaceous-age rocks, but one of the best known members, Ceratosaurus
itself, is a North American theropod from the Late Jurassic. Ceratosaurians, while
not as diverse as the tetanurans, produced many interesting taxa, including the bizarre
carnotaurine abelisaurids and the small, large-clawed (at least in some cases)
noasaurids.
<--Ceratosauria
|--Ceratosauridae
| `--Ceratosaurus
|--Elaphrosaurus
`--+--Spinostropheus
`--Abelisauroidea
|--Berberosaurus
`--+--Noasauridae
| |--Deltadromeus
| |--Laevisuchus
| |--Masiakasaurus
| `--Noasaurus
`-->Abelisauridae
Ceratosauria and Ceratosauridae: Sereno, Wilson, and Conrad (2004)'s new
analysis puts Ceratosaurus as more derived than Elaphrosaurus, but
a fair amount of other analyses have suggested the opposite, or put Ceratosaurus or the
abelisauroids to be closer to the tetanurans, so I'm not going to put one or the
other of these genera closer to the abelisauroids yet.
| Taxon or Taxa: | Time\Place: | Comments: | |
| Ceratosaurus: Marsh, 1884 | C. nasicornis (type) Marsh, 1888 | Kimmeridgian (LJ) of Colorado and Utah; Kimmeridgian (LJ) of Portugal (?C. dentisulcatus) |
This Morrison theropod is famous for its tall nose horn
and less prominent preorbital horns. It also possesses a row of bony nodules down
the spine, which is probably a holdover from the earliest dinosaurs and their
ancestors. To go along with these ornaments, it has a moderate
fin-back. Ossified tendons are found in at least some individuals. The hand
has four metacarpals. The skull has
particularly large teeth, but is
lightly-constructed. |
| C. dentisulcatus Madsen and Wells, 2000 | |||
| C. magnicornis Madsen and Wells, 2000 | |||
| Elaphrosaurus: Janensch, 1920 | E. bambergi (type) Janensch, 1920 | Kimmeridgian (LJ) of Tanzania |
If a skull was known here, it would help
measures immensely. As it stands, Elaphrosaurus is one of the
shortest of known theropods at the hip for its length. It was once considered an ornithomimid
ancestor, but I find that hypothesis hard to believe. The hip structure, for
example, is so different between Elaphrosaurus and a true ornithomimid that it
casts serious doubt on such a proposition. There may be both French and North American ("E. philtippetorum" for the latter, based on a humerus, and a tibia and metatarsal are also apparently referable to this genus) species around at about the same time, as shown by similar isolated remains in those places. The position of Elaphrosaurus is somewhat fuzzy, but new studies find it closest to the abelisauroids, possibly related to the noasaurids. In some ways, it is little changed from the coelophysoids. |
| "E. philtippetorum" (N.N.) Pickering, 1995 | Kimmeridgian (LJ) of Colorado | ||
| Spinostropheus gautieri Sereno, Wilson, and Conrad, 2004 (originally Elaphrosaurus gautieri Lapparent, 1960) | ?Hauterivian (EK) of Niger | Once one of the two dubious EK species of Elaphrosaurus, Spinostropheus has now been recognized as a basal ceratosaurian (more derived than Elaphrosaurus, though). Vertebral remains in the original type have been tied to a long stretch of presacral verts found at the same horizon as Jobaria and Afrovenator. The whole animal would have been on the order of 4 m long. | |
Ceratosauria i.s.: Among other recent discoveries, a new, fairly large ceratosaurian is known from material from the Aptian of Argentina.
