Horns & Spikes Part 3: Jugal "Horns"

The ceratopsian jugal horn is unique in that it comes in two distinct forms: a non-capped form, where the jugal itself flares outward to a point without the assistance of an epijugal ossification; and a capped form, where the jugal flares outward to (usually) a point and is also capped by a separate epijugal ossification, sort of like the nasal horn. What's especially interesting is that the most impressive jugal horns--non-capped forms--are found in the most basal ceratopsians. The crowded genus Psiattacosaurus includes three species with particularly impressive jugal horns: P. sibiricus, P. major, and P. gobiensis.


In P. sibiricus, the jugal horn width (from tip to tip) is greater than the length of the skull. This Russian genus also has the distinction of being the horniest (snicker) psttacosaur. In addition to its impressive jugal horns, P. sibiricus also has small postorbital horns and amazingly large palpebral horns. I would not want to mess with P. sibiricus!


P. major is the largest species in the genus and features a large, triangular jugal horn which, unlike P. sibiricus but like most other ceratopsians, points lateroventrally. While not as wide as the skull is long, the jugal horn width is substantial, and a good swing of P. major's head could cause some serious damage.


Finally, P. gobiensis, the most recently-described species of Psittacosaurus, has large, laterally-pointing jugal horns that are textured in such a way as to suggest a large keratinous covering. Interestingly, P. gobiensis' jugal horns curve gently downward along their length, unlike the straight jugal horns of P. sibiricus and P. major. Among ceratopsians, these species have (so far) the most impressive jugal horns!


Epijugals didn't pop up until later in coronosaur evolution. Protoceratopsids (like Protoceratops, above) have large, flaring jugals with rounded ends and similarly rounded, blade-shaped epijugals. They give the skull a unique look in anterior view.


Ceratopsidae never developed the jugal "spikes" of psittacosaurs or the flaring jugal blades of protoceratopsids. Focusing instead on postoribtal and/or nasal horn development, most ceratopsids simply forgot about their jugal horns. In almost all specimens, the epijugal quickly fuses to the underlying jugal bone, and the two grow outward together. The separation is barely apparant in older animals. Just one chasmosaurine, Pentaceratops, has managed to somewhat revisit the glory days of jugal horns. Pentaceratops has a jugal horn length of about 144 mm. That may seem huge, but keep in mind that that's 14.4 cm, or about six inches--nothing to brag about when your postorbital horns are 56 cm long (about two feet).

Jugal horns are not phylogenetically relevant unless you're talking about the separation between capped and non-capped forms, which seems to be a consequence of higher taxonomy (coronosaur or non-coronosaur). Among ceratopsids, the jugal horns are almost always short and unimpressive, and all those illustrations you see of Triceratops with whopping jugal horns are almost certainly inaccurate, unless the epijugal was, itself, capped by a massive keratinous cover, which I find unlikely. The large jugal horns of protoceratopsids may have had some role in sexual display or species recognition (or sexual dimorphism), and the retention of epijugals was probably inhereted by ceratopsids, but that group never really did anything with them. They may be vestigal structures, overshadowed by the other horns and spikes of the head.

I Don't Have Enough Goddamn Time

This summer has been exceedingly busy. If I'm not running errands, trying to sell the condo, or doing yardwork, I'm eating or doing my meds or at my job. As a consequence, I've developed a significant gaming backlog, I've got a stack of unread technical papers this high, I've got potentially three art projects in the works that I've been unable to put much time into, and my blogging schedule has just gone to hell. If you've been following this humble weblog since its first inception, you'll remember that I used to have a very strict schedule: new posts would go up three times a week, M/W/F, the same as my zithromax schedule. If I felt like it, I'd also blog on the weekend.

See, that was back when I had spare time. Those days are long behind me.

And the helluvit is that I have a bunch of projects I'd like to start, much less complete, my Dinosaurs of Alaska (and other prehistoric beasties) children's book among them. The days seem packed now: get up early, do all my meds, go to work for eight hours, come home, eat dinner, do chores for what seems like eight more hours, watch the news, go to bed. Rinse, repeat. The trend is broken only on weekends, when we do more yardwork.* Every few days, I flip the heat lamp off and cool down the geckos so I can, hours later, pluck Liquid from his log and attempt to force medication into his swollan mouth (he's squirmy) and suck puss from his infected eyeball with an eyedropper. There have been times where I've found and had to remove what appears to be Gak from his eye with my itty-bitty tweezers. What's interesting is that Liquid is infinately more fidgity when I'm trying to give him medicine than when I'm threatening to gouge his eye out. Either he's very smart, very brave, or the medicine just tastes horrible (or all three).

Then there are the peripheral concerns: trying to get a raise during a pay freeze (good luck with that!), planning and coordinating my upcoming trip to the UK (SVP '09, baby!), the constant worry of not being able to sell the condo and having to find new renters, accruing an unhealthy gaming backlog (I forgot how to play a few of them), not having time to train for an upcoming Smash Bros. Brawl tournament, stressing out about affording upcoming games, keeping a close eye on the finances at all times, and finding time to walk two dogs. I think it's just the fact that it's summer in Alaska, and we've got like three months to fix things, do stuff, and sell property. That, and there's not enough goddamn time in the day.

