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| Size comparison of select small non-pygostylian theropods. From left: Palaeopteryx thompsoni, "Ornithomimus" minutus, Parvicursor remotus, Epidexipteryx hui. By Matt Martyniuk, licensed. |
Tuesday, June 5, 2012
The Smallest (Mesozoic) Theropods
The online community of paleontology fans seems to have an unnatural preoccupation with size. Visit any dino-related message board, and you will find near-daily debates over which carnivorous theropod was bigger (or more specifically, longer). Carcharodontosaurus? Giganotosaurus? Tyrannosaurus? Hint: if the length estimates are within a meter of each other, the weight estimates vary widely, and the beasts in question are known only from one or two fragmentary skeletons, the correct answer is "eh, they're all about the same size, give or take." Also, Therizinosaurus and Deinocheirus, being wimpy herbivores, definitely do not count.
Less often, you may come across a discussion about which dinosaur was the smallest. These usually are not the heated debates you find with the mega-carnivores, but rather casual examination of minutiae and bringing up oft-forgotten species. Yes, everybody knows Compsognathus. But did you know the "chicken-sized" type specimen is a subadult or juvenile, and the referred French specimen is much larger? And anyway, Parvicursor was daintier by far. At 34cm (13in) long, Anchiornis was hailed as the smallest dinosaur when it was initially described, but subsequent specimens were larger (up to half a meter in length; still smaller than Compsognathus and slightly shorter than the ~40cm Parvicursor). Epidexipteryx, at a stubby, tail-free 25cm (10in) in length, was the shortest of them all, but its lack of a long tail means that it was a bit larger than the competition in mass. And anyway, isn't Epidexipteryx a bird?
Sunday, April 1, 2012
What Is Enantiornis?
Enantiornis leali was among the first enantiornithes to be found, and the first to be recognized as a member of a unique lineage of "opposite birds" separate from modern birds (Gobipteryx minuta, now recognized as an advanced enantiornithe, was found earlier). But despite being such a widely recognized and historically important member of its namesake group, little can actually be said about this species in terms of ecology or life appearance. Of course, that can't stop us from trying to figure out as much as we can by examining the available evidence and ecological context of these long-dead birds.
Sunday, January 22, 2012
The Debate: Newt Gingrich vs. Jack Horner
This is something I had completely missed until my wife found it linked to on a political blog a few weeks ago. Filmed in 1998, it's a pretty awesome hour-long video of a debate held between paleontologist Jack Horner and then-Speaker of the House Newt Gingrich. The topic: "Were Tyrannosaurus rex active hunters or pure scavengers?" What else?
The debate is actually a follow-up to a previous forum Gingrich did with Horner, both as fundraisers for the Museum of the Rockies. Gingrich, it seems, is an avid armchair paleontologist.
It looks like C-SPAN doesn't let you embed videos, so here's a link to the full debate: http://www.c-spanvideo.org/program/111009-1
What's fascinating about this debate is how it illustrates almost point-by-point a lot of issues I've seen cropping up online lately about the nature of scientific hypotheses, and in particular Horner's approach to them. As some of you may know, Horner recently backpedaled on the whole tyrannosaurs-as-pure-scavengers hypothesis, saying that, from the start, it was merely an attempt to illustrate how the scientific process is supposed to work as opposed to how it often goes in paleo. (Horner explicitly renounced the pure scavenger theory in, among other places, an October 2009 interview on the outstanding Skeptics Guide to the Universe podcast).
Watching this debate unfold, two things surprised me more than I thought they would. First, wow, Gingrich really did his research for this! He comes fully prepared with several examples and analogues to modern ecosystems, many of the same arguments I've seen used in forum debates on this topic, including the fact that there are few if any pure scavengers among modern animals, that hyenas will often take live prey, that vultures can get away with it due to their ability to fly over enormous areas in search of carcasses, etc. Newt knows his stuff, and handily pummels Jack in the debate (though Horner appears to be acting as sort of a gracious host, lobbing him a lot of softballs and overall "letting" Gingrich win.)
The second thing, though, is in Horner's closing arguments. Gingrich easily beats Horner by throwing out analogy after analogy, employing simple logic to demonstrate why his hypothesis "T. rex were not pure scavengers" is superior. But Horner points out, in a way, that it doesn't matter. Debates are antithetical to science. Empirical science is in no way about who has the better argument. It's about who has a more rigorous, testable and ultimately falsifiable hypothesis and can support it with more data, not better analogies.
I have been a T. rex as predators booster since I was 6. But after watching this debate, it is clear to me that Horner has always had a better, more scientific hypothesis and overall approach to the science of paleontology. He is correct that the default assumption has always been that T. rex were active hunters. But at the end of the day, assumptions are not science, and it's a little bit appalling that this assumption has been made an implicit basis of so many statement published in peer-reviewed scientific papers without question. You can never, ever disprove the hypothesis that T. rex were active hunters without a time machine, because it's logically impossible to prove a negative. That's why in science, if we have a positive statement as our hypothesis, it's often necessary to take the null hypothesis (the opposite statement to the one we are testing) and attempt to disprove that in an attempt to support the actual hypothesis. Like string theory, T. rex-as-hunters is an idea that makes logical sense on paper but is unfalsifiable, and therefore not science--just educated speculation.
