Monday, December 05, 2016

Meet the Tyrants, Part III: Albertosaurines and Alioramines

Today's post combines two small clades for purposes of discussion.  We've finally arrived at the "Tyrannosauridae" proper, within "Tyrannosauroidea"—so in layman's English, we can say the "tyrannosaur family proper" within the extended family of "tyrannosaur-form cousins".  Of course, within the tyrannosaur family proper, there are two subfamilies, the albertosaurs and the tyrannosaurs proper.  And then within the tyrannosaur proper subfamily, there is a radiation of alioramine animals as a separate sub-subfamily.

We'll leave the bigger, more robust "classic" tyrannosaurs for the last post in the series, and talk about the albertosaurs and alioramines today.  Although the albertosaur subfamily is nested outside of the classic tyrannosaur subfamily and the alioramines are nested within it, the two groups yet share a number of superficial similarities.  In fact, the alioramines, which have proven difficult until more recent classifications based on more detailed fossils, are only recently confidently classified at all, and Alioramus has often been considered in all kinds of places all over Tyrannosauroidea.  As one could guess, based on the name, Albertosaurus itself is known from Alberta most especially, and as it happens, the entire subfamily is more confined to the northern part of Laramidia, whereas early classic tyrannosaurs may have been more common in the southern reaches of the continent.  Alioramines are an Asian clade.

Gorgosaurus libratus.  Gorgosaurus is the earliest appearing of both groups, most especially common in the Dinosaur Park Formation of Alberta of the Middle Campanian, which would put it at about 75-77 million years ago.  It also may have been present in the Two Medicine and Judith River formations which would expand its range both geographically and temporally, but the remains are a bit spotty and many belong to an albertosaur instead.  Alberta's Dinosaur Park Formation is the only for sure location for Gorgosaurus.  It is a doozy, though—one of the better known formations, with a real wealth of ecosystem data.  Due a combination of continental placement and other climate factors, the earth was quite a bit warmer than it is today, and the sea levels were quite a bit higher.  The Western Interior Seaway flowed across what are today the Great Plains, connecting the waters of the Arctic Ocean to the Gulf of Mexico, and splitting North America into two island continents; the longer and thinner Laramida in the west and Appalachia in the east.  Gorgosaurus was then found in the northern shores of floodplains coming down from the new Rocky Mountains into this seaway.  The climate was sub-tropical—like the Gulf Coast today in terms of weather, with a heavily forested environment with marked seasonality, including periods of drought and flooding, which seems to have been the source of much of the mass bone-beds that make the formation so famous.  The Dinosaur Park Formation is overlain by a marine sediment, so the sea grew at the end of this period and flooded this particular area, at least.

Like everyone else in this subgroup, Gorgosaurus was large, but smaller than the classic tyrannosaurs; up to maybe 30 feet long at the largest, and rhino sized in weight.  Compared to classic tyrannosaurs, he was relatively small, and especially light-weight, with longer legs, and a longer and less tall snout.  Most likely, it was faster, and may have focused on quicker, more lightweight prey—or at least have been capable of attempting pursuits of such.  One of the curious notations about Gorgosaurus is that it appears in the same time and place as another large tyrannosaur; Daspletosaurus which is an earlier and smaller classic tyrannosaur, with a more powerful build and head.  It is unclear exactly how the two coexisted and what—if anything—they did differently to avoid direct, head-to-head competition.  Suggestions that Gorgosaurus was a more specialized hadrosaur hunter while Daspletosaurus was a more specialized ceratopsian or ankylosaurian hunter seems to be mere speculation, and good hadrosaur remains have been found in the stomach region or more than one Daspletosaurus specimen.  The ecological significance of two similar-sized tyrannosaurs in the same formation is as yet unexplained.

A number of very famous Late Cretaceous fossils were found in the Dinosaur Park Formation.  In fact, I have to admit I'm still just a little bit bitter that almost every classic dinosaur kids book that I read as a kid readily threw Dinosaur Park Formation animals into the Maastricthian and suggested that they were hunted by T. rex itself.  This is not the case.  Famous lambeosaurine (Corythosaurus, Lambeosaurus, Parasaurolophus) and somewhat less famous hadrosaurine duck-bills were common, and many famous ceratospsians (Centrosaurus, Chasmosaurus, Pentaceratops, Styracosaurus, Monoclonius, etc.) animals made up what was probably the most likely prey for both tyrants, and both nodosaurid and ankylosaurid armored dinosaurs are known.  There's also famous guys like Struthiomimus and Ornithomimus, assuming that those are actually two separate genera, dromaeosaurs and troodontids (including Dromeaosaurus and Troodon themselves), therizinosaurs, oviraptorosaurs, pachycephalosaurs, and at least one kind of hypsilophodont.  A curious although unexplained phenomena is the fact that of the four largest-bodied and well-known dinosaur groups from the area, subfamilies were common during the Campanian that later either became uncommon or went extinct altogether to be replaced by representatives from another subfamily within the same family that was previously rare: lambeosaurine hadrosaurs replaced with saurolophine hadrosaurs; the centrosaurine ceratopsians seemingly go extinct completely to be replaced by chasmosaurine varieties, the nodosaurs survive in Appalachia, but fade away in Laramidia to be replaced by ankylosaurs, and—of course—the Albertosaurs do not survive the Campanian, and the classic tyrannosaurs replace them.  The cause of this subfamily faunal replacement is not, to my knowledge, at all adequately explained.

As an aside, there are some small numbers of skin impressions of Gorgosaurus.  Although not meant to convey the entirety of the animals body, of course, it's worth noting that the very small sample that we do have has quite fine, bead-like scales, not terribly unlike what would be found on a Gila monster today.

Dinosaur Park formation
Albertosaurus sarcophagus.  As the Bearpaw sea which flooded Dinosaur Park receded again, Gorgosaurus fossils are replaced by the younger Albertosaurus, which was a very similar animal in many respects (there have been numerous attempts over the years to combine the two into a single genus, which is usually resisted.  But they are still similar enough that the attempt continues to prompt reexamination periodically.)  Although similar in many respects to the environment of the Dinosaur Park formation, it is believed that the Two Medicine Formation, in which most albertosaurs are found, was cooler and drier.  They are found in the boundary of the Campanian and early Maastrichtian, about 70 million years ago or so, up until maybe 68 million years ago at the youngest, at which point the more classic Lance and Hell Creek faunas appear to dominate.  That said, in some ways, the fauna in which Albertosaurus operated is still similar to the one in which Gorgosaurus did; chasmosaurine certatopsians like Achiceratops, Eotriceratops, and Arrhinoceratops have replaced the centrosaurines (although Pachyrrhinosaurus is still common) and the lambeosaurine hadrosaurs have similarly seen a great deal or replacement with saurolophine hadrosaurs, including early species of Edmontosaurus, Saurolophus and Hypacrosaurus.  The faunal replacement isn't as advanced as it is in later formations, but it's also clearly happening.  This is the changeover from the so-called Judithian faunal stage to the Edmontonian.

Another interesting fact about Albertosaurus specifically are the group fossil sites that have almost breathlessly been interpreted quite frequently as evidence of pack hunting.  It may or may not have actually been pack hunting (it could be more similar to Komodo-dragon style carcass mobbing, or shark feeding frenzies than true pack-hunting).  Nonetheless, the intriguing possibility of tyrannosaur hunting packs is a tantalizing idea, to say the least, and it has been popular ever since the discovery of the Dry Island bone bed.