I need to hire an intern who will work for free. Any takers?

*See, our yard was overtaken this summer by dandelions (Taraxacum), the mosquitoes of lawn care, and the speed at which they spread throughout our entire backyard is worthy of study. We've been told that dandelion-pickers only succeed in slowing their eventual takeover, as the root of a dandelion can extend 18 inches below the surface, and any part of the root leftover from a surface plucking will regrow a new flower in no time like some kind of perverted floral starfish. Anyway, the picker did nothing, and my wife understandably resorted to chemicals--the kind you buy at the grocery store's gardening department that promises to only kill "broad-leafed plants" like...dandelions. The horrible toxins proceed, however, to kill "all green plants," which includes grass. Half of our yard resembles a post-nuclear world.

According to the always-reliable Interwebs, the next step was to till the unfertile soil, the tortured earth, which we did. Our backyard now looks something like a post-apoloyptic hellscape that has been invaded by Graboids. Great clumps of soil, heavy with dead or dying roots from poisoned blades of grass, were leveled and thrown asunder. The ground successfully (???) tilled, the raking began. After which there was much seeding and laying of hay. The renovation of one's backyard is what you might call a process.

A Jurassic bird from China helps clarify theropod digital homologies

A new article printed in Nature on Thursday (Xu, et al. 2009) describes a Jurassic bird, pushing back the origin of birds almost 80 million years, Protoavis notwithstanding. More importantly, the new animal demonstrates that avian digital homologies have existed for just as long, and that, therefore, theropod dinosaurs cannot have been the ancestors of birds. The digits of theropod dinosaurs—especially maniraptorans, who are most often cited as likely avian ancestors—display a pattern of lateral digit reduction (LDR). In the most basal theropods, all five digits were present, though digit V was reduced to a bony splint. Later in theropod evolution, digit IV was lost as well. In all known tetanurine theropods, the remaining digits are, unquestionably, digits I-III (thumb, index, and middle fingers). This pattern of digit loss is at odds with most other animals, especially mammals, who tend to reduce their digits based on bilateral digit reduction (BDR). Were this the case in theropods, we would expect to see tetanurine theropods with digits II-IV (digits I and V would be lost).

Paleontologists have been at a loss to explain how theropods with digits I-III evolved into birds, which retain digits II-IV (based on embryology studies). The Hox genes are usually blamed for the switch, but genetic correlates do not fossilize well. Paleontologists have further attempted to discredit the studies which demonstrate that birds develop digits II-IV, insisting that the microscopic limb and digit buds in bird embryos cannot be accurately seen, and thus, perhaps birds really do develop digits I-III. This author thinks they doth protest too much, and that it is far more parsimonious to believe that, in fact, the ancestors of birds must have gone through a period of BDR, while theropods, who exhibit LDR, are not suitable ancestors for birds.

The new fossil is unfortunately named Limusaurus by the authors (who wear their bias on their sleeves), who consider it a ceratosaur. Ceratosaursia is a primitive group of theropod dinosaurs including such familiar forms as Ceratosaurus and Carnotaurus. These are big, toothy predators with robust features and powerful jaws. Limusaurus, by contrast, is a very small bird with long legs built for fast running; extremely reduced, splint-like arms; a toothless mouth covered in a rhamphotheca (beak); a single, fused sternal plate; and a gastric mill implying herbivory. These are all features indicative of an avian classification for Limusaurus. What’s more, the presence of a large singular sternal plate suggests that Limusaurus’ ancestors were flighted. Given its suite of basal and advanced features, Limusaurus may, in fact, be a basal paleognathid, obviously more derived than the tinamous, but could be an extremely basal ratite. For these reasons, I believe a name change is in order. To reflect the true phylogenetic nature of the animal, I recommend that Limusaurus be changed to Limuornis.

The authors correctly note that Limuornis retains a stub of digit I in the form of a single rounded metacarpal. This is not surprising in basal bird, and it is even less surprising to see that digit IV is also extremely reduced, though not the same extent, in a flightless animal. Digits II and III, though small, are fully-formed and possess small, blunt claws. The entire arm is very short, reflecting the condition seen in modern ratites. In some ways, the arms are vestigal, but they were probably used in courtship or aggression displays. The long legs and small, raised hallux suggest that Limuornis was strongly cursorial and did not fly or use its arms for prey capture. In fact, prey capture was not probable anyway—a large gastric mill of stones suggests that Limuornis was an herbivorous animal or, at the very most, omnivorous as in modern ratites. As stated above, Limuornis’ finger formula is II-III-IV, like modern birds but unlike theropod dinosaurs. More interestingly, the particular phalangeal formula of Limuornis’ fingers are incompatible with theropod hands. Whereas the tetanurine theropod hand formula is 1-2-3-x-x (metacarpals not included), the formula of Limuornis is x-3-3-x. Even if theropods did have fingers II-III-IV, their phalangeal formula is unlike Limuornis, so hand morphology is again incompatible. A reduction in manual phalanges is not unexpected given Limuornis’ flightless condition.

