Ornithischia
The ornithischians are an amazingly
diverse group of dinosaurs linked by their possession of, among other things: at least
five sacrals (uncertain in most basal forms); pubis rotated backwards, with a
prepubic process of varying development pointing forward; no gastralia; a
palpebral bone in the orbit (prong of bone cutting across outside of eye socket to varying
degrees); ossified tendons at least above the sacral region (lost for tail mobility in stegosaurians); no phalanges on digit V of the foot; a
recession in the jaw margins for "cheeks;" and finally, and to my mind most
importantly, possession of an extra bone at the beak-end of the lower jaw, called a
predentary, which is where the name Predentata, sometimes used for this group, comes from. Ornithischians
were
basally small, bipedal ?herbivores (it is suggested that some ornithischians, particularly
those with heterodonty [differing teeth], and the basalmost forms, were omnivorous, including heterodontosaurids
with their fangs and some marginocephalians; some of the basal ornithischians,
including an unnamed tooth form, may have been carnivorous), who by the MJ had produced an astonishing
variety of body plans, including the spiky quadrupedal stegosaurians, the heavy-set
armored ankylosaurians, and the small, fast basal ornithopods known informally as hypsilophodonts. By the
end of the Cretaceous, more had arrived, like the bulky iguanodonts
and their descendants, the hadrosaurids, the horned
ceratopsians, and the bone-headed pachycephalosaurians.
Although ornithischians derive from bipedal ancestors, almost all of
them show at least some ability to walk on all fours. Ornithischians were primarily
low browsers. Unlike the other main group of classic dinosaurian herbivores, the sauropodomorphs, ornithischians display a wide variety of
bodily features that appear to have been used for display and recognition, like the
vertebral fins and headgear of hadrosaurids and some iguanodonts.
Large-scale ornithischian relationships have been stable and
mostly unquestioned for the last 20 or so years, except for minor quibbles about
where heterodontosaurids go, the position of Scelidosaurus vis-à-vis the
ankylosaurians and stegosaurians, and if hypsilophodontids form a natural
group. There are some changes coming up, though, led by Richard Butler's
work on basal ornithischians and new considerations on all of those tooth taxa
from North America. The main changes will affect the basal taxa and the
hypsils. This below is a version of the most current incarnation, which
isn't too dissimilar to what came before except for Lesothosaurus and the
heterodontosaurids. Using Cerapoda and Ornithopoda may get harder, though.
<--Ornithischia
|--Pisanosaurus
|--Heterodontosauridae
| |--Abrictosaurus
| `--Heterodontosaurinae
| |--Heterodontosaurus
| `--Lycorhinus
`--+--Eocursor
`--Genasauria
|--Thyreophora
| |--Scutellosaurus
| `--+--Emausaurus
| `--+--Scelidosaurus
|
`--Eurypoda
|
|-->Ankylosauria
|
`-->Stegosauria
`--Neornithischia
|--Lesothosaurus
`--+--Stormbergia
`--+--Agilisaurus
`--+--Hexinlusaurus
`--+--Othnielosaurus
`--Cerapoda
|-->Ornithopoda
`--Marginocephalia
|-->Pachycephalosauria
`-->Ceratopsia
Ornithischia to Genasauria: Unlike theropods and sauropodomorphs, there is little good material for basal ornithischians. Most of what was thought to be known for them were distinctive teeth, but after the Revueltosaurus affair these may not even be dinosaurian (check here to find if your favorite tooth taxon has been banished).