Albertosaurus has also become a bit of a waste-basket taxon, as they call them, for any vague or non-diagnostic tyrannosaurian bone fragment.  It's possible that it extended outside of Alberta, and the name has been used to refer to fossils from the East, Wyoming, New Mexico, and even Asia—in many cases, these may well represent sub-adults of classic tyrannosaurs, or some unknown member of the albertosaur family at most, but probably do not actually represent Albertosaurus sarcophagus at all.  A number of specimens now identified as sub-adult or younger classic tyrannosaurs have also been referred to Albertosaurus in the past, sometimes even raising up new species that fall into ignominy as nomina dubia, or doubtful names.   There may well be more varieties of albertosaurs out there, we just have no good evidence of them today.

Due to the temporal and geographical placement of Albertosaurus relative to Gorgosaurus, it has been proposed that the former may be a literal descendant of the latter by some.

Alioramus remotus, altai.  The alioramine tyrannosaurs are not as well known as the others, and it has often been difficult to place them until some very recent studies.  To further complicate things, few of the taxa are known from fully adult specimens, and there has been vigorous debate about whether some of its unusual features are related to its juvenile age set, or if they represent primitive retained characteristics.  Because of this, the Hone cladogram (and the Loewen cladogram) both place alioramines just barely outside Tyrannosauridae proper, but the newer Brusatte and Carr cladogram reaffirm earlier efforts to place it in a basal position within Tyrannosaurinae itself.  Although basal, however, Alioramus is not early.  There are probably two species, A. remotus and A. altai the latter being known from more complete remains.

The alioramines are relatively lightweight and small with remarkably long, pointed snouts compared to other tyrannosaurines.  This snout is what has so mystified researchers especially, since a longer, less tall snout is both a basal trait in Tyrannosauroidea as well as a juvenile trait known from such classic, highly derived tyrannosaurines are Tarbosaurus and T. rex itself.  This snout seems to have been adorned with a very bumpy nose ridge as well.  The discovery of adult specimen Qianzhousaurus from southern China has probably cleared up the fact that this was merely a retained condition in this otherwise quite derived sub-subfamily.

Not only are the members of this family not particularly well known anatomically, but we also know relatively little about them.  Alioramus—both species—seem to be from the Nogon-Tsav beds in Mongolia.  Not much is known about this formation, although it is presumed to be the same age as the nearby Nemegt Formation, and may have shared a great deal of its fauna.  Of course, the Nemegt has also not been radiometric dated, so its exact age is unknown, but probably Maastrichtian, the final period of the Cretaceous and the end of the Age of Dinosaurs.  Believed to be wetter and more lush than the arid Barun Goyot and Djadochta formations on top of which it lies, forested river valleys and floodplains, subject to periodic drought, seem to be the norm.  Although not described in detail, the discoverer reported informally that Tarbosaurus as well as therizinosaurs and ornithomimosaurs were discovered at the same locality.  If the fauna is indeed similar to the Nemegt, which seems likely, then it probably had troodonts, pachycephalosaurs, very large hadrosaurs, titanosaurs and ankylosaurs as well.

Qianzhousaurus sinensis.  Only described in mid-2014, this is the critter that helps to clear up the relationship of the alioramines with the rest of the tyrants.  Long known by the nickname "Pinocchio rex" due to its extended snout, it shows conclusively that long-snouted tyrants were a real thing; an alternative breed of predators that were fairly wide-spread throughout Asia, and not merely a confusing mess of sub-adult classic tyrannosaurs with juvenile features.  As much as 30 feet long, this one was actually an adult, and it is found pretty far away from the Alioramus specimens, in southern China's Nanxiong Formation, which is believed to be from the very latest Maastrichtian.

A list of the relatively few animals known from the Nanxiong Formation is not a "who's who" of late Cretaceous dinosaurs; it's more of a "who did you say again?" Formation with a very diverse set of oviraptors, but also a hadrosaur, a titanosaur, and a therizinosaur.  It's believed to have been similar in many respects to the Nemegt Formation, which we'll talk a lot more about next time when we discuss Tarbosaurus.

All in all, much of the Asian fauna, with the exception of the longer-known Mongolian beds described by the Soviets, there are a lot of questions and a lot of work to be done to really understand better what the Asian late Cretaceous ecosystems were like.  These are not nearly as well known as the earlier Yixian Formation and other similar northern China formations, and a lot more work needs to be done to better describe the alioramines and the environments in which they lived.

Thursday, December 01, 2016

Meet the Tyrants, Part II: Intermediate grade basal tyrannosauroids

Yeah, that's a mouthful.  It's been a pet peeve of mine for many years that cladistic methodology has arbitrarily decided that paraphyletic groups are out, since paraphyletic labels are often incredibly useful and convenient.  How many times, if you read dinosaur books, do you hear references to prosauropods, thecodonts, hypsilophodonts, or for that matter, the word dinosaur itself?  All of the time.  Often, sadly, with a pedantic and somewhat self-righteous caveat that you "shouldn't" because it's "imprecise."  Actually, you could construct paraphyletic groups that are just as precise as monophyletic ones—except that of course nobody does because of the arbitrary rule that you shouldn't.

Be that as it may, there is no paraphyletic label to refer to intermediate grade basal tyrannosauroids because nobody had described any or even knew that the group existed except as some kind of ghost lineage, prior to the advent of phylogenetic analysis.  Heck; as recently as Greg Paul's Predatory Dinosaurs of the World he was still calling tyrannosaurs carnosaurs and presuming that they were descended from the allosaurs or something.  Lacking a preexisting label, and unwilling to type out repeatedly "intermediate grade basal tyrannosauroids" I'm going to just grab the last syllable of that big mouthful and call them 'roids here in this series.  What are the 'roids, exactly?  Essentially they are the members of the tyrannosaur family that don't cluster as part of the early proceratosaur radiation (see last post) and also don't qualify as part of the more derived tyrannosaurid clade either (see next two posts).  So they're intermediate grade; in between the early radiation and the later radiation, but they're kind of strung out rather than forming their own "resolved" family.

As I said last time, I'm using the David Hone cladogram from the book I just read, with only a few minor changes.  That seems to have been based on the Brusatte and Carr 2010 study, for the most part.  Hone also refers to the Loewen et al. 2013 cladogram, which is what Wikipedia has, for whatever reason, as its default cladogram. Brusatte and Carr did a new 2016 cladogram; actually two; based on two methodologies, that give very similar results to each other.  I suspect that, although they're pretty new and haven't yet "sunk in" that these will end up being the standard for some time.  I probably should have used a composite of them instead of the earlier Hone one, but for reasons of convenience to me, the Hone one does the job well enough—besides this whole four post blog series was kicked off because I read Hone's tyrannosaur book.  I'll refer, in the specific animal entries, to where it differs from what was more recently published.