Despite its many derived features, Limuornis retains a few strikingly basal characters, including the presence of a long tail and a triradiate pelvis. Though incomplete, the tail is probably as short or shorter than in Archaeopteryx. While it may seem strange that a basal bird would retain its tail, keep in mind that a variety of basal flying birds, including Archaeopteryx and Jeholornis have long tails, and almost all flightless Mesozoic birds, including dromaeosaurs and troodontids, also have long tails—in some cases longer than Archaeopteryx. A long tail is obviously primitive for Aves, and it may have been lost multiple times within the group, including here, in paleognathes. The triradiate pelvis is a bit more difficult to explain, but, like the long tail, is probably primitive for Aves. Modern ratites have a pelvis that is not unlike Cretaceous flightless birds, though the pubis is more backswept. If a long tail was lost multiple times among birds, can we not say the same for the triradiate pelvis? This structure is potentially primitive for paleognathes as well, and the pubis retrovated to make room for a larger gut as herbivory became more important to the ratite diet. This would explain the differences between ratite pelvic girdles and those of flighted birds—the quadraradiate pelvis is merely a case of parallelism. Notably, the back half of the iliac blade in Limuornis is surprisingly similar to the same structure in emus and ostriches.

Limuornis is an exciting discovery which suggests that crown-group birds originated sometime in the Triassic period or, at the latest, the Early Jurassic. Incompatible finger formula notwithstanding, it is clear that the theropod dinosaurs that lived during the Late Triassic and Early Jurassic are horrible candidates for avian ancestry. They are either too large and ungainly (Dilophosaurus, Herrerasaurus) or too unspecialized (Eoraptor). While it remains possible that birds and dinosaurs share an arboreal common ancestor, I believe that Limuornis demonstrates that birds evolved prior to theropod dinosaurs, and that the two must have evolved from highly disparate groups of tetrapods.

MORE Exciting News

This book comes out November 1st. Mark your calendars! I've rarely been more excited about an upcoming book.

Two Awesome Dinosaur Stories

Two new papers, hot off the virtual presses.

Xing Xu et al, “A Jurassic ceratosaur from China helps clarify avian digital homologies,” Nature 459:940 doi:10.1038/nature08124

Look, ma: no teeth! I remember seeing the presentation for this beastie (Limusaurus) at SVP last year and was very excited about it. You could almost call it the ceratosaur version of an alvarezsaur. It's got long legs, short little arms, a lengthy neck and toothless jaws (and, one would assume, a beak). That's cool enough, right? Well, even better, Limusaurus helps solve the riddle of why non-avian theropods retain digits I-III, but modern birds seem to have digits II-IV. BAND members have pointed to this inconsistency time and time again to invalidate a theropod origin for birds (like any of their "ancestors" have digits II-IV). Limusaurus may solve the problem, as it retains digits I-IV, but digit I is reduced to a single stumpy phalange. The authors suggest that this represents a transition, and that what we see as digits I-III in tetanurine theropods is actually digits II-IV, and that those digits have taken on the characteristics of digits I-III. I don't know how convinced I am of this--Limusaurus is ridiculously derived AS IS. Either way, it's an awesome theropod regardless of what its finger formula implicates.

Paul C. Sereno, Zhao Xijin, and Tan Lin
A new psittacosaur from Inner Mongolia and the parrot-like structure and function of the psittacosaur skull
Proc. R. Soc. B published online before print June 17, 2009, doi:10.1098/rspb.2009.0691

Parrots crack nuts with their big curved beaks, and that's exactly what Sereno, Xijin & Lin are suggesting psittacosaurs did in this new paper. I'm not sure if they're describing another new species within the genus or simply a new well-preserved skull. National Geographic has a brief story about it, and if that's the skull they found, it's a nice-looking skull.

Thanks to Nick Gardner for sending me both papers. I'll have a tongue-in-cheek blog up about Limusaurus in the next few days.

Return of the Boneyard

Welcome back ot the Boneyard, ladies and germs, the blogosphere's premier paleontology blog carnival. It began long long ago on Brian Switek's original Laelaps blog and moved from blog to blog until its once-predictable occurrence was rare. Well today, dear readers, we have an extra-special GIGANTIC double-issue of the Boneyard for you. Let's get right to it--cast your eyes forth at the parade of links before you, and digest the sweet, succulent information contained therein!

The biggest news in the last few months has been the discovery of "Ida," a basal primate from the Messel beds of Germany. The initial media trainwreck irritated some and provided comic fodder for others. Eventually, Brian Switek organized a carnival for for the little gal. Thanks for making this Boneyard a little easier, Brian!