However, Horner's hypothesis can and has been disproved. We now have evidence of healed-over T. rex bite wounds that show that at least occasionally, they bit living prey species. Does this prove T. rex were active hunters? Not necessarily, but it's a major piece of data against Horner's scavenging hypothesis, and that is actually the strength of Horner's position--that it can be tested and shown to be wrong. In science, it's not always better to be right than it is to be rigorous.
So while he may have won the debate, Gingrich was right for the wrong reasons, while his opponent Horner was wrong for the right reasons.
The debate is actually a follow-up to a previous forum Gingrich did with Horner, both as fundraisers for the Museum of the Rockies. Gingrich, it seems, is an avid armchair paleontologist.
It looks like C-SPAN doesn't let you embed videos, so here's a link to the full debate: http://www.c-spanvideo.org/program/111009-1
What's fascinating about this debate is how it illustrates almost point-by-point a lot of issues I've seen cropping up online lately about the nature of scientific hypotheses, and in particular Horner's approach to them. As some of you may know, Horner recently backpedaled on the whole tyrannosaurs-as-pure-scavengers hypothesis, saying that, from the start, it was merely an attempt to illustrate how the scientific process is supposed to work as opposed to how it often goes in paleo. (Horner explicitly renounced the pure scavenger theory in, among other places, an October 2009 interview on the outstanding Skeptics Guide to the Universe podcast).
Watching this debate unfold, two things surprised me more than I thought they would. First, wow, Gingrich really did his research for this! He comes fully prepared with several examples and analogues to modern ecosystems, many of the same arguments I've seen used in forum debates on this topic, including the fact that there are few if any pure scavengers among modern animals, that hyenas will often take live prey, that vultures can get away with it due to their ability to fly over enormous areas in search of carcasses, etc. Newt knows his stuff, and handily pummels Jack in the debate (though Horner appears to be acting as sort of a gracious host, lobbing him a lot of softballs and overall "letting" Gingrich win.)
The second thing, though, is in Horner's closing arguments. Gingrich easily beats Horner by throwing out analogy after analogy, employing simple logic to demonstrate why his hypothesis "T. rex were not pure scavengers" is superior. But Horner points out, in a way, that it doesn't matter. Debates are antithetical to science. Empirical science is in no way about who has the better argument. It's about who has a more rigorous, testable and ultimately falsifiable hypothesis and can support it with more data, not better analogies.
I have been a T. rex as predators booster since I was 6. But after watching this debate, it is clear to me that Horner has always had a better, more scientific hypothesis and overall approach to the science of paleontology. He is correct that the default assumption has always been that T. rex were active hunters. But at the end of the day, assumptions are not science, and it's a little bit appalling that this assumption has been made an implicit basis of so many statement published in peer-reviewed scientific papers without question. You can never, ever disprove the hypothesis that T. rex were active hunters without a time machine, because it's logically impossible to prove a negative. That's why in science, if we have a positive statement as our hypothesis, it's often necessary to take the null hypothesis (the opposite statement to the one we are testing) and attempt to disprove that in an attempt to support the actual hypothesis. Like string theory, T. rex-as-hunters is an idea that makes logical sense on paper but is unfalsifiable, and therefore not science--just educated speculation.
However, Horner's hypothesis can and has been disproved. We now have evidence of healed-over T. rex bite wounds that show that at least occasionally, they bit living prey species. Does this prove T. rex were active hunters? Not necessarily, but it's a major piece of data against Horner's scavenging hypothesis, and that is actually the strength of Horner's position--that it can be tested and shown to be wrong. In science, it's not always better to be right than it is to be rigorous.
So while he may have won the debate, Gingrich was right for the wrong reasons, while his opponent Horner was wrong for the right reasons.
Wednesday, December 21, 2011
Bruhathkayosaurus is Dead. Again.
Above: Working sketches for a speculative B. matleyi reconstruction by Steve O'Connor. Click here for Steve's final drawing.
I don't know how common this knowledge is, but this is the first I've heard of it so humor me while I mourn the possibility of ever re-assessing the intriguingly large sauropod specimen known as Bruthathkayosaurus matleyi.
B. matleyi was known from fragmentary remains of the pelvis and limb bones found in the Tiruchirappalli district of Tamil Nadu, India. It was first described by Yadagiri and Ayyasami in 1989 as species of giant allosauroid. This classification was widely doubted online, but little follow-up work was ever done. The initial description is widely regarded as exceedingly poor in quality and not much can be discerned about the specimen due to poorly detailed drawings and insufficient text. Tom Holtz has even stated that "the hypothesis that this is no more than petrified wood has not been falsified yet to my satisfaction." However, Mickey Mortimer later noted that the tree trunk hypothesis "is questionable given the non-cylindrical bones preserved such as the ilium. Additionally, Chatterjee has personally examined the fossils, and while he has a bad record of misidentifying taxa, I give him enough credit to not confuse a tree for a limb bone."
Sankar Chatterjee did indeed apparently examine the material and told George Olshevsky and Tracy Ford that he believed it to be a titanosaur, as reported in 1999 here.
Holtz responded to these appeals by noting that "not all units are the Dinosaur Park or the Djadokhta. In some preservation is really, really, really crappy. You might get all sorts of authigenic growth on the fossils, or alteration of the original material. In outcrops like that, it isn't out of the question to be fooled into thinking bone is wood and vice versa, especially from simple surficial appearances. This is why a section of the fossil would help resolve if it is bone or wood." So, there's that. We'll now never be able to take that section.