Aviatyrannis jurassica. This little guy is often missed from the cladograms (he's not included in Hone's) and even the new Brusatte and Carr study only includes it in the Bayesian cladogram, not the parsimony cladogram.  Known only from a tiny partial hip, not much can be said about this guy except that he's very small; probably about 3 feet long and as little as 12 lbs. or so—although it may well not have been fully grown.  When I was a kid reading every dinosaur book I could get my hands on, Compsognathus was routinely called "the smallest dinosaur" even though he was larger than this specimen.  (Of course, many smaller dinosaurs have been found since then, especially from some of those ashy lacustrine Chinese formations.)  The Bayesian cladogram calls out Aviatyrannis as a basal—in fact, the most basal—'roid outside of the proceratosaur family.  It was originally referred to Stokesosaurus because of its obvious similarities.  Keep in mind that in the new Brusatte and Carr cladogram, Stokesosaurus and Juratyrant fall outside of proceratosaurus, and just up the tree from Aviatyrannis.  Aviatyrannis is from the Alcobaça Formation, which I don't know very well and can't readily find much information on, from the Kimmeridgian (155-150 or so million years ago) from Portugal.  Although I can't find much info on the Alcobaça Formation, the Lourinhã formation from nearby at late Kimmeridgian and Tithonian (150 to 145 or so million years ago) is remarkably similar to the Morrison formation, even having many of the exact same genera (Torvosaurus, Allosaurus, etc.) and when they're not the same, they are extremely closely related and similar (stegosaurs, brachiosaurs, camarasaurs, diplodocids, hypsilophodonts, early iguanodonts, etc.)  Most likely Aviatyrannis occupied a similar faunal assemblage and ecological niche as its close relative Stokesosaurus.

Dilong paradoxus.  Although a 2014 cladistic study (Porfiri et all) found this within Proceratosauridae, every other cladogram has Dilong as one of the earliest 'roids.  That isn't to take away from the fact that it was very similar in many ways to the proceratosaurs; long, three-fingered arms, relatively unspecialized ankles and legs, etc.  Dilong had the typical tyrannosaur pubic boot, though, and lacked the crests of the proceratosaurs.  Most famously, it was feathered; with a kind of filamentous, "woolly" feathery covering.  At only about 5 feet long and probably no more than 35 lbs or so, Dilong was the size of a medium sized dog.  Coming from the 125 million year old Barremian or earliest Aptian stage Yixian Formation of China, it's also among the most recent of the animals we've reviewed so far (excepting Sinotyrannus who came from the layer immediately above it).  The Yixian is famous for its preservation of small biota—not that such small biota was unique or even unusual, but rather that preservation of them is.  It's a big question how much of the Yixian type biota can be applied more generally across wide swaths of the Mesozoic for which no comparable small critters are known, but probably it can be at least in broad strokes if certainly not with regards to the very specific species found.  The environment is lacustrine, so animals like frogs and toads and loads of fish are well known.  There are a lot of small mammals, and the Yixian has gone a long way towards clearing up all kinds of questions about the state of mammals during the Early Cretaceous; many of the specimens are from lineages which are poorly known elsewhere, like gobiconodonts, eutriconodonts, metatherians, etc. although some more traditional mammals, including a multituburculate and at least one eutherian are known.  Lots of lizards, turtles and pterosaurs are also known.  Heck, we even know a bunch of bugs from the Yixian.

Perhaps most famous from the Yixian fauna, however, is the vast assemblage of otherwise completely unknown primitive birds and bird-like maniraptorans—troodontids and deinonychosaurs, often tiny fliers and gliders and other small creatures.  Not only is the fact that so many of these tiny creatures having been found unusual about the Yixian, but the fine preservation with many feather imprints for many creatures is very unusual and makes the formation justly famous.  Larger creatures include some oviraptorosaurs, therizinosaurs, small bipedal ceratopsians, compsognathids, primitive ornithomimosaurs, small iguanodont-grade ornithopods, etc.  Very little is known about the larger animals, which is very atypical, but there is a medium-sized early hadrosaur (about 20-25 feet long) and some poorly known and probably on the smaller side (based on the remains we have so far, at least, which of course may not have been adults) basal titanosaurs.  All in all, the Yixian is a very unusual formation in terms of preservation bias, but it probably was rather more prosaic that we think, it's just that because it has a very different preservation bias than most, so we see a glimpse into a slice of the Mesozoic world that was commonplace, but rarely preserved in enough detail for us to perceive it elsewhere.

Yutyrannus huali. This is the largest of the animals we've surveyed so far.  In the Hone cladogram, it comes next on the 'roid line after Dilong, but Brusatte and Carr made a strong case for it being a proceratosaur, clustered very closely with Sinotyrannus (on the other hand, they stick Stokesosaurus and Juratyrant together about where Yutyrannus is on the other cladograms.)  Although Yutyrannus is large; allosaur-sized, so around 30 feet long and probably weighing as much as a rhino, it really looks more like a scaled up Guanlong rather than a close analog to the more derived tyrannosaurids.  True; it had a shorter neck and larger head, but that comes with the territory when you're a large, probably apex predator for the region.  Most famously, of course, Yutyrannus is known for being feathery in spite of it's large size.  Coming from the same formation as Dilong, it was pointed out that the Yixian was probably colder during the winter than many other formations surveyed, with freezing temperatures and snow—not unlike continental, temperate forest zones of today.  This may at least partly assist in explaining so many specimens found seems to be so "fuzzy."

Known from three nearly complete skeletons; one adult, one subadult and one juvenile, the animal is relatively well known.  As Brusatte and Carr point out, the proceratosaur analogs are multiple (except for the size)—it had narrow teeth, a fairly lightly built head with little in the way of post-orbital bosses and horns, it had a snout crest (more modest than Guanlong's, but not nothing), unspecialized legs, long arms with three-fingered hands, etc.  In short, it's interesting to see that the tyrants were starting fairly early to reach for apex predator status in their faunas, but because it comes from a "primitive" cluster, it's not really very fair to compare them to the very derived and specialized forms of the more "classic" tyrannosaurs.

Yutyrannus "pack" within its natural setting.
Eotyrannus lengi. One of the earliest described and recognized tyrannosauroids, Eotyrannus went a long way towards starting to clear up all kinds of questions about how the tyrants formed, and where.  It was believed prior to this discovery that the tyrants had an Asian origin—this was before the discovery of all of the Asian feathered specimens, and before the recognition of Proceratosaurus and Stokesosaurus as members of the group.  Eotyrannus comes from the exact same time as Dilong and Yutyrannus, but from the opposite side of the Eurasian continent, in England's Wealden Group on the Isle of Wight.  The specimen is known from one fragmentary but intriguing skeleton that is probably a juvenile or subadult about 13 feet long or so—so the full size of the adult animal is unknown.  Some of the very classic Barremian-Aptian European dinosaurs are known from the same formation: Hypsilophon, Iguanodon, Polacanthus, some poorly known diplodocid and rebbachisaurid sauropods, etc.  Predators of the fauna are a little more poorly known, or at least were until more recently, but some spectacular discoveries in the last decade or two have actually opened up various previously unrecognized families, such as Baryonix, Neovenator, and... of course, Eotyrannus.  Small predators probably include, although the material found is difficult to interpret, various maniraptorans, dromaeosaurs, compsognathids, etc. In this regard, the fauna is a bit different than what we see in Asia, and more like what we see in North America at this time—a large carnosaur (Acrocanthosaurus), several sauropods of lingering Jurassic vintage lineages (Sauroposeidon), small, medium and large sized ornithopods, a spinosaurid, etc. Or, for that matter, it's not terribly different from Gondwana-style faunas as found in South America and Africa from the Aptian-Albian either.  But all of this was about to change; sauropods were going to fade in the northern hemisphere, carnosaurs were going to become much more rare if not completely absent, ankylosaurids and ceratopsians and more derived ornithopods like hadrosaurs were all on the rise.  And hunting them, instead of carnosaurs, were going to be tyrannosaurs.

Of course, we're not quite sure how we got there yet.  Sadly, there is a pretty big gap between these last periods of the Early Cretaceous and the last periods of the Late Cretaceous.  We actually know relatively little about the Cenomian, the Turonian, the Coniacian or the Santonian periods; a gap of nearly twenty million years.  There's one more early, somewhat primitive 'roid left to talk about before we hit on this gap, though.