| Taxon or Taxa: | Time/Place: | Comments: |
| Pisanosaurus mertii Casamiquela, 1967 | early Norian (LTr ) of Argentina | This small animal is based on a partial skeleton and skull that show an ur-ornithischian. It has been allied to heterodontosaurids, "fabrosaurids," and hypsilophodonts, but is much too primitive; the condition of the sacrum and direction of the pubis are not certain (but it seems to be propubic, like classic saurischians). |
| Eocursor parvus Butler, Smith, R.M.H., and Norman, 2007 | Norian (LTr) of South Africa | Based on a partial skeleton including most of the lower jaw, portions of vertebrae, and much of the girdles and limbs minus the forearm and some of the hands, feet, and coracoid, Eocursor is the best-represent early ornithischian. It clearly is opisthopubic, but unfortunately the tip of the lower jaw is poorly preserved, so the presence or absence of a predentary is unknown. The animal was about 1 meter long, but given the preservation of the verts, I'd guess it wasn't fully grown. It looks a lot like a Lesothosaurus with big heterodontosaurid hands. |
Ornithischia to Genasauria i.s.: Years ago, when all ornithischians were thought to have sprung from some sort of ornithopod or another, most of these animals would have been known as the most basal ornithopods, the "fabrosaurids." Something like these animals is known from the Late Triassic Lower Elliot Formation of South Africa.
| Taxon or Taxa: | Time/Place: | Comments: |
| Alocodon kuehnei (N.D.) Thulborn, 1973 | late Callovian (MJ) of Portugal | This animal is based on teeth which show some resemblance to "othnieliid" teeth, so it may belong in that "family". Alternatively, it could be a thyreophoran. |
| Fabrosaurus australis (N.D.) Ginsburg, 1964 (?Lesothosaurus) | Hettangian (EJ) of Lesotho | Based on a partial dentary, this animal may well be the same as Lesothosaurus. Fabrosaurus in fact is the name usually used for restorations of what is actually Lesothosaurus in older dinosaur books. |
| Gongbusaurus shiyii (N.D.) Dong, Zhou, and Zhang, 1983 | Oxfordian (LJ) of China | This is an indeterminate tooth taxon, somewhere close to Genasauria. It could be an ankylosaurian. |
| "Gongbusaurus" wucaiwanensis Dong, 1989 | Oxfordian (LJ) of China | This animal is known from a partial skeleton that may or may not belong in the same genus as the tooth-taxon Gongbusaurus. It appears to be another "fabrosaurid"-type ornithischian, but is in need of restudy. |
| Taveirosaurus costai (N.D.) Antunes and Sigogneau-Russell, 1991 | late Campanian-early Maastrichtian (LK) of Portugal | Based on teeth, this may be a small, late, basal European pachycephalosaurian, or a juvenile nodosaurid. |
| Trimucrodon cuneatus (N.D.) Thulborn, 1973 | Kimmeridgian (LJ) of Portugal | This is an indeterminate basal ornithischian based on teeth. It could be a heterodontosaurid, and may be close to Echinodon. |
| Xiaosaurus dashanpensis Dong and Tang, 1983 | Bathonian-Callovian (MJ) of China | Possibly a basal ornithopod (although some have suggested that it could be a very basal marginocephalian, which wouldn't be all that different from a very basal ornithopod), this animal's remains are scrappy (the type is a partial maxilla, two cervicals, four caudals, and a hindlimb; a referred specimen is two teeth, a dorsal, two sacrals, a rib, a femur, and a phalanx; neither specimen can be found at this time) and just distinctive enough to stave off the dreaded nomen dubium. In the past, it has been mostly thought of as a "fabrosaurid." At this point, it's safest to consider it among the uncertain basal ornithischians, probably sorting out somewhere around Agilisaurus and Hexinlusaurus. More material is needed. |
Heterodontosauridae: Heterodontosaurids are known mostly from skull material, with distinctive tusks found in most individuals. They have been wild-cards, given the paucity of other early ornithischians to compare them to, and have been put with the ornithopods, or closer to the marginocephalians due to their jugal (cheek-area) bosses and tusks, the latter of which are also known in pachycephalosaurians, and some pelvic and hand details. The most recent hypothesis is that they are a very basal offshoot of Ornithischia, which I'm using here. This is still a hot issue, or at least it would be if ornithischians were charismatic predators. Heterodontosaurid material may also be known from the Sinemurian-Pliensbachian (EJ) of the southwest U.S., the Norian (LTr) of Argentina, and the LTr of Europe.