From primates, we move to sauropods--those giant dinosaurs whose heads how crane ever higher thanks to those SV-POW guys. They've actually got a bunch of (what amounts to) supplementary information on their humble blog. Here's what they have to say about heads, DinoMorph, necks in living animals, more against DinoMorph, the sadness that is T.rex's neck (drink!), the implications, and of course, horizontal semicircular canals. And just in case you're wondering what Darren Naish had to say about it all, well, here you go.

Darren also recently discussed the "Birds Come First" theory of avian evolution, and why it FAILS.

Traumador, that adorable tyrannosaur, has been busy lately. He's posted a watchable blog about microfossils as well as a handy field guide to himself, terrifyingly huge monitors, obscure pterosaur Cearadactylus, one of the stranger therapsids I've run across, and finally, giant ground sloths. Talk about diversity!

Speaking of therapsids and diversity, have you seen the new Art Evolved! show about Permian synapsids! There's a secondary "example gallery" lower down. Give it a shot, you'll learn something! Nima has a great breakdown of what a synapsid is over on his blog, too. Speaking of synapsids, Bora reports on a wonderful new mammoth fossil that's been in the news. And while this isn't necessarily paleontology, John McKay details the strange case of Teutobochus. Fascinating!

And of course there's plenty of art to go around. Angie has some really beautiful work up on her blog these days: Cryolophosaurus, a wonderful CG Maiasaura, and a three-part series on a CG Parasaurolophus. Wonderful work, Angie! Can't wait to see your next Art Evolved entry, if you're doing one...Mo Hassen describes three recent creations over at his blog, as well. Finally, Dave Bressan posted some awesome pictures up at his blog that you would all do well to check out.

Have to point these out: David Hone has written two wonderful posts about those whackjobs who comment on our paleo blogs telling use that pterosaurs are lizards or birds are crurotarsians or theropods are paraphyletic or whatever. This is required reading, people! Here's his first volley, and here's the fatal blow. Absolutely wonderful posts. David's blog is just plain awesome, and if you're not reading it currently, you should.

As for me, I written very little in the past few months, but I'd like to think that what I HAVE written has been high quality (cognitive dissonance, I'm sure). Check out my post on everyone's favorite fuzzy ornithischian, Tiangulong, Puijila, and the first two parts of my Horns & Spikes series: postorbital horns and nasal horns!

I'll keep adding posts through the weekend if you folks give me the links! Hope you enjoyed this exciting edition of the Boneyard!

You Say Goodbye...

And I say "heaven-o."

Hat-tip to PZ for alerting me to this story.

The Coming of the Boneyard

Remember, dear readers: The Boneyard, the interweb's premier paleo blog carnival, is coming once more to my humble blarg on June 12th. If you have submissions to submit, I suggest submitting them via either the comments or by email. I will, of course, be mining my blogroll and beyond for interesting posts (the Ida carnival, for instance) but I always like to see original content. Which reminds me, now that I'm back from E3, I really need to get going on my own Boneyard entry...

WPFR E3 '09 Awards

Now that E3's been over for two days and I've had some time to digest and reflect on what I've seen, I can finally hand out some coherent shows awards. Keep in mind that the awards given on this blog do not reflect the views of Nintendo World Report, which will probably be giving out its own Nintendo-centric awards (understandably) within the next week or so. So without further ado, let me present you with the When Pigs Fly Returns E3 2009 Awards!

Best Nintendo Wii Game: New Super Mario Bros. Wii

I played more of this game more than any other on the show floor. New Super Mario Bros. Wii is a sequel to the DS game from a few years ago, and in general it's the same game. Supposedly, the game features more than 80 levels, more than twice what the DS game had. A few new gameplay devices really set this one apart, though. First, Yoshi is in here and you can ride him. What's more, he's back to eating berries a la Super Mario World. New items like the Penguin Suit make up for the horrors that were the Micro Mushroom and Koopa Shell. Most impressively, New Super Mario Bros. Wii will feature four-player local co-op. Players compete for Star Coins, lives, and normal coins and are ranked at the end of each stage. You can pick up other players, jump off each other's heads, and do the Galaxy spin move by shaking the Wii Remote. It's undeniably fun, but like Four Swords Adventures, I can see it becoming frustrating if you're playing with people who don't know what they're doing. For the rest of us, though, New Super Mario Bros. Wii is gonna be freakin' sweet when it launches this holiday. Oh, did I mention that the Koopa Kids are making a triumphant return? That basically seals the deal.

Best Nintendo DS Game: Mario & Luigi: Bowser's Inside Story

The big news was The Legend of Zelda: Spirit Tracks, but I honestly didn't like it very much. Being a huuuge fan of Mario & Luigi: Superstar Saga on the GBA, I was excited to play Bowser's Inside Story, the third game in the series. The game looks absolutely stunning, with ridiculously detailed, expressive sprites. You get to play as Bowser and the Mario Bros. The boys have been shrunked down and inhaled by the Koopa King, and you've got to help or hinder his exploits by switching between Bowser and his enemies. The gameplay is roughly similar to that of Partners in Time, but controlling Bowser is a hoot. Looks like Fawful is back, which may be the best news ever. I don't have a release date for you, aside from before the end of the year.

