Pachycephalosauria

    Known in vernacular as the "bone-heads," the pachycephalosaurians are members of a bizarre group of ornithischians with greatly thickened skulls.  It has been thought that the derived members came in two families,  Homalocephalidae and Pachycephalosauridae, and this still may be the case once more remains are found, but no support has been found for Homalocephalidae in recent analyses.  "Homalocephalids" were known as "flat-headed," while the pachycephalosaurids are "dome-headed."  Within Pachycephalosauridae the members are classified by skull characters (as that's often all there is to go on), including closure of certain holes. 
    Whom the closest relatives of pachycephalosaurians are is a subject that has long been debated.  In the past, stegosaurians, ankylosaurians, and ornithopods have been suggested.  Today, they fit relatively comfortably with the ceratopians in Marginocephalia.
    Pachycephalosaurians are reliably known only from the Cretaceous of the northern continents; a supposed large Gondwana taxa, known as Majungatholus, has turned out to be an unusual thick-headed abelisaurid, Majungasaurus (ironically, there may be tiny pachycephalosaurians from the late Carnian [LTr] of India, although the odds aren't great, as getting bone expansions on the skull has been a popular evolutionary option for hundreds of millions of years among dozens of groups; wait for the paper, I guess).  They are typically small bipeds with large skulls relative to body size, and a variety of bony ornamentation decorating the skull.  They are known from rather poor remains, usually only thickened frontoparietal domes for pachycephalosaurids (these, the frontal and parietal, being the bones involved in the domes); as a result, certain parts, like hands, have not yet been discovered.  They are often reconstructed with five fingers, a reasonable number considering the basal number of manual digits in Cerapoda is five, and bipedal cerapodans typically have either four or five.  "Homalocephalids" are currently known from better skeletal material than pachycephalosaurids.  
    The purpose of the immensely thick skulls has long been debated.  The actual brain of a pachycephalosaurian is very small in relation to the size of the skull (although the parts relating to the sense of smell are expanded, suggesting a keen snout), so the massive skull was not for holding a large brain.  Recently, the common interpretation had been that the animals used them to fight amongst themselves in the manner of bighorn sheep.  "Homalocephalids" would have pushed each other, and pachycephalosaurids would have rammed each other.  There are some problems with this idea, including the fact that the round domes of pachycephalosaurids may not have allowed good contact, and could have sent the heads of ramming opponents twisting in directions detrimental to the health of the neck.  Thus, the commonly-illustrated proposal is losing favor to the idea that the domes were mainly for display.
    With the proposal of sheep-like head butting came the proposal the animals lived like bighorn sheep as well, in herds in upland areas.  Neither idea is improbable or contradictory to known evidence; for example, North American pachies are usually known only from worn skull remains, suggesting the bones traveled a long distance to the lowlands.  Males and females have been tentatively identified from Stegoceras skull remains.
    Because pachycephalosaurian teeth bear a passing resemblance to those of troodontids, an inaccurate illustration of a Troodon tooth led to the idea Troodon was a pachycephalosaurian and the same as Stegoceras.  In old works this group may be called Troodontidae.
    Current pachy research is divided into two camps, which lately seem to publish a big paper a year largely reversing what the other side said (this isn't strictly true, but sometimes it feels like this).  The rhetoric has reached the "overheated" stage, and a third-party review by someone who knows these animals is in order, but I think there's only two groups of pachy researchers at this point.  What I've got is a sort of mismatch guaranteed to upset partisans of both groups.  A new monkey wrench is the possibility that "flat-headed" pachies are fully-domed taxa that haven't hit puberty.  The flagship sequence for this new interpretation is spiky Dracorex going to small domed and more spiky Stygimoloch going to less spiky but big domed Pachycephalosaurus.        