| Taxon or Taxa: | Time/Place: | Comments: |
| Abrictosaurus consors Hopson, 1975 (Thulborn, 1974 [originally Lycorhinus consors]) | Hettangian (EJ) of Lesotho and South Africa | Abrictosaurus is known as the "tuskless"
heterodontosaurid, although it gained that reputation through a mislabeled
figure. Its type actually does have tusks; two in the premaxilla are
known, although the prominent dentary tusk is indeed lacking. It may be that it is actually a juvenile (although juvenile Heterodontosaurus had tusks), as suggested by its steeply-sloping face, or female of another taxon. However, it also may have unusually small hands compared to Heterodontosaurus, which doesn't seem like a growth or sex difference. |
Heterodontosauridae i.s.:
| Taxon or Taxa: | Time/Place: | Comments: |
| Echinodon becklesii Owen, 1861 | early Berriasian (EK) of England | Echinodon (sometimes seen as Saurechinodon in older references) has long been an enigmatic ornithischian. At various times, it has been considered a "fabrosaurid," basal ornithischian, and basal thyreophoran (based on possible armor, which appears to belong to a turtle's limbs). It instead appears to be a heterodontosaurid. It is based on jaw remains, with many other jaws and teeth referred. One or two maxillary teeth may be caniform. |
| Fruitadens haagarorum Butler, Galton, Porro, Chiappe, Henderson, and Erickson, 2009 | early Tithonian (LJ) of Colorado | At long last, the "Fruita Echinodon"
is unveiled to the public. Those of you who have followed the
popular and technical literature since the 1980s will be aware of numerous
brief references to a "possible second species of Echinodon"
from the Fruita area of Colorado. As advertised, it turns out to be
a basal-ish heterodontosaurid. Unlike some other heterodontosaurids,
it lacked upper "canines," and had replacement teeth. It
is particularly known for its diminutive size, measuring 65-75 cm long and
less than a kilogram (1 to 2 lbs estimated) at about 5 years of age.
Material known includes jaw remains, a femur, a tibia, a fibula, and a partial humerus, from at least four individuals of varying stages of growth. It and fellow late heterodontosaurids Echinodon and Tianyulong seem to have hit upon the strategy of being generalists, and were less specialized in the skull than Heterodontosaurus. Fruitadens has even been proposed as an omnivore. |
| Geranosaurus atavus (N.D.) Broom, 1911 | Sinemurian (EJ) of South Africa | Geranosaurus is an indeterminate heterodontosaurid based on jaws. |
| Tianyulong confuciusi X.-T. Zheng, H.-L. You, X. Xu, and Z.-M. Dong, 2009 | early LJ of China | Tianyulong has attracted quite a bit of attention for its integumentary impressions, including short fuzzy stuff over the body and longer filaments on the tail. It's also noteworthy in being the first Asian heterodontosaurid that hasn't turned out to be a sphenosuchian croc. It is known from the partial skeleton of a subadult about 70 cm long, including a partial skull and jaws, some presacrals, a good-sized chunk of the tail, a scapula, the upper arms and an ulna, part of the hips, and much of the legs. The partial skull is like that of Heterodontosaurus in general profile, with no antorbital fenestra. The tail has ossified tendons, but they look oddly un-ornithischian. Since I don't mind looking like an idiot (you have many years of this site for evidence), I'll say that the tail looks vaguely dromie. Of course, early heterodontosaurids lacked ossified tendons, so there's no reason that heterodontosaurid ossified tendons should look like those in ornithopods, for example. Aside from the suitably large caniform teeth, the cheek teeth are pointed pegs. |