Best Nintendo DSiWare Game: FlipNote Studio

FlipNote is going to be freeware on the DSi before too long, so that alone is awesome. The best part is that it's a simple, yet robust, piece of animation software for the Nintendo DSi. I created three simple animations: a T.rex yawning, a sauropod eating, and a blob bouncing. What's nice is that subsequent flipbook pages save a grayed-out version of the previous frame so you can work off that and not from scratch. You'll be able to send your animations to other DSi's (locally) and upload your creations to a special website. This little piece of software absolutely blew me away, and is reason enough to buy a DSi if you've been holding off.

Best Nintendo Non-Game: Wii Sports Resort

This game was at E3 last year, and seemed to be little more than a tech demo for Nintendo's upcoming Wii Motion Plus accessory. This year, however, the games are much more robust. The Wii Motion Plus is a very powerful tool, integrating a gyroscope and allowing players to have a lot more control over their in-game motions. I played Archery, Frisbee, and Table Tennis, and I was impressed with the control in each one. In Frisbee, it feels like you're throwing the thing, and this is coming from somebody who plays ultimate frisbee. Table Tennis is great because you can put a lot of spin on the ball, and the game can differentiate between a forehand and backhand shot. Archery challenges you to hit an ever-distancing target (you hold the Wii Remote as though it's the bow). I was surprised how much I enjoyed the games, and it's coming out in just a few weeks, bundled with the Wii Motion Plus.

Best Nintendo Announcement: Metroid: Other M

Some fans may decry the newest entry in the Metroid series as heresy given Team Ninja's involvement, but those people are idiots. You decried the Metroid Prime games too because of Retro Studio's involvement, only to eat your words when the games turned out to be awesome. Other M looks to be a more traditional 2D game (2.5D) but looks more action-oriented. These are the people who brought you Ninja Gaiden, after all. But that's not a bad thing. It's a prequel game that explores Samus' involvement in the military under Adam Malokavich from Metroid Fusion. I started tearing up when I realized that we were watching a Metroid trailer: you can check it out at NWR. Tycho and Gabe summed up my feelings pretty well.

Most WTF Announcement: The Wii Vitality Sensor

What the fuck? A pulse/O2 sensor for your Wii Remote? Nintendo President Iwata was really psyched about this bizarre device. His hope is that gaming can become a part of everyone's lives, even people who will never play games. So let's give 'em a heartbeat sensor for your index finger. I can see how the Vitality Sensor would work with a wide range of games, from Wii Fit to a stealth game, but Iwata didn't talk about any pratical uses for the device. It left most of us saying, predictably, "what the fuck?"

Most Disappointing Nintendo Announcement: No Mario Galaxy 2 in 2009

It's because they don't want it eating into the sales of New Super Mario Bros. Wii, which is a terrible reason. The games will appeal to different people, and as a bonus, Nintendo fans like me will buy them both. Miyamoto said that the game is basically finished but they're holding off until 2010. They'd better release that thing quick. Now I have to finally beat Mario Galaxy with Luigi (which, at times, is really hard).

Best 3rd Party Nintendo Game: Maramusa: The Demon Blade

It's like Odin Sphere without the alchemy. It's beautiful, and plays extremely well. What sets the game apart from other hack 'n' slash games is that your character accrues a number of swords during his or her journey, and when they break in combat, you must use your other swords. Merely unsheathing a sword does damage to opponents and can be worked into combat strategically, and you don't have to wait for a sword to break before switching. Swords "heal" over time, so you never outright lose a weapon. They all have different special attacks, though, so discovering them best combat options for a given situation is tantamount to success. Need I mention again that Maramusa looks stunning? High-res 2D sprites and incredible particle effects combine to create possibly the prettiest game at the show.

Now we'll do the Sony awards. You could basically call these the "non-Nintendo" awards because in many cases, these are multi-platform games. I'll be talking about them from the Sony perspective because I don't own an Xbox 360, and Halo: ODST has not compelled me to do so.

Best PS3 Game: Batman: Arkham Asylum

I want this game so bad. I was already excited before the show, but after having played it, this really is a must-buy. The game's three big names (Batman, the Joker, and Harley Quinn) are all voiced by their animated series counterparts, which kicks ass. Combat is fairly simple but will surprise you. Press Square plus a direction (toward an opponent) allows you to pull off a big branching combo which uses a dynamic location-based animation string. Batman will punch an approaching thug in the face, then kick a dude in the side, then elbow the guy behind him, all with the Square button! Counters can be executed with Triangle to keep the combo going. The booth attendant told me that during big fights, you can string up a 100+ combo! The Dark Knight can also toss batarangs to destroy environmental objects and briefly knock out bad guys. The meat of the gameplay involves Detective Vision, which shows where enemies in a room are, whether they're armed or not, and what environmental objects you can interact with. By using this ability, you can plan out attacks and get through big rooms without even being seen! It's so cool. There are few things more awesome than silently dropping from a gargoyle behind a thug, sneaking up behind him, and knocking him out before leaping back into the darkness. The game launches in October, and the special edition includes a full-size batarang! Whoa!