While B. matleyi was a near-mythical celebrity among "semi-apocryphal gigapods", its legend loomed larger than (published) reality. While most online sources (such as the DML posts quoted above) had long since agreed that the specimen was probably a gigantic sauropod and not a gigantic carnosaur, no actual published reference to the species as a sauropod existed until five years ago (Krause et al. 2006).
And what a sauropod it was, maybe! Obviously with such a paltry footprint on the scientific literature, reliable size estimates for such a poorly described specimen are hard to come by. Luckily, some researchers have done the best they could with the available data and determined that, if B. matleyi was indeed a titanosaur with similar proportions to say, Argentinosaurus, it would have been very large indeed. Matt Wedel over at SV-POW has estimated the size of this animal in life at 139 tons. Mickey Mortimer has estimated its length at up to 34 meters. That would position it as one of the largest species of land animals ever, second only to Amphicoelias fragillimus, possibly.
And now, it appears that B. matleyi has suffered the same fate as its atlantosauroid rival for the record. In the comments at another SV-POW post about semi-apocryphal gigapods, Wedel reports that the type and only specimen of B. matleyi was at some point washed away in a flood.
UPDATE: Thanks to Fabrizio in the comments pointing out a source closer to the horse's mouth. Artist "palaeozoologist" at DeviantArt posted an apparent personal correspondance from Kumar Ayyasami last January, in which he reported that the specimen was lost in heavy rains several years ago. (There's some more discussion of the specimen and the author's publication record here, including the suggestion that Dr. Ayyasami may now be deceased--that is, if you can get past the inexplicable Ali G speak). So it sounds like not only was the specimen poorly described, but nobody had bothered to actually collect it from the field site in the ~15 years since its discovery, and it was (predictably and inevitably) lost to erosion.
Any hope of verifying the stupefying claims about this species' size now seem to be lost. And unlike A. fragilimus, which was described and well-illustrated by a mostly reputable source with no obvious errors, the poor state of the B. matleyi description will forever doom this creature to the realm of dubious claims. After all, given the poor state of the description, it seems possible that a simple scale bar error or other mix-up could have tainted the data, and therefore all of our size estimates.
So here's to Bruthathkayosaurus matleyi, a beast (or possibly, a tree?) that died 70 million years ago, raised its spectral head (or crown?) again for one tantalizing moment and then, like Hitchcock's Ornithichnites, sunk back beneath the earth before we could really learn anything about it.
I don't know how common this knowledge is, but this is the first I've heard of it so humor me while I mourn the possibility of ever re-assessing the intriguingly large sauropod specimen known as Bruthathkayosaurus matleyi.
B. matleyi was known from fragmentary remains of the pelvis and limb bones found in the Tiruchirappalli district of Tamil Nadu, India. It was first described by Yadagiri and Ayyasami in 1989 as species of giant allosauroid. This classification was widely doubted online, but little follow-up work was ever done. The initial description is widely regarded as exceedingly poor in quality and not much can be discerned about the specimen due to poorly detailed drawings and insufficient text. Tom Holtz has even stated that "the hypothesis that this is no more than petrified wood has not been falsified yet to my satisfaction." However, Mickey Mortimer later noted that the tree trunk hypothesis "is questionable given the non-cylindrical bones preserved such as the ilium. Additionally, Chatterjee has personally examined the fossils, and while he has a bad record of misidentifying taxa, I give him enough credit to not confuse a tree for a limb bone."
Sankar Chatterjee did indeed apparently examine the material and told George Olshevsky and Tracy Ford that he believed it to be a titanosaur, as reported in 1999 here.
Holtz responded to these appeals by noting that "not all units are the Dinosaur Park or the Djadokhta. In some preservation is really, really, really crappy. You might get all sorts of authigenic growth on the fossils, or alteration of the original material. In outcrops like that, it isn't out of the question to be fooled into thinking bone is wood and vice versa, especially from simple surficial appearances. This is why a section of the fossil would help resolve if it is bone or wood." So, there's that. We'll now never be able to take that section.
While B. matleyi was a near-mythical celebrity among "semi-apocryphal gigapods", its legend loomed larger than (published) reality. While most online sources (such as the DML posts quoted above) had long since agreed that the specimen was probably a gigantic sauropod and not a gigantic carnosaur, no actual published reference to the species as a sauropod existed until five years ago (Krause et al. 2006).
And what a sauropod it was, maybe! Obviously with such a paltry footprint on the scientific literature, reliable size estimates for such a poorly described specimen are hard to come by. Luckily, some researchers have done the best they could with the available data and determined that, if B. matleyi was indeed a titanosaur with similar proportions to say, Argentinosaurus, it would have been very large indeed. Matt Wedel over at SV-POW has estimated the size of this animal in life at 139 tons. Mickey Mortimer has estimated its length at up to 34 meters. That would position it as one of the largest species of land animals ever, second only to Amphicoelias fragillimus, possibly.
And now, it appears that B. matleyi has suffered the same fate as its atlantosauroid rival for the record. In the comments at another SV-POW post about semi-apocryphal gigapods, Wedel reports that the type and only specimen of B. matleyi was at some point washed away in a flood.