Heterodontosaurinae:
| Taxon or Taxa: | Time/Place: | Comments: |
| Heterodontosaurus tucki Crompton and Charig, 1962 | Hettangian-Sinemurian (EJ) of South Africa | Heterodontosaurus is the best-known heterodontosaurid. A mostly complete skeleton is known for it, and shows it had relatively large, powerful hands and arms, including a strong thumb claw. Juveniles also had tusks. This animal seems to have lacked continuous tooth replacement like that of most toothed dinosaurs. |
| Lycorhinus angustidens Haughton, 1924 (including Lanasaurus scalpridens Gow, 1975) | Hettangian (EJ) of South Africa | This heterodontosaurine, based on a dentary fragment, was first considered to be a primitive relative of modern mammals. Even with the referral of Lanasaurus, it is not particularly well-known, with only jaw material behind it. |
Genasauria i.s.:
| Taxon or Taxa: | Time/Place: | Comments: | |
| "Acanthopholis": | "A." macrocercus (N.D.) Seeley, 1869 | late Albian (EK) of England | Both of these animals are chimerical: "A." macrocercus is based on ankylosaur scutes and ornithopod vertebrae. "A." stereocercus is based on ornithopod dorsal vertebrae and ankylosaurian tail vertebrae and armor. |
| "A." stereocercus (N.D.) Seeley, 1869 | |||
| "Anoplosaurus" major (N.D.) Seeley, 1878 | late Albian (EK) of England | "A." major is based on an ankylosaurian neck vert and three ornithopod tail verts. | |
| "Dysganus" peiganus (N.D.) Cope, 1876 | late middle Campanian (LK) of Montana | The type tooth of this obscure species has been considered as a hadrosaurid or ceratopsid, but is more likely an ankylosaurian (or, less likely, a hypsilophodont). | |
| Heishansaurus pachycephalus (N.D.) Bohlin, 1953 | ?early Campanian (LK) of China | This animal is a hard-headed ornithischian of some kind, either an ankylosaurian (seems more likely at this point) or a pachycephalosaurian. | |
| Peishansaurus philemys (N.D.) Bohlin, 1953 | early Campanian (LK) of China | It may be that this animal is a juvenile ankylosaurian, or a pachycephalosaurian. | |
| Serendipaceratops arthurcclarkei Rich and Vickers-Rich, 2003 | EK of Australia | This finally gives recognition to the informally-discussed Australian "neoceratopsian" ulna. It was thought to have been similar to Leptoceratops (the gold standard for non-Protoceratops basal neoceratopsians), but realistically cannot be diagnosed beyond Genasauria. | |
| "Tianchungosaurus" (N.N.) Zhao, 1983 | ?J of China | This could be an early basal pachycephalosaurian, or possibly a misspelling of Dianchungosaurus, which was thought to be a heterodontosaurid but is really a crocodilian. | |
Thyreophora: Thyreophora was revived in the 1980s to contain the armored dinosaurs, which by the early 1990s consisted of the ankylosaurians and stegosaurians. There are few characters which support this clade, but one, the possession of keeled, dorsal, parasagittal armor, running head to tail, is pretty strong. However, some researchers believe that the stegosaurians may not belong, which would leave the ankylosaurians alone here with some basal taxa. More basal thyreophoran material may be known from the EJ of China.