First; one other curious fact.  Porfiri et al did a cladogram in 2014 that recovered Eotyrannus as a megaraptor.  Otherwise, the megaraptors, while phylogenetically a bit bouncy in general, are assumed to be derived neovenators, i.e., late-appearing cousins of Allosaurus.  Based on this, there's been a bit of a swan song floating around in paleontological circles in general that the megaraptors might be a derived radiation of tyrannosauroids of some kind.  I think this is almost certainly a red herring.  The tyrants have been a bit like a Pied Piper of sorts for drawing comparisons with any other therapod—and even occasionally some thecodonts: Sankar Chatterjee once implied that the tyrannosaurs were actually direct descendants of rauisuchians like Postosuchus.  A different carnosaur group, the carcharodontosaurids are often claimed based on one or more similarities to possibly be related to the tyrants, and abelisaurs are as well—but this is the cart coming before the horse, as based on many other synapomorphies, the carcharodontosaurids are a yet different radiation of late-appearing Allosaurus cousins and the abelisaurids are very late appearing descendants of Ceratosaurus.  I had thought, briefly, of cataloging the megaraptorans as part of this catalog because of that claim, even though I don't personally take it very seriously, but I'm now thinking it makes more sense for me to eventually do a catalog of carnosaurs too—my other favorite lineage of big dinosaurian therapods.

Xiongguanlong baimoensis.  This medium-sized carnivore is from China's Aptian-Albian period (125-100 million years ago.)  The exact age isn't clear, but I'm angling towards the earlier end of that big 25 million year range as being more likely; which makes the problem noted just above even worse; instead of a nearly 20 million year gap, it could be closer to 45.  There are still a number of 'roids left to cover, but after Xiongguanlong they are much later in time, and cluster in terms of features much more closely with the derived tyrannosaurids.  Although Xiongguanlong is about the same size and similarly built to Eotyrannus, with a relatively gracile form, longer neck, smaller head, long arms, three-fingered hands, etc.—and like them, in the Brusatte and Carr cladograms, he clusters near the proceratosaurs and represents a similar "grade" of 'roid—he also includes the first appearance of some "tyrannosaurid grade" developments; strengthened skull with some specific struts for muscle attachment, broadening of the base of the skull, ventrally convex maxillae, assymetric serrations on the maxillary teeth, etc.  This is kind of obscured by the fact that it also had a very long snout, quite unlike the tall and deep snout of most derived tyrants, and also lacked the D-shaped spike-like teeth in favor of narrower, more "knife-like" teeth that earlier forms had.  The earlier theory that tyrants evolved in Asia is probably not true; we have proceratosaurs and other primitive forms all over the place, but the specific features of the derived forms might well have, as Xiongguanlong is the first to really have some of them.  However... we don't really know a lot about what followed Xiongguanlong as we don't have any other 'roids until we get to the Campanian period of 80 million years ago that are sufficiently complete that we can tell much about them.

Bagaraatan ostromi.  Brusatte and Carr didn't even include this specimen, although Hone and Loewen do—and probably for good reason.  We really don't even know if this is a 'roid at all.  Thomas Holtz called it one, but with equal authority it's been called a troodontid and a maniraptoran.  It might even be chimerical, which would explain the difficulty in placing it.  If it was a 'roid, it is in the middle of the gap, from the Nemegt Formation of 70 million years ago in Mongolia.  It's known from partial legs, hip and tail, and a bit of jawbone—with features cherry-picked, it appears, from all over Therapoda from birds to even ceratosaurs.  It would have been relatively smallish; probably smaller than Eotyrannus, but other than that, it's probably not worth talking too much about because it's more confusing rather than helpful to consider it.  If it is a 'roid, it doesn't really seem to be indicating much in the way of development towards the derived tyrants, so it might have been a weird off-shoot doing it's own thing.

Raptorex kriegsteini.  This is another problematic specimen.  While undoubtably part of the tyrant lineage, exactly what it is or even when it is from is questionable.  It was initially interpreted as from the Yixian formation (sadly, it wasn't excavated by professionals, so the details have to be guessed at.)  It was later decided that it probably came from the Iren Dabasu of the late Campanian, or a "similar" formation.  It's a very small juvenile hindquarters that cannot be confidently paired with any adult specimens, so many have put out calls to declare it a nomen nudum or undiagnosable specimen.  The latest may seem to be that it is an extremely young specimen of an animal very much like Tarbosaurus.

Alectrosaurus olseni.  Another difficult specimen, Alectrosaurus is known properly from a foot, and maybe some very poorly preserved other bits and pieces from the Iren Dabasu formation of Mongolia from the Campanian.  Other material has been referred to it, including some skull and shoulder elements, from the Bayan Shireh formation (of uncertain age—maybe earlier) although this material may not belong to this genus.  So, what we have really is merely a good, solidly tyrannosaur leg from the Campanian.  Early Campanian according to some (83 million years ago) and Campanian-Maastrichtian (roughly 75 million years ago) according to others.  Most workers leave this out of cladograms entirely, because although it's definitely tyrannosaur, it comes up all over the place, with as many as eight equally valid positions, so its relationships and even its specific features are simply too poorly known for it to be very useful.

Dryptosaurus aquilunguis. Originally named Laelaps, this is famous from a beautiful and dramatic Charles Knight painting of two individuals fighting (Leaping Laelaps from 1897.)  It has since fallen out of the public consciousness a fair bit.  It wasn't recognized as tyrannosaurian until 2005 when the discovery of Appalachiosaurus gave it some context.  Those two are the only known large predators from Appalachia; North America was divided by an epicontinental sea into two separate "island continent" landmasses in the Late Cretaceous; eastern Appalachia and western Laramidia.  Sadly, Appalachia is not well known in terms of its fauna.  Other than some hadrosaurs and nodosaurs it's not 100% clear what else lived on Appalachia, but it's curious that nodosaurs (which were otherwise replaced by ankylosaurs on Laramidia) and these two interesting late-appearing 'roids (instead of derived tyrannosaurs) are here, as well as a very different assemblage of hadrosaurs than are found in Laramidia.  There are also some scanty remains that have been referred to ornithomimosaurs and dromaeosaurs, and there even appears to be one very atypical and specialized ceratopsian.  Overall, Appalachia was significantly different than Laramidia, and has a rather unique "island continent" phenomena to it in terms of fauna that survived here that did not in other, more connected continents.

Leaping Laelaps
Dryptosaurus has some features in common with more basal 'roids; a more gracile build generally, smaller, less strengthened head, longer arms, three fingers with long claws (although some proposals suggest that only two were functional in any way), etc.  Curiously, in one of the two phylogenies in the recent Brusatte and Carr paper, Dryptosaurus comes up nestled within Tyrannosaurinae as a close relative of Alioramus.  The authors don't actually believe this result, however—they account for the fragmentary nature of the skeleton thus giving fewer points of comparison as well as the long ghost lineage of the Appalachian 'roids as obscuring exactly what was going on with the radiation in the east, most likely.  It was probably about 25 feet long and weighed a ton and a half.  Although somewhat smallish for an apex predator of the Maastrichtian, as far as we know, it was the largest guy on the block in its locality.