Best PSP Game: LittleBigPlanet

If you liked LittleBigPlanet on the PS3 (and how could you not?), you'll love the PSP sequel, which tosses new continents at you (I played in Australia) but retains the basic gameplay, physics engine, and aesthetic of the console version. It will also retain Stephen Fry as the narrator and a robust level creator. Unforunately, at this time, you can't share levels between the PS3 and PSP games, but Media Molecule is working on a fix for this. I'm very excited about this game, as I loved the first game. Worth mentioning in this category is Soul Calibur: Broken Destiny, which is identicle in every way (including graphically) to Soul Calibur 4, which is just mind-blowing. It plays a little slower, but the game is still in development. The game will feature a smaller roster, but that roster includes a few new characters including--get this--Kratos from God of War. While not especially original, Broken Destiny is amazing from a technical perspective and is definately worth checking out.

Best Game That Keeps Having Problems: Brutal Legend

Tim Schafer is a genius. He gave us Escape from Monkey Island, Grim Fandango, and most recently, Psychonauts. Now he's made a brilliant, hilarious, beautiful game called Brutal Legend that tells the story of a rock-tastic roadie named Eddie who is transported to a world where every rock/metal cover you've ever seen is part of an enormous overworld. The demo was incredible and completely sold me on the game. However, it's already had problems finding a publisher after Activision dropped it (probably because they couldn't annually whore it out) and now EA has picked it up, but Activision just sued because now they regret the decision. According to Tim Schafer, "If Activision liked it, they should've put a ring on it." Well said, sir. The game is supposed to launch in October, but Activision is trying to delay it with their lawsuit.

Best Game Not Demo'd: Modern Warfare 2

I don't particularly care for Activision, but they are publishing Modern Warfare 2, so I guess that's pretty nice. The sequel to my favorite FPS of all time comes out on October 11th, and I'm totally psyched. The Activision booth just had a repeating trailer for the game, and it looks incredible. It will follow the same kind of "summer movie" flow that the first one did, with a new storyline. Your character in CoD4 ("Soap" McTavish) is now your superior officer, and Al'Ahsad (who you killed in the first game) is now seen as a martyr in the Middle East, and tensions are yet again on the rise thanks to Al'Ahsad's second-in-command, Mackevoy. You'll travel to Rio, Siberia, and what appears to be Europe during Modern Warfare 2, and Infinity Ward aims to top the thrills and heart-pounding moments of Call of Duty 4 this November.

Biggest Third-Party Disappointment: No Jade at the Ubisoft Conference

I don't really give two craps about Raving Rabbids and Avatar. I was impressed by Assassin's Creed 2 (loved the first game), and TMNT: Smash Up was kind of disappointing. I was banking on Ubisoft showing off Beyond Good & Evil 2 after that leaked trailer on Kotaku a few weeks ago (I'm convinced it's BG&E2), but there was no such luck. Even worse, the Ubisoft booth had a poster for the original BG&E up, demonstrating that it was one of their key franchises, but nobody at the Ubisoft booth knew where Jade was this year. I can only hope that they announce something about the game by the end of the year. I was also surprised that Ubisoft didn't roll out a teaser trailer for a sequel to Prince of Persia.

Most Useless Hardware Upgrade: PSP Go!

This long-rumored PSP reboot is finally here, and it was playable on the show floor. The PSP Go is a slider in that you slide the screen up to reveal the face buttons, which are now ridiculously small and packed together. The screen itself is smaller and more square than the current PSP model, and the system does away completely with a UMD drive, which means its software is exclusively DLC. Supposedly, Sony is trying to figure out a way for players to use their existing UMD-based games on the new system, probably with an optional peripheral. I played LittleBigPlanet on the PSP Go and quickly came to dislike the small, closely-packed button layout. The analog nub is virtually in the center of the system shell, and the face buttons are small and shallow--they reminded me of the GBA SP buttons. The PSP Go features 16 gigs of flash memory, which is nice I guess, but I get by just fine with my 2 gig Memory Stick Duo. The Go's biggest stumbling block, I'm afraid, is its price point: $250. Sony does NOT know how to price their hardware.

Game of Show: Selasphorus rufus

Despite all those awesome games, the best part of the whole show was seeing my very first hummingbird, fluttering around the flowers outside the convention center. Based on the coloration (and a little online research), I tentatively identified the tiny bird as Selasphorus rufus, which ranges down the West Coast. What's most awesome is that it seemed totally unafraid of people, and I was able to get mere inches away from it with my DSi camera. As soon as I find myself an SD card reader for my computer, I'll post the pictures.