| Taxon or Taxa: | Time/Place: | Comments: |
| Scutellosaurus lawleri Colbert, 1981 | Sinemurian-mid Pliensbachian (EJ) of Arizona | This animal was at first thought to be an armored "fabrosaurid," but as that group was disbanded, Scutellosaurus was reclassified as a basal thyreophoran, providing possibly the best example of what an ankylosaurian precursor would look like. It is small, with a very long tail, limb proportions not too dissimilar from Lesothosaurus, and many small armor studs, nodules, and plates. Remains from several individuals are known. |
| Emausaurus ernsti Haubold, 1991 | early Toarcian (EJ) of Germany | This recently-described animal is best known from a virtually complete skull that has some similarities to that of Huayangosaurus. |
| Scelidosaurus harrisonii Owen, 1860 | Sinemurian-mid Pliensbachian (LJ) of England and ?Arizona | Known from a couple of mostly complete skeletons, this animal superficially looks very much like a smallish nodosaurid. It has unusual "tricorn" armor nodules directly behind the skull. Interestingly, mixed in with what is now the type material were hindlimb elements from a "megalosaur"-type theropod. This theropod material was unwittingly designated as the type at one point. Fortunately, the type was officially redefined and no longer includes this extraneous material, now informally named "Merosaurus". Before the type was removed from its matrix, Scelidosaurus was suggested to be anything from an ankylosaurian to a stegosaurian to, most unusually, a fleet bipedal early ornithopod. Possible Scelidosaurus remains suggest that its armor was covered by a keratinous layer, a la turtles' shells. |
Thyreophora i.s.:
| Taxon or Taxa: | Time/Place: | Comments: |
| Lusitanosaurus liasicus (N.D.) Lapparent and Zybszewski, 1957 | Sinemurian (EJ) of Portugal | This dubious basal thyreophoran is known from a skull fragment that is similar to the corresponding elements of Scelidosaurus. |
| "Omosaurus" hastiger (N.D.) Owen, 1877 | early Kimmeridgian (LJ) of England | Based on two spikes thought to be from what we now know as a stegosaurid genus, this species does not appear to be stegosaurian. |
| Tatisaurus oehleri (N.D.) Simmons, 1965 | Sinemurian (EJ) of China | This poorly-known genasaurian is based mainly on a partial jaw. It is not the same as Scelidosaurus, as is sometimes claimed, and there is no evidence that it was a stegosaurian. It has also been regarded at times as a heterodontosaurid or ornithopod. |
Eurypoda i.s.
| Taxon or Taxa: | Time/Place: | Comments: |
| Brachypodosaurus gravis (N.D.) Chakravarti, 1934 | Maastrichtian (LK) of India | Based on a heavily-built ?humerus, this animal is usually tossed off as a dubious ankylosaurian, but may be a stegosaurian. |
Neornithischia: Here again we see a spray of "fabrosaurs," this
time grading into hypsil-type animals. Agilisaurus and Hexinlusaurus appear to have lacked ossified
tendons on the tail, an absence which would have made their tails much more
flexible than most other ornithischians.
"Othnieliidae"
was never much more than a "vanity project" for Othnielia (now Othnielosaurus). There may be a new, undescribed Spanish
"othnieliid".
| Taxon or Taxa: | Time/Place: | Comments: |
| Lesothosaurus diagnosticus Galton, 1978 | Hettangian (EJ) of Lesotho and South Africa | Lesothosaurus is the best-known basal ornithischian, with several skulls and most of a skeleton known. It is a "step up" from Pisanosaurus, and may be the same as the dubious Fabrosaurus. The arms appear to be rather short, with small hands. |
| Stormbergia dangershoeki Butler, 2005 | Hettangian (EJ) of South Africa and Lesotho | Stormbergia is the "large lesothosaur" of long-standing rumor. Three individuals, including one juvenile, are known from partial postcranial remains. Befitting its status in rumor, it was indeed a very large basal ornithischian, getting up to around 2 meters long, not unlike a scaled-up Lesothosaurus but with less of the proportions of a runner. It may just represent adults of Lesothosaurus, though. |
| Agilisaurus louderbacki Peng, 1990 | Bathonian-Callovian (MJ) of China | This small ornithischian may be a very basal ornithopod. Its body plan was suited for agility, as the generic name suggests (although maybe not as much as contemporaneous Hexinlusaurus, which had a longer tibia relative to its femur). It is known from a complete articulated skeleton and skull, which is very tall (I've got to think that there's probably some funky crushing there) with a long rod over the eyes. |
| Hexinlusaurus multidens Barrett, Butler, and Knoll, 2005 (He and Cai, 1983 [originally Yandusaurus]) | Bathonian-Callovian (MJ) of China | Hexinlusaurus has been through quite a few names before it found where it fit. It started as a species of Yandusaurus, which was more of a hypsil-type animal. It then moved on to Agilisaurus, which still wasn't quite right but at least was closer and from the same formation. Gregory Paul considered it a possible species of Othnielosaurus, which again was not quite right. It is known from a nearly complete skeleton with skull, probably not fully grown, and another skeleton and skull have been referred to it; apparently 8 other individuals were referred here, but cannot be located. It is one of those things that was somewhere between a "fabrosaurid" and a hypsil, a small, bipedal, running animal that probably was mainly an herbivore. |
| Othnielosaurus consors Galton, 2006 (originally Laosaurus consors Marsh, 1894) | Kimmeridgian (LJ) of Colorado, Utah, and Wyoming | Othnielosaurus is what is usually represented
in illustrations of Nanosaurus (and now Othnielia, to tell
the truth). A common Morrison
dinosaur, it is known from several
specimens, including an articulated skeleton missing only the hands, skull, and part of
the tail, other partial skeletons, and a possible juvenile individual.