Appalachiosaurus montgomeriensis.  Found in the Turnipseed Dinosaur Site (what a great name!) from the Demopolis Chalk Formation in Alabama, this is the best known eastern therapod, from about 40% of a skeleton.  Although the skeleton indicates an animal about the same size as Dryptosaurus, i.e. a little over 20 feet long, it also is not fully grown and seems to indicate a subadult that was maybe 2/3 of its adult size.  It's from smack dab in the middle of the Campanian, about 77-79 million years ago.  The Demopolis Chalk formation is actually a marine environment and was deposited under a good 100 ft. or so of water.  Many of the dinosaur fossils from Appalachia seem to have been bodies that tumbled down from the foothills or lowland plains in river floods and were washed out to sea, which makes putting them in their proper context a little bit difficult.  A decent-sized hadrosaur (Lophorhothon) is known, as well as some nodosaurian, dromaeosaurian and ornithomimid remains that were probably contemporaneous.

Appalachiosaurus was initially believed to be a very eastward trending Albertosaurus, but subsequent investigation into the remains uncovered its true nature as a derived, yet non-tyrannosauridan tyrannosauroid; or a late-breaking and advanced 'roid.  As I said above, it also provided enough information to compare with Dryptosaurus which led to the belief that Appalachia may have had "island continent" faunal patterns, with "relics" of groups that were otherwise replaced across the Laramidian-Asian intermittently connected landmass.  It also raises questions as to whether or not any connections of note happened between Laramidia and Appalachia—it kind of appears not, or if there were, not very much participated in faunal exchange.

Although it appears earlier, it appears also to be more derived in its own Appalachian 'roid evolutionary path than Dryptosaurus, with a fairly longirostrine skull that has some features in common (against all expectations) with Alioramus from Asia.  The forelimbs are not confidently known, so it's unclear if they were long or reduced, or if they had two or three fingers.  A lot more work, if it can be pulled off, to uncover the secrets of Appalachian dinosaurs would be very welcome.  We're just starting to realize how potentially interesting the Appalachian island continent may have been in the Campanian and Maastrichtian.

Speaking of Alioramus, in the Hone cladogram, it's located next, just outside of Tyrannisaurida and closely related to Appalachiosaurus.  However, that wasn't always the case; in fact, Alioramus was often seen as very tyrannosaurine in nature specifically, but showing juvenile characteristics, such as a long snout, light, smaller build, long legs, etc.  Many even proposed that it was merely a juvenile Tarbosaurus.  With the discovery of a second Alioramus species and the similar (albeit larger) Qianzhousaurus, it was recognized that it was indeed an atypical radiation of early derived tyrants that were lighter, probably faster, and longer-nosed than the deep-snouted classic tyrants, which means that it is not closely related to Appalachiosaurus after all.  The Brusatte and Carr phylogenies show this clearly, and it is one major change that I've adopted to the Hone cladogram that I've otherwise using for purposes of this series of blog posts, in part because it makes it easier for me to lump the alioramines with the albertosaurines for my next post, and then finish up in a fourth post with the classic tyrannosaurines.

The Brusatte and Carr cladogram also removes Bihastieversor from within Tyrannosaurinae and puts it just outside Tyrannosaurida.  The logic here is that the Loewen (and even their own earlier work, which is what Hone is repeating) gave too much diagnostic importance to the "tallness" of the skull.  Once that's corrected for, it makes sense based on the rest of the analysis to remove deep-snouted Bihastieversor from deep within the tyrannosaurines based on the coincidence that it has a tall head, and put the alioramines within the tyrannosaurines in spite of their shorter, longer snouts.  That said, this post is long enough, so I'm going to do Bihastieversor along with the rest of the guys it more closely superficially resembles, the way Hone has written it up in his book.

There are also a few other specimens that are sometimes referred to as belonging to the 'roids, but they are generally known from too fragmentary and non-diagnostic material to really be useful, and most cladograms don't include them because they merely introduce noise rather than helpful info.  These specimens include Labocania from Campanian Baja California, which may be an abelisaurid, or even some other type of therapod altogether, Timimus from Australia's famous Dinosaur Cove locality, which was clearly some kind of coelurosaur, but may not have been any type of tyrant (see swan song discussion under Eotyrannus above) and Timerlengia from the Turonian of Uzbekistan.  The latter probably is a tyrant of some kind, presumably a slightly later fairly close relative of Xiongguanlong, which should be exactly the kind of "missing link" both in phylogeny and in time that we'd love to find.  Sadly, the remains are too scrappy to tell us very much; merely two braincases, and some other bones here and there that have been somewhat dubiously referred to the genus.

Wednesday, November 30, 2016

Meet the Tyrants, Part I: Proceratosaurs

The tyrant lizards are a rather diverse family, and can be divided roughly into five groups, with a sixth that is somewhat dubious and who's relationship to the rest of the family (if it is indeed one at all) is uncertain.  Due to the strange way in which cladograms are constructed, one of those five groups is kind of "everyone else who doesn't fit into one of the other four"; i.e. "basal" tyrannosauroids.  For this first catalog, I'll be covering the proceratosaurs, which are the earliest appearing and generally the most primitive, showing marked gross morphological similarity to other basal coelurosaurs, like say, Ornithomimus or the Coeluridae.  In fact, the Coeluridae have occasionally been recovered in some studies as very basal tyrannosauroids (I won't do that for my purposes here; I think it most likely that they are basal coelurosaurs, and possibly the sister group to tyrannosauroids at best; which seems to be the most common belief among practicing paleontologists today.)  Although I don't accept that placement, the point is that guys like Tanycolagreus, Ornitholestes, and Coelurus, who are found in the Morrison Formation, are very similar to guys like Stokesosaurus, which is a tyrannosauroid and in fact they have often all been confused for one another at various times without very careful analysis. This highlights the environment, if you will, in which the tyrants got their start and the type of animal that the earliest tyrants really were.

Because different cladograms have different data points, i.e., sometimes basal tyrannosaurs are included in proceratosauridae, which in other cladograms will not be, etc., I've had to settle on one way of doing things.  For convenience, because it breaks up the genera into more or less equally sized groups (more or less) I'm going to be following Hone's "standard" cladogram, more or less, with a few changes from the very recent Brusatte and Carr cladogram that I posted in my last post, and the Loewen cladogram (specifically, the inclusion of Alioramini within Tyrannosaurinae.  So, proceratosaurs with this post, then the rest of the tyrannosauroids who don't manage to belong to Tyrannosauridae specifically.  There really should be a better "paraphyletic" family name for this group in common usage (like we have for iguanadonts, or hypsilophodonts or prosauropods) but there isn't, so rather than type out "basal tyrannosauroids" all the time, I'll probably call them 'roids for short.

Proceratosaurus bradleyi. The earliest (or tied for that position anyway) tyrant is known from an incomplete skull and lower jaw, missing the top/back of the skull, but we have the mandibles, the nares, and a bit of the forward skull roof.  It's from the Bathonian (Middle Jurassic; 167 or so million years ago) in the Cotswold region of southwest England.  It's remains aren't much to go on, in fact until just a very few years ago, it wasn't known what kind of animal this really was.  Because it had a nasal crest, it was presumed to be a small ancestor to the Morrison therapod Ceratosaurus, hence the name.  Later, it was recovered as a generic "coelurosaur" and considered to be similar to Morrison therapod Ornitholestes.  Based on the size of the skull, the entire animal was fairly small; similar to a medium to large sized dog like a collie or labrador and would not have come up to a normal person's waist.  It was recovered quite early (1910) and at the time, little care was made to attempt to link the skeletons to a fossilized ecosystem, if it were even possible, so the environment and surrounding fauna in which it operated is unknown—although nearby fossils from the same age (roughly) include Megalosaurus, primitive sauropods (possibly Cetiosaurus) and some indeterminate ornithopods.  Also from the same general area (although probably a few million years earlier) are the indeterminate Iliosuchus remains.  Nobody knows for sure what these are, because they're too fragmentary to be diagnostic, but they may represent an even earlier (and smaller) earlier tyrannosauroid of some kind.