UPDATE: Thanks to Fabrizio in the comments pointing out a source closer to the horse's mouth. Artist "palaeozoologist" at DeviantArt posted an apparent personal correspondance from Kumar Ayyasami last January, in which he reported that the specimen was lost in heavy rains several years ago. (There's some more discussion of the specimen and the author's publication record here, including the suggestion that Dr. Ayyasami may now be deceased--that is, if you can get past the inexplicable Ali G speak). So it sounds like not only was the specimen poorly described, but nobody had bothered to actually collect it from the field site in the ~15 years since its discovery, and it was (predictably and inevitably) lost to erosion.
Any hope of verifying the stupefying claims about this species' size now seem to be lost. And unlike A. fragilimus, which was described and well-illustrated by a mostly reputable source with no obvious errors, the poor state of the B. matleyi description will forever doom this creature to the realm of dubious claims. After all, given the poor state of the description, it seems possible that a simple scale bar error or other mix-up could have tainted the data, and therefore all of our size estimates.
So here's to Bruthathkayosaurus matleyi, a beast (or possibly, a tree?) that died 70 million years ago, raised its spectral head (or crown?) again for one tantalizing moment and then, like Hitchcock's Ornithichnites, sunk back beneath the earth before we could really learn anything about it.
Monday, December 5, 2011
Waddle, _Achillobator_, Waddle!
Above: Revised illustration of Achillobator giganticus with corrected leg proportions. Scale bar = 500 mm. By Matt Martyniuk, all rights reserved.
"Fast for a biped?"
"Cheetah speed. Fifty, sixty miles an hour if they ever got out into the open, and they're astonishing jumpers."
This quote from the original Jurassic Park film did much to cement the image of dromaeosaurids, the raptor* dinosaurs, in the public consciousness as fleet-footed hyper predators. Despite being nearly 20 years old, this portrayal has by and large remained unchanged in popular culture, with raptors often stock monsters with near-supernatural murderous abilities in everything from tongue-in-cheek xkcd comics to (I hope) tongue-in-cheek made for SyFy movies.
*Yes, I'm going to commit a cardinal sin and refer to dromies as "raptors". "Raptor" in ornithology refers to most predatory birds, even those that hunt on the ground (the Raptor Research Foundation considers Secretarybirds to be raptors). Since dromaeosaurids were both predatory and birds under any sane definition of the word, there should be no problem referring to them as an extinct group of raptors.
But were raptors really particularly fast? Bipedal running speed in digitigrade animals (that is, those that walk on their toes like birds rather than their ankles like humans) is usually roughly determined by the ratio of the lower leg bones (the tibia/tibiotarsus) to the upper foot bones (the metatarsus). The longer the upper foot is in length compared to the lower leg, the faster an animal could run. Therefore, we would expect the fastest theropod dinosaurs to be those with the longest metatarsi relative to tibiae.
A prime example of a theropod specialized for running very very quickly are the parvicursorines. This specialized group of alvarezsaurids (strange theropodan insectivores with stout, powerful arms each bearing one very large claw) has among the longest lower leg to upper leg ratio of any Mesozoic dinosaur group. Looking at the statistics compiled by Mickey Mortimer at The Theropod Database (a phenomenal resource I turn to so often I really should just make it my browser's home page), the type specimen of Parvicursor remotus (see leg diagram here) has a femur 52.6 mm long, a tibiotarsus about 75.6 mm long, and a metatarsus 58 mm long. The functional lower leg is 113 mm long, well over twice the length of the upper leg. More importantly, the lower leg and upper foot bones were fairly close to being equal in length. This animal was clearly a speed demon.
How does this compare to raptors? If, as Jurassic Park claimed, raptors were exceptionally fast, we would expect them to have similarly long lower legs. The terrifyingly human-sized raptors in JP were a Hollywood invention, but we do know of raptor species slightly smaller and slightly larger than they were.
On the smaller side were the famous Deinonychus antirrhopus. According to TTD, the femora of one relatively complete specimen measured 248 mm, with a tibia 324 mm long, and a metatarsus 151 mm long. Again, the total lower leg length is nearly double the upper leg. But the upper foot bones were only half as long as the lower leg bones.
The "fast raptor" meme was started by John Ostrom himself, when he first described Deinonychus in 1969. This was merely speculation on his part, as the hind limb was not completely known in the first specimens. Ostrom actually changed his opinion in later papers, finding that the femur was shorter than he'd initially thought, and that the foot bones were surprisingly short compared to other dinosaurs. Not only was Deinonychus not particularly fast, it probably could not have been nearly as fast as most other small theropods, including modern flightless birds, let alone cheetahs.
Another very popular type of raptors are advanced giant dromaeosaurines (Utahraptor and Achillobator). I recently found myself revising an older drawing of an Achillobator giganticus, which were, as mentioned above, only slightly larger than the Jurassic Park raptors. Many young dinosaur fans are very attached to these species in part because they're much larger than most other raptors, and because they had a slightly anthropomorphized novel written about them by Bob Bakker shortly after JP was released (Raptor Red). As a result, these big birds have a cachet in the collective consciousness similar to the generally more famous Deinonychus and Velociraptor--that of super-fast, agile and intelligent predators.