I've Been Away

You'll all probably noticed that it's been pretty quiet here at When Pigs Fly Returns lately. For those of you who aren't checking Facebook, I just finished covering E3 for Nintendo World Report here in sunny Los Angeles, California. I'll tell you all about it when I get back, but long story short: I met a lot of my colleagues (who are AWESOME), two of them got engaged (congrats to Neal and Lauren!), there was a rousing late-night round of PictoChat (hmmm), and an incident involving coleslaw. Justin, my brother-in-law, was kind enough to put me up for the last few days I'm here. I get back Monday, so stay tuned for more details!

Horns & Spikes Part 2: Nasal Horns!

The ceratopsid nasal horn is comprised of towering, fused nasal bones. In chasmosaurines, a small epinasal bone sits atop the nasals until it is incorporated into the bony structure of the horn. The epinasal is most obvious in certain juvenile and subadult Triceratops specimens. Unlike the postorbitals, nasal horns may not be primitive for Ceratopsidae—Zuniceratops lacks any kind of nasal horn, and the plasticity of the nasal structure in both ceratopsid subgroups suggests that they formed independently in both groups. We’ll talk about chasmosaurine nasal horns first, as they are the most standardized and vanilla.

Chasmosaurine nasal horns are never all that tall, and while they appear early in life, the horn itself does not become truly noticeable until late in life. Certainly the suggestion of a horn is there in subadult animals—in fact, it forms a noticeable “step up” from the rostrum to the forehead (see growth series below). However, aside from the small epinasal, a true horn does not develop until the chasmosaurine reaches maturity. In almost all genera, the horn is angled anteriodorally, forming a “ski slope” toward the rostrum. Once chasmosaurines reach maturity, the nasal horn continues to thicken and elongate—in old adult Triceratops specimens, the horn points subhorizontally. Some chasmosaurines, such as Torosaurus, place less emphasis on the nasal horn, seemingly content with developing a narrow, almost rounded horn. Agujaceratops follows a similar pattern. Even as its postorbital horns rise ever higher, the nasal horn remains short and rounded, though definitely more distinct than in Torosaurus.

Nedoceratops (= Diceratops, Diceratus) apparently has no nasal horn, but a close inspection of the type (and only) skull reveals that in fact it has a very small one. Lacking an epinasal, the stepped structure of the horn may suggest that the individual is simply a subadult who has not developed its nasal horn yet. On the other hand, it may also be related to Torosaurus (as a recent phylogeny suggests), which itself has an abbreviated nasal horn. Much has been made of Arrhinoceratops’ alleged lack of nasal horn (thus its name), but rest assured that a nasal horn is present, just partially broken off. In fact, its nasal horn may have been quite impressive.

Centrosaurine nasal horns are divided basically into two types: horns and bosses. A small banana-shaped boss is present on Albertaceratops, and two veeery small horns are present on an undescribed taxon from Utah (seems they could easily be sections of a broken boss). Horns are generally more prevalent than bosses among centrosaurines, though they come in some weird shapes. Centrosaurus (= Monoclonius) has a large, singular horn which curves every which way in different individuals. As you can imagine, early American fossil hunters gave different genus/species names based on different horn shapes. The result is that Centrosaurus has perhaps the most complex taxonomic history of all ceratopsids (except maybe Triceratops). Most skulls assigned to Monoclonius have been sunk into Centrosaurus, or have been unassignable to specific genera given what we now know of ceratopsid ontogeny. Styracosaurus albertensis was once restored with a towering nasal horn, but a reappraisal of the genus by Michael Ryan showed that this is wrong—Styracosaurus actually had a mid-sized nasal horn that may not have been all that pointy.

A sampling of ceratopsid nasal horns. From top to bottom: Triceratops prorsus (YPM 1922); Torosaurus latus (MOR 981); Pachyrhinosaurus lakustai (TMP 1989.55.1234); and "Monoclonius flexus" (AMNH 5239).

Midway between horn and boss is Einiosaurus, a mid-sized taxon from the Two Medicine Formation whose horn goes through quite a transformation as it ages. In young animals, the horn is short and slightly recurved. Once adulthood is reached, though, the horn quickly dominates the face as it grows up and then dips forward. It ends up looking something like a hook. The horn is laterally narrow, but its anterior-posterior breadth is impressive, and the horn narrows to a blunt point. Further toward the bossy end of things is Achelousaurus, from the same formation, but quite distinct (for an interesting interpretation of their differences, see Sampson 1995). Achelousaurus is a basal member of the Pachyrhinosaurini. As it reaches adulthood, all horny growths on the nasal and postorbital are reabsorbed, leaving large, rough-textured bosses in their place. The living bony tissue was probably fairly brittle toward the surface, and it would have been protected by a large keratin or horny covering. I like the suggestion that pachyrhinosaurine bosses were covered with oxen-like horn—the texturing is consistent.