<--Pachycephalosauria 
      |--?Stenopelix
      `--+--Wannanosaurus
           `--+--Goyocephale
                `--+--Homalocephale
                     `--Pachycephalosauridae
                          |--Stegoceras
                          `--Pachycephalosaurinae
                              |--+--Colepiocephale
                              |    |--Hanssuesia
                              |    |--Prenocephale
                              |    |--Sphaerotholus
                              |    `--Tylocephale
                              `--Pachycephalosaurini
                                  |--Pachycephalosaurus
                                  `--Stygimoloch

Pachycephalosauria: Being given a generic name ending with a conjugation of "-tholus" has been unlucky for pachies: Stenotholus turned out to be Stygimoloch, Ornatotholus may be just a juvenile Stegoceras, Majungatholus turned out to be an abelisaurid, Gravitholus is dubious, and Sphaerotholus has been challenged not more than a few months after being coined.  The moral of the story is use "-cephale" instead.
    The old Homalocephalidae is now considered a lineage of progressively more derived animals.  Several of them are interesting in their possession of a dentary fang similar to that of the heterodontosaurids

Taxon or Taxa: Time\Place: Comments:
?Stenopelix valdensis Meyer, 1857 Barremian (EK) of Germany This animal is a marginocephalian and apparently a basal pachycephalosaurian based on hip characteristics, although it was thought for a while that it was too primitive to be either a pachycephalosaurian or ceratopian.  It is based on a headless partial skeleton.
Wannanosaurus yansiensis Hou, 1977 Campanian (LK) of China Wannanosaurus is a small animal known from partial skull and postcranial remains.
Goyocephale lattimorei Perle, Maryanska, and Osmolska, 1982 late Santonian-early Campanian (LK) of Mongolia Goyocephale is one of the better known pachycephalosaurians, with a skull and most of the skeleton known.  It is known to have had a dentary fang.
Homalocephale calathocercos Maryanska and Osmolska, 1974 (?Prenocephale) early Maastrichtian (LK) of Mongolia Like Goyocephale, a partial skull and postcranium are known, showing this animal to have been a tubby beast.  An interesting feature, besides the knobbly head, is the greatly expanded lateral processes on the tail vertebrae, which when combined with the relatively wide-set hip bones could allow for the guts to continue back behind the hips for a short distance.

Pachycephalosauria i.s.

Taxon or Taxa: Time\Place: Comments:
Alaskacephale gangloffi Sullivan, 2006 late Campanian (LK) of Alaska Briefly described in 2005, this is based on a squamosal like that of Dracorex, but lacking the long spikes.  
Dracorex hogwartsia Bakker, Sullivan, Lucas, Larson, and Saulsbury, 2006 (?Pachycephalosaurus) late Maastrichtian (LK) of South Dakota This is the flat-headed dragon-looking pachy unveiled in 2005.  It's got big supratemporal fenestrae, so it was not very derived compared to other pachies, especially at its late age (it was a Lancian-age animal).  The type skull, however, is from an individual that wasn't fully grown, so that could have an influence on where it plots out.  I'd guess it was close to full growth, though, because of the spectacular array of spikes and nodes, features usually interpreted as display structures, and one does not do much displaying before reaching breeding age.  It reminds me of a thescelosaur in a Halloween mask of Stygimoloch.  Besides the skull, you also get the atlas and two other cervicals, but only if you act today!
Of course, the species name refers to the main school of magic in the Harry Potter series (Hogwarts), where dragons have made appearances.  Interestingly, as someone on the Dinosaur Mailing List pointed out, there is a character named Draco who is a "butt-head", although not in the grand old pachy fashion (paleontologists are suckers for puns, other types of wordplay, and convoluted etymologies).
Of course, it may have just been a juvenile Pachycephalosaurus.
?"Palaeoscincus" latus (N.D.) Marsh, 1892 late Maastrichtian (LK) of Wyoming This indeterminate tooth taxon could be some sort of pachycephalosaurian.

Pachycephalosauridae:

Taxon or Taxa: Time\Place: Comments:
Stegoceras validum Lambe, 1902  late middle Campanian (LK) of Alberta and Montana Known from a partial skeleton and dozens of partial skull domes, this animal is one of the more common Judithian dinosaurs.  Like most pachycephalosaurians, it is small (for a nonavian dinosaur).  Its premaxillary teeth are wideset and its muzzle wide compared to Prenocephale, indicating different feeding preferences (possibly indicating that Prenocephale was a more selective forager).

Pachycephalosauridae i.s.:

Taxon or Taxa: Time\Place: Comments:
"Stegoceras": "S." browni (N.D.) Wall and Galton, 1979 late middle Campanian (LK) of Alberta Also known as Ornatotholus, and previously (sometimes) regarded as the only North American "homalocephalid", this taxon may instead with its lightly-thickened dome represent a juvenile individual of Stegoceras.
"S." breve Lambe, 1918 late middle Campanian (LK) of Alberta Based on juvenile remains (frontoparietal domes), this material was long assigned to S. validum.  It may represent the basalmost species of Stegoceras, or an example of Prenocephale or its relatives.
"Troodon" edmontonensis (N.D.) Brown and Schlaikjer, 1943 latest middle Campanian- Maastrichtian (LK) of Alberta A rare and poorly known animal, this creature lived alongside Pachycephalosaurus.  Its remains, frontopartietal domes, were long assigned to Stegoceras, but are not generically distinct.  It may be referable to Prenocephale.