However, when finishing up my revised drawing, I had to double check the proportions several times to make sure I wasn't screwing it up. To my amazement, the legs, particularly the lower legs and upper foot bones, looked almost laughably short. Again according to TTD, the femur of A. giganticus measures 505 mm long, the tibia 490 mm long, and the metatarsus a paltry 234 mm long--less than half the length of the tibia. Not only is the metatarsus much, much shorter than the tibia, the entire lower leg in only marginally longer than the femur! The first thing that struck me wen looking at my own reconstruction was that this looked like the dromaeosaurid equivalent of Majungasaurus, those abilisaurids with the ludicrously short legs (which, coincidentally like Achillobator, have been suggested to be made up of chimeric specimens). It is also reminiscint of another stout-legged dromaeosaurid species, Balaur bondoc. While Balaur have been suggested to be possibly herbivorous due to these strange proportions, partial jaws and some teeth of Achillobator confirm that they were carnivores. But with legs like those, it's hard to imagine these creatures behaved the way the public imagines raptors to have done, chasing down fast moving prey. Frankly, it's hard to imagine Achillobator doing much beyond waddling across Nemegtian lake shores hunting turtles in epic slow-motion chases.
For the record, no described specimen of Utahraptor preserves both a femur, a tibia, and a metatarsus, so it's impossible to say whether or not they had the same squat proportions (unless somebody has some more detailed information on the numerous undescribed specimens in the BYU collections). For now, it would be safe to assume that they, too, would have been a laughing stock if they were caught trying to run.
Ok, but why would predatory animals have such stubby legs? There is a lot of evidence that dromaeosaurids were specialized for hunting big game, often animals larger than themselves. Deinonychus are infamous for their association with large ornithopods Tenontosaurus tilletti, and while evidence suggests they mainly targeted juveniles (Forster 1984), these were still much larger in terms of weight than even adult Deinonychus. Velociraptor are known to have grappled with the larger Protoceratops, and even a rumored specimen of a Microraptor apparently preserves evidence that they tackled prey larger than themselves. The short legs, especially the short foot bones, seem to be linked with the function of the large sickle-claw, making lack of speed a trade-off for improved ability to grapple and kill large game. Dinosaurs like Tenontosaurus and Protoceratops probably weren't particularly fast-moving themselves, making this sacrifice in speed worthwhile for the chance to down a massive animal that could provide a whole lot of food.
In the end, while the raptor chase scenes in Jurassic Park remain some of the most exciting parts of the movie, they require a significant amount of disbelief to be suspended; after all, it would be more realistic but slightly less suspenseful if Laura Dern had been able to evade the raptors in the power bunker by simply breaking into a light jog.
Sunday, November 20, 2011
An Alternate History for "Archaeoraptor"
For those who didn't figure it out, yesterday's mystery bird teaser was a bit of a trick question. In fact, the bird, which I restored on a lark when working on other species, never existed at all. It was, in fact, a restoration of the famous fossil chimera, "Archaeoraptor liaoningensis". Two commentors did get it half right, guessing Yanornis, though the long tail would make that identification impossible (as euornithines, Yanornis had short tails with retractable fans of feathers). You'll notice I modified the image a bit since yesterday: Mickey Mortimer pointed out that Yanornis were not fully toothed, and while the premaxilla had teeth, there is a small edentulous anterior portion which may have supported small beaks.
Despite being the most famous fossil forgery to come out of China, the "Archaeoraptor" debacle really is a success story for the peer review process. You can read a summary of the history of the specimen here. The upshot is that several researchers were suspicious of the chimeric specimen to begin with, and though they rather dubiously chose to submit it for publication anyway, their paper was subsequently rejected by two major journals, Nature and Science, before unfortunately being reported on without review in National Geographic magazine.
In fact, it's not surprising that so many researchers were suspicious of the specimen's validity to begin with. At it combines the skull, body and wings of a Yanornis martini (a songlingornithid euornithine) and the tail of a Microraptor zhaoianus (a microraptorian dromaeosaurid), the "Archaeoraptor" specimen would have been a very odd anomaly had it been real, given our knowledge of bird evolution (even in 1999, when it was first revealed to the public). Note that many news reports have stated that the hind limbs of the composite specimen also come from a Microraptor or even a third, unidentified type of bird, Zhao et al. (2002) noted that they also come from a Yanornis martini. Anyway, I broke out Professor Farnsworth's What-If Machine to try to suss out how our current understanding of bird evolution would be different if the composite nature of the original "Archaeoraptor" specimen had never been discovered.
Intuitively, it seems like a bird with the long tail of a dromaeosaurid combined with the advanced wing configuration and well-developed breastbone of an euornithine doesn't make much sense and would have eventually caused a rather extraordinary rearrangement of the bird family tree. To test this idea, I employed the services of phylogeneticist extraordinaire Mickey Mortimer, who had coincidentally just finished coding Yanornis martini and entering it into a combined Theropod Working Group matrix. Mickey very helpfully humored my request and combined the relevant anatomy Yanornis and Microraptor into a single taxon, and plugged the result into a theropod phylogeny which included only other taxa known in 1999.