The king of nasal bosses, however, is Pachyrhinosaurus, the largest of the centrosaurines and the most northerly: the taxon is known from the North Slope of Alaska. There are currently two recognized species: P. candensis and P. lakustai, both from Canada, but occur in different stratigraphic layers. Both show remarkable facial reconstruction with age. P. lakustai is the better-known animal, and a monograph was recently published on it. An ontogenetic series demonstrates the bizarre nature of the boss: babies and juveniles have respectable nasal horns and short postorbital bumps. As they grow, the nasal horns do not grow upwards, but laterally, eventually spilling over the lateral edges of the nasal bones (see illustration below)! In P. canadensis, the nasal and postorbital bosses eventually grow together to form a gigantic bony mass, but in P. lakustai, they all remain separate structures. In some individuals, the ballooning nasal boss begins to reabsorb and becomes concave.

Last time, I lamented how postorbital horns are not good phylogenetic markers, but nasal horns may be. All members of the Chasmosaurinae have basically the same nasal horn structure with different sizes and shapes depending largely on ontogenesis. The horn is almost always ski-sloped, fairly short, and sometimes rounded or blunt. The nasal horn often “steps up” the skull from the rostrum. The most impressive nasal horn among chasmosaurines may belong to Triceratops itself. Even its closest relatives (Torosaurus and Nedoceratops) have pitiful nasal horns by comparison. Even so, chasmosaurine nasal horns seem to have common features, and nasal horn morphology could be useful in establishing at least whether a ceratopsid is a chasmosaurine or not. As for centrosaurines, nasal horn morphology is strikingly different from chasmosaurines. The nasal horn is expressed just as often as a boss! When a horn is present, it grows vertically rather than angling forward, and it often dominates the face. In pachyrhinosaurs, a short nasal horn is present in juvenile animals but quickly becomes a boss in subadults. However, these bosses are very distinct from the small, smooth banana-shaped boss in Albertaceratops.

Horn growth in chasmosaurines, as exemplified by Triceratops. Note the "stepped-up" look of the rostrum in everyone except MOR 004.

How did ceratopsids use their nasal horns? In addition to figuring out that Triceratops used its postorbital horns for intraspecific combat, Andrew Farke also discovered that Centrosaurus must not have fought the same way, as the frills he studied carried virtually no scars attributable to intraspecific battle. This is not so surprising—without large postorbital horns, it’s difficult to see who centrosaurines head-to-head in quite the same manner, but it would appear that even the elongate nasal horn did not inflict injury on conspecifics. Absent any pathological evidence, it becomes very difficult to figure out what the nasal horns were used for. In chasmosaurines, they may have simply served as age indicators—Horner & Goodwin demonstrated that nasal horn morphology changes significantly along with postorbital morphology during growth in Triceratops. Like the angle and curve of the postorbitals, chasmosaurines may have looked to the development of the nasal horn to assess age and fitness amongst themselves. But what of centrosaurines, who exhibit a far greater range in nasal horn morphology?

Sexual dimorphism is often invoked. In fact, it’s been suggested that Centrosaurus apertus is simply the female of Styracosaurus albertensis—an idea that has largely been ignored. Sampson discussed the possibility that Einiosaurus and Achelousaurus were sexual dimorphs of the same species, but decided that without a large, mono-specific sample size (like a river crossing bonebed), there was just no way to be sure. Indeed, sample size is the biggest roadblock to determining sexual dimorphism among any extinct organism. Sexual dimorphism has been shown in Protoceratops andrewski only because its bones are littered across the Mongolian desert, and sample size is not a problem. When you only have a few partial skulls or skeletons, though, demonstrating sexual dimorphism becomes much more difficult if not impossible. Even where monospecific bonebeds are known, as in Pachyrhinosaurus lakustai, sexual dimorphism could be confused with ontogenesis. Is this individual a female, or just a subadult male? Or a subadult female? There is often no way to know for sure.

Horned centrosaurines may have used their horns as part of the overall display, and perhaps they lacked any kind of practical purpose aside from perhaps defense. Like the giant postorbitals of Triceratops, I can’t see Styracosaurus turning its horn away from a charging tyrannosaur. One wonders, however, about the bone strength of such horns. Could a predator be impaled without risk of injury to the horn itself? A recently-described Triceratops horn shows evidence of having been bitten off by a tyrannosaur! Pachyrhinosaurs, with its broad, horn-covered boss, would have been better protected against attack, as head-ramming would be a more viable alternative to impaling from both an injury prevention and practical standpoint. The bosses may also have been used in intraspecific combat—shoving matches or aggressive charges are well within the realm of possibility. It would be really cool if some pachyrhinosaur skeletons showed broken and/or healed ribs, which you’d think would be caused by intraspecific combat. Unfortunately, there are no postcranial details in the P. lakustai monograph, but hopefully somebody can investigate that in the future!

Overall, nasal horn morphology is more variable than postorbital horn morphology and may hold more value phylogenetically.