Pachycephalosaurinae:  All animals at this level are known to have closed the supratemporal fenestrae (holes that would have been found on the top-rear of the skull) except for Tylocephale, which is incomplete in that area.

Taxon or Taxa: Time\Place: Comments:
Colepiocephale lambei Sullivan, 2003 (originally Stegoceras lambei Sternberg, 1945) early middle Campanian (LK) of Alberta Colepiocephale has a dome which looks somewhat triangular in top view, and has an oblique slope in lateral view. The name literally means "knuckle head" (which leaves me waiting for Curleyia howardi-"nyuck, nycuk, nyuck!").
Hanssuesia sternbergi Sullivan, 2003 (originally Troodon sternbergi Brown and Schlaikjer, 1943) late middle Campanian (LK) of Alberta and Montana Hanssuesia has a very round dome in most views.
Prenocephale prenes Maryanska and Osmolska, 1974 (?Homalocephale) early Maastrichtian (LK) of Mongolia Prenocephale is known from material including a skull similar to that of Stegoceras, but with heavy ornamentation and closed supratemporal fenestrae. 
Sphaerotholus: Williamson and Carr, 2002 (?Prenocephale) S. goodwini (type) Williamson and Carr, 2002 early late Campanian (LK) of New Mexico Based on "bowling ball"-type domes, these are derived pachycephalosaurids.  Beyond that, I am confused by the current state of pachy taxonomy.
S. buchholtzae Williamson and Carr, 2002 late Maastrichtian (LK) of Montana
Tylocephale gilmorei Maryanska and Osmolska, 1974 (?Prenocephale) ?middle Campanian (LK) of Mongolia This animal is known from an incomplete skull.  It could be the same as Prenocephale.

Pachycephalosaurinae i.s.:

Taxon or Taxa: Time\Place: Comments:
Gravitholus albertae (?N.D.) Wall and Galton, 1979 late middle Campanian (LK) of Alberta Based upon a solitary, enigmatic, and probably pathological dome, this large-domed animal probably resembled Hansuessia.
"Microcephale" (N.N.) Sereno, 1997 LK of Alberta Yet to be described, this animal is apparently based on remains including very tiny skull caps.
"Troodon" bexelli Bohlin, 1953 ?late Campanian (LK) of China Currently a memorial to the practice of assigning pachycephalosaurian species to Troodon, this derived pachycephalosaurine requires a new name.

Pachycephalosaurini: This group is diagnosed by nasal nodes and squamosal nodes, which in Stygimoloch become short horns around the rear of the head.

Taxon or Taxa: Time\Place: Comments:
Pachycephalosaurus wyomingensis Brown and Schlaikjer, 1943 (originally Troodon wyomingensis Gilmore, 1931) (?Dracorex, ?Stygimoloch) late Maastrichtian (LK) of Wyoming, South Dakota, and Montana The largest known pachycephalosaurian, at over five meters in length, Pachycephalosaurus had a set of nodules at the rear of the skull and at least four short spikes on the snout, possibly for display or rooting.  It is rather rare, known only from skulls.
An old skull fragment, named Tylosteus ornatus (Leidy, 1872), is usually suppressed in favor of Pachycephalosaurus, but recent study has suggested that it just might be its own thing, more like Dracorex (or just another juvenile). 
Stygimoloch spinifer Galton and Sues, 1983 (including Stenotholus kohlerorum Giffin, Gabriel, and Johnson, 1987) (?Pachycephaloasurus) late Maastrichtian (LK) of Montana and Wyoming Stygimoloch is most famous for the horns projecting from its squamosals.  These would have given the three-meter-or-so long animal a very threatening profile, something similar to the frill spines of centrosaurine ceratopids like Styracosaurus.  The skeleton known as "Sandy," once thought to belong to Pachycephalosaurus, more likely belongs to Stygimoloch.  Its skull has a high, narrow, thin dome.

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