Kileskus aristotocus. Aside from the possible Iliosuchus, Kileskus is the next proceratosaur, found in Middle Jurassic (probably also Bathonian) strata in southern Siberia.  Not only is it about the same age as Proceratosaurus, curiously it's also about the same size (maybe a little larger, although based on the sample size of one for each, I wouldn't make much of that), and mostly known from very similar fossils; most of the head and a handful of foot and hand bones.  Like many of the proceratosaurids, it had a bony head crest along the nose, although apparently not as dramatic as that of Guanlong.  Little is known of the region in which these fossils were found, so little can be said about its environment or contemporaries as of yet.  That said, its discovery in 2010 was the catalyst that led to the recognition of proceratosauridae as a valid family, and as an early radiation of the tyrant's lineage.

Guanlong wucaii. A little bit later, at 160 million years ago in the Oxfordian of Chinese Turkestan, we find Guanlong.  This guy is known from two relatively complete specimens; a fairly complete adult and an even more complete subadult.  Because of the completion of the specimens, they are useful in reconstructing what more incomplete forms were probably like (like Proceratosaurus and Kileskus.)  At about 10 feet long, much of it tail, Guanlong was about the same size as the former two as an adult, and it shows that tyrants still had fairly long, grasping three-fingered arms.  Its crest is quite tall and delicate, significantly larger in the adult than in the subadult.  In fact, quite a few changes occurred physically between the adult and subadult, including the spreading and growing of the crest, the younger specimen had a comparatively larger orbit, hand, and smaller pubic boot.  Like other rather undifferentiated coelurosaurs, it had a comparatively small head (compared to later tyrannosaurs, certainly), longer neck, slender build and jaws, and overall probably fairly speedy build.  It is from the Shishugou Formation, which is relatively well known with many sauropods, a stegosaur, a tiny, extremely basal ceratopsian (Yinlong) and larger carnosaurs and other primitive tetanurans—all in all, a very similar environment to the Morrison at least in terms of comparable fauna.  It might have competed directly with basal coelurosaur Zuolong which was almost exactly the same size and build, although the stratigraphy isn't completely clear to me; they might not have overlapped.  The environment was a marshy woodland not far from a small, occasionally active volcanic mountain range, and the deposits tend to be fluviatile and lacustrine with a fair bit of volcanic ash.  Because this type of environment is the best for preserving fossils, that isn't necessarily meant to mean too much, though—most dinosaur-bearing fossil beds were similar (with some notable exceptions) but that isn't meant to mean that dinosaurs didn't live in places were erosion was happening, merely that their remains are extremely unlikely to have been preserved.

Stokesosaurus clevelandi.  This Morrison Formation fossil is known from only pretty scanty remains; the holotype is a broken pelvis from the Tithonian of the Brushy Basin member at the Cleveland-Lloyd quarry in central Utah (150 million years ago.)  Another smaller ilium was recovered from South Dakota a few million years earlier, and some other material has been occasionally referred to the genre, but in general, all of that is dubious, and many of those have since been rescinded.  Little is therefore known of Stokesosaurus, but it appears to be, like the three above, a fairly small creature, about 10-13 feet long as an adult, maybe.  For contemporaries, Stokesosaurus has the entire rather famous panoply of Morrison dinosaurs: Allosaurus, Diplodocus, Apatosaurus, Stegosaurus, Ornitholestes, etc.  If you were a kid growing up loving dinosaurs in the 70s and 80s like I was, almost every creature referred to the Jurassic that you've heard of was a contemporary of Stokesosaurus.  The Morrison was a large semi-arid basin with seasonal monsoon-like weather patterns where rivers flowed from the newly rising Rocky Mountains to the epeiric Sundance Sea.  Fern savanas made up much of the environment, and riparian forest valleys, and seasonal flooding and volcanic ash were periodic threats; although aeolian dune-like conditions persisted in the southern end of the formation.

Juratyrant langhami.  Although discovered in 1984, this fella wasn't described until quite a bit later; 2013, specifically.  Although initially believed to be con-generic with Stokesosaurus, it was later determined that many of the features that unite them were in fact quite common among all kinds of Jurassic tyrannosauroids.  Cladistic analysis still shows the two of them as the closest to each other among the proceratosaur family, however, with Sinotyrannus (see below) the next closest relative.  Juratyrant is quite a bit more complete than Stokesosaurus, and combined with the rather large distance between them, it was decided that it was too speculative to assume that they were the same genus, so the name Juratyrant was erected.  It comes from the Kimmeridge Clay formation of Dorset (although not from the Kimmeridgian period of the Jurassic) Tithonian age, about 150 million years ago, like the Morrison, but a more coastline habitat.  A number of dinosaur remains are known from the same formation, but few of them are complete enough to be diagnostic; although indeterminate larger therapods (Megalosaurus?), sauropods (Cetiosaurus?), and ornithopods and the stegosaur Dacentrurus are known from the region.   I've seen size estimates for Juratyrant that range from 10-20 feet, so it would have been (so far) possibly the largest of our specimens; although built lightly and probably rather speedy, it would have weighed maybe as much as the American black bear.

Sinotyrannus kazuoensis. The largest and latest of the proceratosaurs, Sinotyrannus is from the Jiufotang Formation from the Aptian of the Early Cretaceous, 120 million years ago.  This is the formation that is famous for it's lacustrine birds and feathered dinosaurs, including two-winged tiny maniraptorans and what-not that were buried in volcanic ash following eruptions nearby—newly famous critters like Confuciusornis and Microraptor.  Although famous for small fossils, which are normally quite rare, the Jehol biota captured here does not reveal it's secrets with regards to larger fauna very easily.  Psittacosaurus is known from the area, small caudipterids, some scanty ankylosaur remains, etc. No doubt many other animals lived here, and the slightly younger Yixian Formation (which I'll have to refer to later) has a more balanced faunal view.  Sinotyrannus is the largest known predator from the entire formation and one of the largest from the entire 11 million year old Jehol biota spread.  It was initially reported rather sensationally as "as big as late Cretaceous tyrannosaurs like T. rex" but that's not really true.  Although known from reasonably good remains, it's not complete, of course, and estimates for its length range from 20 to over 30 feet, with a weight of up to close to two tons—although probably considerably less (like I said, the early estimates were pretty wild.  I'd think it'd be comparable in weight to a large grizzly most of the time, possibly more in exceptional individuals.)

Note: According to the 2016 Brusatte and Carr cladogram, Stokesosaurus and Juratyrant are 'roids, not proceratosaurs, while Yutyrannus on the other hand is a proceratosaur instead of a 'roid.  The "standard" cladogram prior to that had that reversed, so we'll talk about Yutyrannus next time.  I suspect that over time, the Brusatte and Carr cladogram might gain more steam, but like I said earlier, the one presented by Hone was more convenient for me in terms of breaking the groups up into more or less like-sized chunks.