It was some sort of basal hypsil-type thing, although what sort remains
obscure for the moment. To make a long story short, Othnielia was named for Nanosaurus rex, which was just based on a femur that was not distinctive, whereas Laosaurus consors was based on a partial skeleton that was referred to Othnielia. Galton decided to go with the partial skeleton and not the femur. Otherwise, Othnielosaurus is the same thing you knew fondly as Othnielia. Since the Morrison is getting crowded with basal ornithopods (Drinker and Nanosaurus agilis) and ornithopod-like critters (the Fruita heterodontosaurid), it was a good idea to go with the most distinctive of the old remains. |
Neornithischia i.s.: This grouping is for taxa that appear to fit somewhere in the vicinity of Othnielosaurus, basal Cerapoda, and the hypsilophodonts.
| Taxon or Taxa: | Time/Place: | Comments: |
| Drinker nisti Bakker, Galton, Siegwarth, and Filla, 1990 (?Othnielosaurus) | Kimmeridgian (LJ) of Wyoming | Closely resembling Othnielosaurus, Drinker is named after Edward Drinker Cope, contrasting Othnielosaurus, which is named after his arch-rival, Othniel Charles Marsh. It may end up as a species of Othnielosaurus. Drinker had unusual long spreading toes, suggesting it could negotiate swampy terrain well. Bob Bakker has reported finding the remains of over thirty individuals in what might have been a burrow. |
| ?Nanosaurus agilis Marsh, 1877 | Kimmeridgian (LJ) of Colorado and Utah | At one time considered a "fabrosaur"-type ornithischian, Nanosaurus is most likely a basal ornithopod or in the cerapod-to-ornithopod limbo zone, although it could be related to Fruitadens. It is based on molds of its teeth, with other material, including an ilium, femorae, tibiae, and a fibula, referred to it. |
| "Nanosaurus" rex (N.D.) Marsh, 1877 | Kimmeridgian (LJ) of Colorado | It was a nice run while it lasted. Marsh named this species for a femur, that Galton later deemed worthy of its own genus which, when combined with a couple of good partial skeletons, ruled the Morrison of our imaginations for nearly three decades as Othnielia (and you thought I was going to say Allosaurus). However, the femur wasn't that distinct, while the skeletons were, so a Lagosuchus\Marasuchus has been pulled, with the skeletons transferred to Othnielosaurus consors (from an obscure species of Laosaurus that had better remains for a type specimen), leaving Othnielia, and "Nanosaurus" rex, to the dustbin of history (which is too bad, because it was always funny to have an ornithopod with rex as its species name; it's like naming a tiny dog Rex). |
| Phyllodon henkeli (?N.D.) Thulborn, 1973 | early Kimmeridgian (LJ) of Portugal | This is a tooth taxon of uncertain affiliation, from the cerapod-basal ornithopod nexus. It could be a valid genus akin to Drinker. |
Cerapoda i.s.: Cerapoda comes from Ceratopsia+Ornithopoda, and, as the name suggests, contains both, plus the pachycephalosaurians. The name is rather unfortunate, in that mispronunciation can cause confusion with Sauropoda, which is composed of quite different dinosaurs.