I'll let Mickey explain the analysis in more detail:
"Okay, so what I've done is taken my Theropod Working Group analysis and restricted it to taxa described by 1999 (when "Archaeoraptor" was supposed to be described) and coded by the TWG [Theropod Working group - Ed.] by 2005. I've also limited it to those characters taken from the first TWG analysis (Norell et al., 2001), since I already posted those results on my blog (http://theropoddatabase. blogspot.com/2011/06/theropod- working-group-matrix-recoded. html)
and Scott understandably doesn't want more of the Lori results being
distributed before publication [Click here for the scoop on Lori, Scott Hartman's unpublished Morrison troodontid -Ed.]. So data-wise, this is equivalent to
results I've already posted and with the exception of new specimens of
old taxa (like the new Caudipteryx specimens in Zhou et al., 2000),
which would be far too tedious to correct for, is representative of our
knowledge in 1999. "Archaeoraptor" is represented by the Yanornis
codings mixed with Microraptor tail codings. Many of those Yanornis
codings are from specimens besides the Archaeovolans holotype, but I'm
not going to take the time to go through and see exactly which
characters can be coded from only that specimen. In total, 11 codings
were changed between Yanornis and "Archaeoraptor". The entire matrix is
51 taxa and 210 characters."
The result of Mickey's analysis was that "Archaeoraptor" ended up as the sister taxon to Confuciusornis, but with some caveats. First, no other pygostylians (the group uniting Confuciusornis and modern birds)were included, because Confuciusornis is the only pre-1999 taxon coded by the TWG. A complete test of "Archaeoraptor"'s 1999 relationships would require the addition of taxa like Patagopteryx, Songlingornis (which is a potential close relative of Yanornis), Cathayornis, Iberomesornis, etc. Because the only plesiomorphic character of "Archaeoraptor" relative to Yanornis and other euornithines proper are the "longer tail, elongate distal caudal prezygapophyses and rectangular proximal caudal centra", "Archaeoraptor" looks, for 1999 standards, like what we might have expected the sister taxon of Pygostylia to be like. Nevertheless, because the bulk of the specimen is so derived, Mickey reckons the presence of the long tail might not pull it away from euornithines even if other basal pygostylians were included (the long tail would then have to be considered a bizarre reversal).
Mickey also tested a few alternate hypotheses to see how much support they would get. Perhaps the obvious conclusion for a scientists faced with an oddity like "Archaeoraptor" would be that it is probably a sister group to Pygostylia as mentioned above, or somehow intermediate between dromaeosaurids and pygostylians. Mickey tested this by forcing "Archaeoraptor", Confuciusornis, and dromaeosaurs to clade together, which resulted in only three extra steps: this would have been considered a pretty sound hypothesis given the small pygostylian sample. Interestingly, this arrangement would also have resulted in unenlagiines as basal eumaniraptorans and put Archaeopteryx in Troodontidae. Removing "Archaeoraptor" from the mix decreased the likelihood of a Dromaeosauridae+Pygostylia clade by five steps, which Mickey notes is still "plausible" but certainly less likely, and shows that "Archaeoraptor" would have created the kind of link between dromies and more advanced birds touted by the original NatGeo article.
This leads me to speculate that "Archaeoraptor" may have provided "evidence" for the modular evolution of pygostylians directly from some traditional dromaeosaurs. Modular evolution refers to cases where major traits of a descendant group appear in a taxon which simultaneously retains major "primitive" or plesiomorphic traits of the ancestors taxa. One famous fossil example of this phenomenon is Darwinopterus modularis, which is a long-tailed pterosaur with a generally primitive, "rhamphorhynchoid" body plan, but which has a characteristically pterodactyloid-type skull. Had "Archaeoraptor" been accepted as real, we may now have believed that characteristic euornithine traits evolved first in the skull, forelimbs and torso of taxa which possessed otherwise dromaeosaurid tails (this is almost the exact opposite of how we view bird evolution today, with tail shortening coming very soon after the split between dromaeosaurs and pygostylians). As Mickey found, without more discoveries of basal pygostylian and ornithothoracine birds, our cladograms may have rendered dromaeosauridae paraphyletic with respect to modern birds.
Ok, but how would all this stand up over the years after 1999, with more and better specimens of basal birds from a variety of lineages? From Mickey:
"Here's where I'd expect the inclusion of more pygostylians to have an effect though, since right now with only Confuciusornis, "Archaeoraptor" is effectively the most basal pygostylian. But if we had an omnivoropterygid, an enantiornithine, and other ornithuromorphs in there, Yanornis' birdy characters would nest it with those and make its dromaeosaurid-like tail a reversal."
So, the discovery of more and better basal bird specimens may have been enough for "Archaeoraptor" to be regarded more and more as a curious side-branch of the avian family tree, less and less relevent to bird evolution as a whole: a euornithine with some rather inexplicable reversals. This would be similar to the way the relevance of the Piltdown Man to human evolution was exponentially reduced by numerous valid specimens before it was finally found to be a hoax.
It wouldn't be DinoGoss without a discussion of taxonomical minutiae, and this is a question that has popped up on the net several times before: if "Archaeoraptor liaoningensis" included the holotype specimen of Microraptor zhaoianus, why isn't the former an objective senior synonym (that is, a name that is synonymous by virtue of being based on the exact same specimen, not a different specimen later assigned to an already named species) of the later?