Tyrannosaur Chronicles

I'm finishing up (probably today; I've only got about 35 pages to go) David Hone's The Tyrannosaur Chronicles, a very recently published (this year) popular science book about the tyrannosaur family.  It's quite good, and I certainly recommend it, but there are a few comments I have about it.  First; I find it curious that he presents the Loewen phylogeny as if it's an unusual alternate compared to the "standard" phylogeny, but on Wikipedia, the Loewen phylogeny of Tyrannosauroidea is the only one presented.  There's a lot of interesting "what-ifs" in the details of the phylogeny of the Tyrannosaurini clade in particular floating around.  A very recent phylogeny that I like is included here as an image; one thing that it also does is show in graphical form more or less where and when the various species lived, which is really important.  One recent theory (too recent to have appeared in the book, unless it's in the last few pages, which I doubt) is the notion that tyrannosaurines in particular underwent two distinct radiations; a North American (Laramidan) one, and an Asian one.  The North American radiation gave us classics like Daspletosaurus and as well as more recent finds like Teratophoneus, Lythronax and the odd Nanuqsaurus.  T. rex himself, while appearing in North America, of course, would be an interloper in this scenario; a member of the Asian clade that includes Tarbosaurus and Zhuchengtyrannus and who came to North America across the land bridge and basically out-competed the local tyrannosaurines and replacing them through the Maastrichtian.

One odd find in the cladogram attached, which is at odds with most that I've seen, is that it shows Bistahieversor as a sister-group to Tyrannosauridae rather than nestled comfortably within the tyrannosaurines.  Either way, however, it points to something different going on in the north vs the south of Laramida.  It may not require an Asian Invasion of T. rex itself (the invasion might have gone the other way, for instance—Zhuchengtyrannus and Tarbosaurus might be North American invaders of the same type as T. rex—perhaps descended from Daspletosaurus.

In any case, I encourage you to have a look at the various phylogenetic proposals and come to your own conclusions.  Even if you don't like the way the tyrannosaurines are positioned in the one I've attached here, you've got to admit that the graphical presentation of the geography and timeline are extremely useful.

Another thing that I was unaware of, because I hadn't really been paying close attention is the raising of the subfamily alioramines which would have played a similar ecological role, I suppose, in Asia as the albertosaurines did in North America; smaller, more slender predators which presumably had either a different hunting strategy or different prey targets than the tyrannosaurines.  Similar to leopards in the same region as lions or tigers, if you will, or coyotes or golden jackals in the same region as wolves.

However, the most disappointing thing that The Tyrannosaur Chronicles didn't include was a good catalog of the various members.  This was always one of my favorite parts of Greg Paul's old Predatory Dinosaurs of the World, and a brief catalog with time-frame, size, location and some description of both the animal itself and the environment in which it lived would be welcome.  I guess I'll have to do it myself; it shouldn't be too hard to start with Infogalactic, look at the primary sources from there to add, if necessary, and put my own little catalog together.

So... I guess that means I've got at least a few more PALEONTOLOGY tagged post coming up in the very near future...  Maybe while I'm at it, I'll even add the Megaraptorans just in case they end up belonging to the same family tree as well (as opposed to Neovenatoridae, as they are traditionally albeit hesitantly placed today.)

Tuesday, November 22, 2016

Ad Astra Iconics (not with stats... yet)

Crew of the Wayland Wyrd (or Double Dub for short)

  • Donovan Flint; (Dono, Donnie) — a knight, trained by his father as an independent.  His father was killed by Shadow Knight assassination team years ago.  He is the nominal owner of the Double Dub, a repurposed and heavily modified (to the point that it’s original model is almost completely unrecognizable) UX-83 corvette with an expanded cargo hold.  While not particularly angry, he does have a bit of a chip on his shoulder about the Hezun Empire, but mostly he resents any authority.  He’s somewhat friendly with the Monarchy (from whom he has a Letter of Marque, although he doesn’t use it too often), as long as he stays in the fringes where the Monarchy’s influence is relatively light.  His main goals are to keep themselves in enough money to keep flying and maintain their freedom and nomadic lifestyle, so he tends to make shipping runs (sometimes of questionable legality and political expediency) between what are essentially little backwaters.  Dono is fairly tall, with light brown hair and hard gray eyes.  He tends to have a laconic smirk on his face.  Think somewhere between Clint Eastwood’s Man With No Name and Han Solo, but a bit younger and played by someone that looks like Richard Spencer.
  • Surt — an alien, a former soldier of the Monarchy, and an electro-organic Tearexian hulk.  Surt is an incredibly large example of the species; at over 8 feet tall and 600 lbs.  He has bright red skin and dark hair; his eyes have a black sclera but an almost chrome-colored iris.  There is a organic electronics system that the Tearexian hulks have evolved and are born with.  Physically this means that he has a tiny series of markings, mostly on his torso and limbs that have the appearance of a circuit board schematic.  Usually they are very fine and difficult to see, but sometimes they flare up brightly.  This organic electronics system in the Tearexian hulks offers them some modest benefits to purely regular organic beings (like humans) but mostly it’s the very large size and strength of the hulks that makes them in demand.  Surt is a former janissary of the Monarchy, who would probably be considered a deserter by them if anyone cared to chase him down, although Dono makes sure to keep the crew in the backwaters and fringes where this isn’t much of a serious risk.  Surt is relatively gentle and friendly most of the time, but when provoked, becomes a seriously dangerous opponent.  He carries a massive chaingun sometimes, and fights with two high tech scramasaxes.
  • Jeiry Edwards — The pilot.  He grew up on a hot, dry world drag racing and hot-rodding speeders, and was press-ganged by a Revanchist merchant marine crew.  His talents as a pilot were quickly recognized, and he also showed the ability to learn ether magic, so he was trained as a full pilot.  In spite of this investment, he was always angry at the Revanchists for kidnapping him, so when Dono raided the ship he was on—at the time, Dono did not have ether drive capabilities (although his ship did have an ether drive) so he was only raiding locally, Jeiry eagerly jumped on the opportunity to ditch his erstwhile crew.  Jeiry is a bit nerdy.  He doesn’t socialize that much with the crew (and when he does, he’s wont to talk too much about things nobody else is very interested in) and he seems to get along better with the mechanical aspects of the Double Dub than with his crewmates, in many respects.
  • Had Kulan — an Idachar spy; he was a double agent who was informing handlers from both the Monarchy and the Revanchists on the activities of the Hezun Empire.  When his cover was blown on Omad Gan, he was nearly killed.  Luckily for him, Dono was running an operation of his own in the system and took Kulan with him, making him the third member of the crew to be on the run from someone at least.  Kulan is an alien human of the Idachar race, which is the majority race in the Hezun Empire.  This means that his skin is a chalky kind of gray color, as are his eyes and even his hair.  Kulan is actually an accomplished spy; not just in terms of procuring information as a double agent, as he did, but also he is trained as a cat burglar and assassin.
  • MP15-22 (Empie) — a bot.  He belongs to an older, slightly obsolete class of combat bot, but as with the ship itself, Empie has been fairly heavily “hot-rodded” into high performance.  Painted black and bright yellow, with a nearly featureless face-plate, Empie is surprisingly sarcastic in the way that he talks, and crows frequently about his emotionlessness—belied by his smugness, of course.