| Taxon or Taxa: | Time/Place: | Comments: |
| Albalophosaurus yamaguichiorum Ohashi and Barrett, 2009 | Hauterivian-Valanginian (EK) of Japan | Albalophosaurus is known from a partial skull. It may be a basal ceratopsian, but it is not dissimilar to ornithopods, either. Thus, it gets to sit here until more material is known. |
| Changchunsaurus parvus Zan, Chen, Jin, and Li, 2005 | ?Aptian (EK) of China | This cerapod is based on a skeleton with skull, and a couple of referred skull fragments. It had five premaxillary teeth, a jugal boss, relatively short toothless gaps at the tip of the premaxilla and between the premax and maxilla, and premax bottom margin at about the same height as that of the maxilla. The long predentary and form of the quadrate remind me of Thescelosaurus. Changchunsaurus appears to be very comparable to Jeholosaurus. |
| Claorhynchus trihedrus (N.D.) Cope, 1892 | late middle Campanian (LK) of Montana | Based on partial premaxillae and a predentary, this dubious species has unaccountably been a football, sometimes classified as a hadrosaurid, sometimes as a ceratopsid as originally described. |
| Jeholosaurus shangyuanensis Xu, Wang, and You, 2000 | Barremian-early Aptian (EK) of China | A new Yixian dinosaur, this appears to be something similar to a hypsilophodont, perhaps a representative of a localized east Asian offshoot. It is based on a nearly complete but dorsally compressed skull, along with postcranial remains, with another skull and additional postcranial remains referred. |
| "Magulodon muirkirkensis" (N.N.) Kranz, 1996 | late Aptian-early Albian (EK) of Maryland | This animal is based on teeth which may have belonged to either a "dryosaur" or basal neoceratopsian. |
| Micropachycephalosaurus hongtuyanensis (?N.D.) Dong, 1978 | early-mid Maastrichtian (LK) of China | Famous for having the longest generic name of any classic dinosaur, this animal was ironically very small for a dinosaur. As its name suggests, it was originally described as a pachycephalosaurian, but its remains are too poor to refer it to that group. |
| Notoceratops bonarelli (N.D.) Tapia, 1918 | early Maastrichtian (LK) of Argentina | This animal is based on a now-missing dentary, originally considered that of a ceratopsian. As no ceratopsian remains have ever been discovered in South America, it has been suggested that the bone actually belongs to a hadrosaurid, as hadrosaurids are known from the area. |
Marginocephalia i.s.: The marginocephalian families, Pachycephalosauria and Ceratopsia, are linked by several characters, including the bony shelf found at the back of the skull, which gives this group its name and forms a distinctive frill in ceratopsians.
| Taxon or Taxa: | Time/Place: | Comments: |
| Ferganocephale adenticulatum Averianov, Martin, and Bakirov, 2005 | Callovian (MJ) of Kyrgyzstan | Ferganocephale is a tooth taxon that was interpreted as a pachycephalosaurian, extending the pachy record back into the middle Jurassic. The teeth are unusual for not having marginal denticles (little pointy bits on the edges, if you like). Although the authors admit that there could be some damage if the teeth were swallowed and partially digested, they note a lack of enamel removal. These teeth, one of which was originally tentatively considered stegosaurian, have a good solid unspecialized ornithischian look to them, with a leaf-like shape, except they don't have denticles, which makes them look a bit odd. They share with pachies weak to nonexistent ridges and denticles, along with a few other characteristics, although the teeth could also belong to a basal ceratopsian. Either way, it would be the oldest-known well-supported marginocephalian. |
| Stenopelix valdensis Meyer, 1857 | Barremian (EK) of Germany | This animal is apparently a marginocephalian. Every so often someone comes up with a new case as to why it's a pachycephalosaurian or ceratopsian, but without a head the partial skeleton has proven difficult to pin down. |
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