| Taxon or Taxa: | Time\Place: | Comments: | |
| "Ceratosaurus" roechlingi (N.D.) Janensch, 1925 | Kimmeridgian (LJ) of Tanzania | "C." roechlingi is based on some odds and ends (quadrate, fibula, caudals, astragalus) of a ceratosaurian. | |
| "Labrosaurus": | "L." stechowi (N.D.) Janensch, 1920 | Kimmeridgian (LJ) of Tanzania | This is a tooth taxon with some similarities to Ceratosaurus, and has sometimes been assigned to it. |
| "L." sulcatus Marsh, 1896 | Kimmeridgian (LJ) of Colorado | Again, this tooth may belong to Ceratosaurus. | |
| ?Lukousaurus yini Yang, 1948 (?N.D.) | Hettangian-Sinemurian (EJ) of China | This beast, based on a partial skull with distinctive preorbital horns, may not be a dinosaur. If it is, the form of its antorbital fenestra shows some resemblance to Ceratosaurus' and the abelisaurids'. | |
| "Megalosaurus": | "M." bredai (N.D.) Seeley, 1883 | Maastrichtian (LK) of the Netherlands | Otherwise known as Betasuchus, this theropod, described as similar to Sarcosaurus, was once considered to be Europe's only ornithomimid. |
| "M." ingens (?N.D.) Janensch, 1920 (?Ceratosaurus) | Kimmeridgian (LJ) of Tanzania | This theropod is based on a whopping big tooth, suggesting an owner that weighed around five metric tons. It is similar to Ceratosaurus teeth. | |
| "M." meriani (N.D.) Greppin, 1870 | Tithonian (LJ) of Switzerland | "M." meriani is based on a Ceratosaurus-like tooth, mostly interesting for its time and place (big theropods are rare for the LJ of Europe, except in Portugal). | |
Ceratosauridae i.s.:
| Taxon or Taxa: | Time\Place: | Comments: |
| Genyodectes serus Woodward, 1901 | ?Aptian-?Albian (?EK) of Argentina | Genyodectes is based on part of the front end of some theropod jaws. Sometimes it has been suggested that it , may be synonymous with the better-known Abelisaurus, but new study indicates that it was actually a ceratosaurid. More surprising, given the years that it was tossed off as theropod scrap, is that when someone (Rauhut) went to look at the material, it turned out to be distinctive. The teeth are closely packed, and the maxillary teeth are flattened side-to-side and very tall (and quite a bit longer than the pmx teeth). |
Abelisauroidea: The abelisauroids were primarily a late Cretaceous
Gondwanan group. They are mostly large and some
members grew exotic headgear (Carnotaurinae). At least some members are known to
have reduced their hands and arms to almost nothingness.
Several undescribed abelisauroids
have been mentioned in the literature recently, including a late Cenomanian
(LK) form from Argentina.
| Taxon or Taxa: | Time\Place: | Comments: |
| Berberosaurus liassicus Allain, Tykowski, Aquesbi, Jalil, Monbaron, D. Russell, and Taquet, 2007 | Pliensbachian-Toarcian (EJ) of Morocco | Not the same as the EJ Moroccan theropod mentioned by Gregory Paul back in 1988's Predatory Dinosaurs of the World (that's something else, under prep), Berberosaurus is based on a partial postcranial skeleton of a subadult comparable in size to Dilophosaurus or Elaphrosaurus. Most of the remains pertain to the hindlimb or vertebrae. It is the earliest known abelisauroid, and indicates an early diversification for the group. Tazoudasaurus was a contemporary. |
Abelisauroidea i.s.:
| Taxon or Taxa: | Time\Place: | Comments: |
| ?Chuandongocoelurus primitivus (?N.D.) He, 1984 | Bathonian-Callovian (MJ) of China | Rather obscure, Chuandongocoelurus is based on a partial skeleton, probably chimeric as the presacral verts appear to be too large for the rest of the skeleton, of a subadult individual. It may be close to Elaphrosaurus. |
| Genusaurus sisteronis Accarie, Beaudoin, Dejax, Fries, Michard, and Taquet, 1995 | Albian (EK) of France | Based on a partial skeleton, this taxon is unusual in its combination of young geologic age and conservative features, although some features suggest that it might be related to Carnotaurus. |
| Ilokelesia aguadagrandensis Coria and Salgado, 2000 | Cenomanian-early Turonian (LK) of Argentina | This is a new, incomplete abelisauroid with a very interesting name, known from skull fragments, some vertebrae, and hindlimb material first found before at least 1991. It may be a basal sister group to Abelisauridae+Noasauridae. The caudal verts' transverse processes are T-shaped. |
| Ozraptor subotaii (?N.D.) Long and Molnar, 1998 | Bajocian (MJ) of Australia | As its name makes clear, this is an Australian predator. Little else is known at this time; although some workers have suggested dromaeosaurid affinities for the type distal tibia, new research indicates that it actually was an early abelisauroid, becoming the first Australian abelisauroid and the earliest known example. |
| Pycnomenosaurus nevesi Kellner and Campos, 2002 | ?Santonian (LK) of Brazil | This is a new abelisaurian from Brazil, apparently the first such named member of the clade from that country. It is based on rather scrappy remains, including a tibia, parts of a fibula and pubis, and a handful of verts, uncovered in the early 1950s, and may be an abelisaurid. |
| Tarascosaurus salluvicus (?N.D.) Le Loeuff and Buffetaut, 1990 | early Campanian (LK) of France | Tarascosaurus, a rare non-Gondwanan abelisauroid, may be related to Abelisaurus. It is based on a femur and two dorsals, which may not belong to the same taxon. |
| Xenotarsosaurus bonapartei Martinez, Gimenez, Rodriguez, and Bochatey, 1987 | late Cenomanian- early Turonian (LK) of Argentina | This animal is based upon most of a hindlimb and some verts. Although often tossed off as an abelisaurid, its scanty type material lacks important elements for diagnosis. |
Noasauridae: This family is composed of some of the smaller ceratosaurians (ignoring Deltadromeus and the possible member Bahariasaurus). It was long thought that at least some of them paralleled sickle-clawed coelurosaurians like the dromaeosaurids and troodontids by developing a sickle claw, but it is more likely that the "foot" claw actually went to a finger. You know, the same thing has happened to Baryonyx, Megaraptor, Fukuiraptor (all once considered to be dromaeosaurids, however briefly), and Dryptosaurus (if I remember correctly; I'll have to check), and it probably won't be the last time; we paleontologists just like to put claws anywhere but the hands, it seems, even though the more we look, the more it seems like big manual claws aren't that uncommon among dinosaurs. Heck, even Iguanodon's thumb spike was first interpreted as a nasal horn!