The crux of the argument is that the original National Geographic article that released the "Archaeoraptor" name did not satisfy the criterion for publication set forth by the International Code of Zoological Nomenclature (ICZN).The article, "Feathers for T. rex", was written by Christopher Sloan for the November 1999 issue of the magazine. While Sloan did technically coin the name "Archaeoraptor liaoningensis" by publishing it in a widely-distributed magazine (and the ICZN does not mandate that names can only be created in peer-reviewed journals), he did not make it clear that he was intending to formally erect a new taxon (usually done by specifying ("new genus and species", "gen. et sp. nov.", or some variation). Furthermore, he explicitly referred to the fact that a formal description of the species was forthcoming. Most observers have interpreted this as falling short of the ICZN requirements for naming taxa, and I don't know of any good reason to disagree.
A more complicated factor is that, among the subsequent publications on the chimeric fossil, some authors did try to formally name the taxon and designate a lectotype (when a type specimen is found to actually represent two or more individuals, a lectotype must be chosen from among them to officially bear the name that originally applied to the lot). Noted BANDit Storrs Olson published an article in a 2000 issue of the Backbone newsletter of the US National Museum of Natural History in which he attempted to remove the tainted name from his own area of study, fossil birds. This is before many BANDits (the "birds are not dinosaurs" crowd) collectively reversed their positions and decided that dromaeosaurs ARE fossil birds after all, thus becoming MANIACs ("maniraptorans are not in actuality coelurosaurs"). Olson therefore designated the tail specimen as the lectotype. This would seemingly make "Archaeoraptor" the official senior synonym of Microraptor.
Not so fast. Olson did not actually describe the specimen or convey intent to coin the name. Like Sloan, he referred the creation of the name to other authors (in this case, Sloan himself, in the mistaken belief that Sloan's article DID coin the name). So Olson effectively specified that the nomen nudum "Archaoraptor" should refer to the tail, but failed to officially create the name, let alone specify its lectotype.
And that is why "Archaeoraptor liaoningensis" remains a nomen nudum; at least, just until some cheeky bastard decides to formally attach the name to something for reals. Technically, nomina nuda are still up for grabs nomenclature-wise...
Despite being the most famous fossil forgery to come out of China, the "Archaeoraptor" debacle really is a success story for the peer review process. You can read a summary of the history of the specimen here. The upshot is that several researchers were suspicious of the chimeric specimen to begin with, and though they rather dubiously chose to submit it for publication anyway, their paper was subsequently rejected by two major journals, Nature and Science, before unfortunately being reported on without review in National Geographic magazine.
In fact, it's not surprising that so many researchers were suspicious of the specimen's validity to begin with. At it combines the skull, body and wings of a Yanornis martini (a songlingornithid euornithine) and the tail of a Microraptor zhaoianus (a microraptorian dromaeosaurid), the "Archaeoraptor" specimen would have been a very odd anomaly had it been real, given our knowledge of bird evolution (even in 1999, when it was first revealed to the public). Note that many news reports have stated that the hind limbs of the composite specimen also come from a Microraptor or even a third, unidentified type of bird, Zhao et al. (2002) noted that they also come from a Yanornis martini. Anyway, I broke out Professor Farnsworth's What-If Machine to try to suss out how our current understanding of bird evolution would be different if the composite nature of the original "Archaeoraptor" specimen had never been discovered.
Intuitively, it seems like a bird with the long tail of a dromaeosaurid combined with the advanced wing configuration and well-developed breastbone of an euornithine doesn't make much sense and would have eventually caused a rather extraordinary rearrangement of the bird family tree. To test this idea, I employed the services of phylogeneticist extraordinaire Mickey Mortimer, who had coincidentally just finished coding Yanornis martini and entering it into a combined Theropod Working Group matrix. Mickey very helpfully humored my request and combined the relevant anatomy Yanornis and Microraptor into a single taxon, and plugged the result into a theropod phylogeny which included only other taxa known in 1999.
I'll let Mickey explain the analysis in more detail:
"Okay, so what I've done is taken my Theropod Working Group analysis and restricted it to taxa described by 1999 (when "Archaeoraptor" was supposed to be described) and coded by the TWG [Theropod Working group - Ed.] by 2005. I've also limited it to those characters taken from the first TWG analysis (Norell et al., 2001), since I already posted those results on my blog (http://theropoddatabase.
The result of Mickey's analysis was that "Archaeoraptor" ended up as the sister taxon to Confuciusornis, but with some caveats. First, no other pygostylians (the group uniting Confuciusornis and modern birds)were included, because Confuciusornis is the only pre-1999 taxon coded by the TWG. A complete test of "Archaeoraptor"'s 1999 relationships would require the addition of taxa like Patagopteryx, Songlingornis (which is a potential close relative of Yanornis), Cathayornis, Iberomesornis, etc. Because the only plesiomorphic character of "Archaeoraptor" relative to Yanornis and other euornithines proper are the "longer tail, elongate distal caudal prezygapophyses and rectangular proximal caudal centra", "Archaeoraptor" looks, for 1999 standards, like what we might have expected the sister taxon of Pygostylia to be like. Nevertheless, because the bulk of the specimen is so derived, Mickey reckons the presence of the long tail might not pull it away from euornithines even if other basal pygostylians were included (the long tail would then have to be considered a bizarre reversal).