Tuesday, November 15, 2016

GMing advice

As a teaser from my FANTASY HACK m20 game, here's a portion of the text in Appendix I; basically it's meant to indicate a quick summary of my GMing advice of many years, distilled down to just a couple of pages worth of text.  This should be pretty basic stuff to anyone who's run for any length of time, but the audience that this is written for is primary newer GMs.
Successfully running a game of FANTASY HACK m20 (or any other role-playing game) is a challenging yet rewarding endeavor.  Besides; someone has to do it, otherwise there's no game for anyone!  If you accept the challenge of being the Gamemaster, there are a few things you should know.  This section has a very small bit of advice, based on my own experience, followed by a fair number of tools that can help you.  In no particular order: 
You're a player too.  Although your task, and therefore what makes the experience rewarding and fun for you is perhaps a bit different than for the other players, this is still a game and you should be enjoying yourself too.  GMing is not a chore; it's not a job, it's not what the guy who gets the short stick has to do.  If you are not motivated and having a good time, the game will suffer because of it for everyone.  If so, consider giving the reins to someone else. 
Be fair and be consistent.  One of the things that the players need most is feeling like they can make decisions for their characters based on reasonable risk assessment.  In other words, they need to feel like they understand the way the world works (and most likely they expect it work like the real world does.)  Although this is one of those "perception is reality" kind of things, especially on a highly rules-light game like m20, the players will rely solely on your judgement about how likely things are to be successful.  If they can't get a handle on that because your rulings and DCs are inconsistent, or if they are consistent but out of whack with their expectations, either one, it will create the strong impression that the game is arbitrary and therefore unfair, which will dramatically reduce the enjoyment that your players feel. 
Be varied and interesting.  There is a wealth of sources in terms of ideas for your adventures.  Don't ever feel bad about borrowing from any and every source you can imagine; books, TV shows, movies, video games, whatever.  Just don’t borrow the same kinds of elements from the same kinds of sources.  Even Gary Gygax wrote (although this is often forgotten) that the game was not supposed to have been mere dungeon-crawling, and some versions of the game stressed doing other things (not that this was often appropriated by the players.)  FANTASY HACK m20 is flexible enough to be used for all kinds of activities, and it actually is not designed specifically to be a dungeon-crawling game at all.  In addition, if you pay attention to your players, you will before long find it easy to judge when they are engaged and entertained, and when they are more bored or frustrated.  Pay attention to this and give them more of what they like and less of what they don't.  They may not all be on the same page about what their favorite aspect of the game is (and they may be in different moods to do different things at different times anyway) but some situational awareness is crucial for good GMing. 
Be generous and say yes.  Although I personally dislike games that are overly concerned with the acquisition of character wealth and powers, in general, players tend to be happier when they get what they came to the table for, rather than feeling like it's denied them.  This doesn't mean give them "stuff" necessarily; but it does mean allowing them to indulge what they want to do as a character.  FANTASY HACK m20 is meant to emulate swashbuckling action stories.  Think of a well-known example like the Star Wars franchise.  Do the characters ever get bogged down looking for equipment that they don't have access to?  While getting passage to Alderaan is a key plot element of the first movie, it's easily accomplished.  When Luke needs a lightsaber, he has one.  When the characters have the opportunity to have a speeder bike chase, they're readily available.  How does Luke even get his X-wing that he flies for most of the movies?  I dunno.  It's there when he needs it.  This is the kind of story that I intend to emulate.  Hoarding of gear, doing tedious accounting and shopping are not at the heart of this kind of story; they are things that are typically breezed over because they are tedious and boring.  Now; some players actually do enjoy that kind of thing, so I don't recommend excising it entirely.  But I do recommend a focus more on the action, role-playing and the solving of interesting problems than I do on making things arbitrarily difficult for the characters.  That's the spirit of swashbuckling adventure stories, after all. 
Let the PCs dictate the game.  Don't overplan, because you will tend to get locked into your plans the more time you spend on them.  This isn't your novel that the other players get to have a minor role in.  This is their game, and you're supposed to represent the environment and the setting.  Let them be the stars of the game, not anything that you’ve created.  Let them decide what kinds of characters and what kind of party to create; don't passive-aggressively punish them for not picking your ideal of a "balanced party" or whatever.

Don't give them simply one solution to problems and ensure by fiat that anything else fails.  If you are too prone to trying to not let the PCs have their head, as the saying goes, then maybe you should rethink being the GM.  If that’s the only way you can enjoy the game, then you are probably not equipped to be the GM.  Being a successful GM means always remembering that it’s their game.  You'll have plenty of interesting and fun things to do, and honestly, you'll probably be a great deal more entertained by seeing what they come up with then you will be trying to ram them into your own ideas of what they should do. 
That said, it's also my experience that few groups have enough initiative, especially at early stages of the game, to know what to do from scratch if you give them total freedom.  Usually they will wander around aimlessly and even with a great deal of frustration "trying to find the game."  Once they are able to anchor themselves a bit more into the setting and their characters, they are much more capable and willing to take the reins, start making things happen that they initiate, and pursuing character goals that they themselves have set, rather than plot goals that you have created for them.  So ease them into it, but when they're ready to take control, absolutely let them do so.
Be prepared with things to do if the players seem lost, bored, or just need some kind of motivation.  To quote Raymond Chandler, "when in doubt have a man come through a door with a gun in his hand."  An ambush by brigands, thieves, highwaymen, cut-throats, or dangerous wild animals is sometimes just the thing to get the game going again when it flags.  Have a list of names appropriate for your setting that you can draw from to give NPCs that you didn't anticipate the feel of being more than a hastily constructed expedient.  If the PCs ignore threats or certain events, in the back of your mind think about what is happening while they're not intervening.  Make their decisions (or lack thereof, as the case may be) have consequences that they can see in game.  Maybe they still won't care, (although you should take that as an indication that you're probably not presenting them with the kind of game that’s engaging or interesting to them if so.) 
 More likely, they'll take the perceived failures personally and be more motivated to keep them from happening again.  Nothing gets players more motivated than a rivalry with an NPC that has gotten the best of them at least once in the past.  In short, make your setting feel like a real world, not just an environment for them to interact with.  This is the big benefit of table-top RPGs over computer ones; you can have flexibility to do all kinds of things that a computer programmer could not anticipate, and you can react to PC actions that they wouldn't even be able to do in a computer game.  Do not make the mistake of sacrificing this advantage for your own convenience; your game will suffer from being too much like a computer game… but without the nice graphics. 
But again don't over-prepare. You don't need gigantic campaign settings the size of a continent.  You don't need a lot, actually.  A very brief outline of what you think is likely to happen over the next session or two, including a few details about some NPCs, monsters, and locations that the PCs are likely to encounter is usually sufficient.  I rarely type up more than a page of outline, and it usually ends up lasting for several evenings worth of play.  But in order to do this well, you simply have to practice.  Don't be afraid of not running the best game ever when you're starting.  You'll probably do better than you think, and even if you don’t, you’ll get valuable practice and experience and be better at it next time, if you pay enough attention to your group to notice what went well and what did not. 
The Secret Roll. As GM, you probably need a few details about your characters—a single line will suffice, but have the character and player names, their stat modifiers, AC, and skill modifiers and level noted at least.  There are always times when as GM you will want to make rolls for the character that the player is not aware of, or at least cannot see the result of, because a failure would give them knowledge that their character could not have.  A great example of this is where another NPC is trying to sneak up on the character, or when the characters are traveling and may get lost but not realize it while traveling through the wilderness.

Monday, November 14, 2016

Fantasy Hack m20 v1.1

I've made a few minor corrections and formatting fixes to FANTASY HACK m20.  It is now up to v. 1.1, as this is the first modification made since it's launch.  I don't anticipate major (or even minor) changes to follow, but I thought I'd announce the update just in case.

UPDATE: As is often the case, as soon as I make an update, I find more things that need doing; from the simple (correcting some spelling or grammar errors that I didn't see before) to the slightly more expedient (alphabetizing the spell list.)

So... as soon as I released v1.1, I needed to update it again to v1.2.

Wednesday, November 09, 2016


We interrupt our normal gaming and science fiction discussion to comment very briefly on current events.

Note: physiognomy works.  There's an exception or two here, but mostly you do not see any male that looks like a man, nor any female that looks like a woman.  Or at least not an attractive one...

And this... holy cow.