| Taxon or Taxa: | Time\Place: | Comments: |
| Deltadromeus agilis Sereno, Dutheil, Iarochene, Larrson, Lyon, Magwene, Sidor, Varricchio, and Wilson, 1996 (?Bahariasaurus) | early Cenomanian (LK) of Egypt and Morocco | Known from a partial postcranial skeleton and other
postcranial bits, Deltadromeus is considered to be one of the swiftest
theropods. It also got fairly big, as elements once assigned to the
possibly closely-related Bahariasaurus that actually belong to this animal bear
out. Its coracoids (in the shoulder girdle) are unusually large. Deltadromeus started out its public career as the most complete basal coelurosaurian. However, new evidence suggests that, instead, it is the largest known noasaurid. |
| Laevisuchus indicus Huene and Matley, 1933 | Maastrichtian (LK) of India | This theropod, based on three cervicals, has at times been considered a "coelurid." However, recent information indicates this is a small abelisaurian. |
| Masiakasaurus knopfleri Sampson, Carrano, and Forster, 2001 | mid Maastrichtian (LK) of Madagascar | Quite possibly the first dinosaur to be described in the new millennium, Masiakasaurus is also garnering fame from its specific name, honoring Mark Knopfler of the band Dire Straits. It is a rather bizarre little creature, known from several individuals and roughly 40 percent of the skeleton, including the odd jaws: this creature had unusual forward-jutting teeth, like a fishing animal. There may be two morphs ("robust" and "gracile"). |
| Noasaurus leali Bonaparte and Powell, 1980 | late Campanian-Maastrichtian (LK) of Argentina | This small theropod was long thought to be unusual in that it seemed to independently come up with a sickle-claw system similar to that of the dromaeosaurids and troodontids, without belonging to either of those two groups by virtue of its basal skull and very different mechanism for the claw's action. Well, there may be a good reason for why the mechanism is different: the foot claw may be a hand claw, given that Masiakosaurus has enough foot remains to show it lacked such a claw. |
Noasauridae i.s.:
| Taxon or Taxa: | Time\Place: | Comments: |
| ?Bahariasaurus ingens Stromer, 1934 (?N.D.) (?Deltadromeus) | ?Albian-early Cenomanian (EK-LK) of Egypt and ?Niger | Bahariasaurus, like Spinosaurus and Carcharodontosaurus, had its type material destroyed in WWII. Not all of the material assigned to it belongs to it, including some material that has since been referred to Deltadromeus, which may well be synonymous. |
| ?Coeluroides largus (N.D.) Huene and Matley, 1933 (?Jubbulpuria) | Maastrichtian (LK) of India | Originally considered a "coelurosaur", hence the name, this dinosaur is based on primitive caudals with delta-shaped transverse processes, originally described as dorsals. Ironically, it seems to have been a large theropod, despite how its name attempts to affiliate it with Coelurus; also ironically, its caudals are said to resemble those of tiny Ligabueino. It could be the same as Jubbulpuria. |
| ?Jubbulpuria tenuis (N.D.) Huene and Matley, 1933 (?Coeluroides) | Maastrichtian (LK) of India | This theropod has been called a "coelurid" in the past simply because of its small size. What it really is, is hard to determine from its meager remains, although its caudals appear to belong to something like Ligabueino. |
| ?Ligabueino andesi Bonaparte, 1995 | Barremian (EK) of Argentina | On of the earliest known and smallest abelisauroids, Ligabueino is known from partial postcranial remains. |
| ?Velocisaurus unicus Bonaparte, 1991 | Santonian (LK) of Argentina | This fleet animal is known from a partial hindlimb. The toe proportions have led some to suggest it may have fed like a chicken, through scratching at the ground. |
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