Mickey also tested a few alternate hypotheses to see how much support they would get. Perhaps the obvious conclusion for a scientists faced with an oddity like "Archaeoraptor" would be that it is probably a sister group to Pygostylia as mentioned above, or somehow intermediate between dromaeosaurids and pygostylians. Mickey tested this by forcing "Archaeoraptor", Confuciusornis, and dromaeosaurs to clade together, which resulted in only three extra steps: this would have been considered a pretty sound hypothesis given the small pygostylian sample. Interestingly, this arrangement would also have resulted in unenlagiines as basal eumaniraptorans and put Archaeopteryx in Troodontidae. Removing "Archaeoraptor" from the mix decreased the likelihood of a Dromaeosauridae+Pygostylia clade by five steps, which Mickey notes is still "plausible" but certainly less likely, and shows that "Archaeoraptor" would have created the kind of link between dromies and more advanced birds touted by the original NatGeo article.
This leads me to speculate that "Archaeoraptor" may have provided "evidence" for the modular evolution of pygostylians directly from some traditional dromaeosaurs. Modular evolution refers to cases where major traits of a descendant group appear in a taxon which simultaneously retains major "primitive" or plesiomorphic traits of the ancestors taxa. One famous fossil example of this phenomenon is Darwinopterus modularis, which is a long-tailed pterosaur with a generally primitive, "rhamphorhynchoid" body plan, but which has a characteristically pterodactyloid-type skull. Had "Archaeoraptor" been accepted as real, we may now have believed that characteristic euornithine traits evolved first in the skull, forelimbs and torso of taxa which possessed otherwise dromaeosaurid tails (this is almost the exact opposite of how we view bird evolution today, with tail shortening coming very soon after the split between dromaeosaurs and pygostylians). As Mickey found, without more discoveries of basal pygostylian and ornithothoracine birds, our cladograms may have rendered dromaeosauridae paraphyletic with respect to modern birds.
Ok, but how would all this stand up over the years after 1999, with more and better specimens of basal birds from a variety of lineages? From Mickey:
"Here's where I'd expect the inclusion of more pygostylians to have an effect though, since right now with only Confuciusornis, "Archaeoraptor" is effectively the most basal pygostylian. But if we had an omnivoropterygid, an enantiornithine, and other ornithuromorphs in there, Yanornis' birdy characters would nest it with those and make its dromaeosaurid-like tail a reversal."
So, the discovery of more and better basal bird specimens may have been enough for "Archaeoraptor" to be regarded more and more as a curious side-branch of the avian family tree, less and less relevent to bird evolution as a whole: a euornithine with some rather inexplicable reversals. This would be similar to the way the relevance of the Piltdown Man to human evolution was exponentially reduced by numerous valid specimens before it was finally found to be a hoax.
It wouldn't be DinoGoss without a discussion of taxonomical minutiae, and this is a question that has popped up on the net several times before: if "Archaeoraptor liaoningensis" included the holotype specimen of Microraptor zhaoianus, why isn't the former an objective senior synonym (that is, a name that is synonymous by virtue of being based on the exact same specimen, not a different specimen later assigned to an already named species) of the later?
The crux of the argument is that the original National Geographic article that released the "Archaeoraptor" name did not satisfy the criterion for publication set forth by the International Code of Zoological Nomenclature (ICZN).The article, "Feathers for T. rex", was written by Christopher Sloan for the November 1999 issue of the magazine. While Sloan did technically coin the name "Archaeoraptor liaoningensis" by publishing it in a widely-distributed magazine (and the ICZN does not mandate that names can only be created in peer-reviewed journals), he did not make it clear that he was intending to formally erect a new taxon (usually done by specifying ("new genus and species", "gen. et sp. nov.", or some variation). Furthermore, he explicitly referred to the fact that a formal description of the species was forthcoming. Most observers have interpreted this as falling short of the ICZN requirements for naming taxa, and I don't know of any good reason to disagree.
A more complicated factor is that, among the subsequent publications on the chimeric fossil, some authors did try to formally name the taxon and designate a lectotype (when a type specimen is found to actually represent two or more individuals, a lectotype must be chosen from among them to officially bear the name that originally applied to the lot). Noted BANDit Storrs Olson published an article in a 2000 issue of the Backbone newsletter of the US National Museum of Natural History in which he attempted to remove the tainted name from his own area of study, fossil birds. This is before many BANDits (the "birds are not dinosaurs" crowd) collectively reversed their positions and decided that dromaeosaurs ARE fossil birds after all, thus becoming MANIACs ("maniraptorans are not in actuality coelurosaurs"). Olson therefore designated the tail specimen as the lectotype. This would seemingly make "Archaeoraptor" the official senior synonym of Microraptor.
Not so fast. Olson did not actually describe the specimen or convey intent to coin the name. Like Sloan, he referred the creation of the name to other authors (in this case, Sloan himself, in the mistaken belief that Sloan's article DID coin the name). So Olson effectively specified that the nomen nudum "Archaoraptor" should refer to the tail, but failed to officially create the name, let alone specify its lectotype.
And that is why "Archaeoraptor liaoningensis" remains a nomen nudum; at least, just until some cheeky bastard decides to formally attach the name to something for reals. Technically, nomina nuda are still up for grabs nomenclature-wise...
Saturday, November 19, 2011
Who's That Birdie?
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